A Field Guide To Tropical Plant Families (Roland Keller)

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 (eBook)

https://doi.org/10.1007/978-3-031-05942-1
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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 mod-
ifications 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 Anthro-
pocene 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, ‘super-
foods’, 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’ jour-

nals, 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 the-
saurus 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 eco-
nomic plants with notes on the larger genera and, particularly helpfully, dis-
cussion 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 vege-

tative characters. When assessing species composition of a vegetation,
majority of plants is not flowering but still they are residents of the com-
munity, 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 Acknowledgements
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.
Contents
Part I User Manual and Architectural Concepts
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.1 Scope of the Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2 Why Consider Families? . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.3 Photos or Drawings? . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.4 Sources and Tests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.5 Structure of the Book . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 How to Use This Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
3 Architectural Concepts and Key to the Models . . . . . . . . . . . . 13
3.1 Concepts and Statements . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.2 Key to the Architectural Models . . . . . . . . . . . . . . . . . . . 18
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Part II Identification Keys
4 Key to the Groups of Families of Trees and Shrubs . . . . . . . . 29

5 Keys to Families of Trees and Shrubs . . . . . . . . . . . . . . . . . . . 37
6 Supplementary Keys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... 95
6.1 Key A—Tribes and Main Genera
of the Euphorbiaceae s.l. . . . . . . . . . . . . . . . . . . . . . .... 95
6.2 Key B—Tribes and Main Genera of the Malvaceae .... 103
6.3 Key C—Families of the Order
Sapindales + Picramniaceae . . . . . . . . . . . . . . . . . . .... 106
6.4 Key D—Tribes and Main Genera
of the Leguminosae . . . . . . . . . . . . . . . . . . . . . . . . . .... 111
6.5 Key E—Some Genera of the Rubiaceae . . . . . . . . . .... 119
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... 120
7 Keys to the Groups of Families of Lianas and Woody
Epiphytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
8 Keys to Families of Lianas and Woody Epiphytes . . . . . . . . . 123
xi
xii Contents
Part III Description of the Families
9 Acanthaceae to Asteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

9.1 Acanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
9.2 Achariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
9.3 Actinidiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
9.4 Anacardiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
9.5 Ancistrocladaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
9.6 Anisophylleaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
9.7 Annonaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
9.8 Apocynaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
9.9 Aquifoliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
9.10 Araliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
9.11 Araucariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
9.12 Arecaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
9.13 Aristolochiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
9.14 Asteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
10 Bignoniaceae to Buxaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
10.1 Bignoniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
10.2 Bixaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
10.3 Boraginaceae (incl. Cordiaceae, Ehretiaceae) . . . . . . . . . 178
10.4 Burseraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
10.5 Buxaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
11 Cactaceae to Cycadaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
11.1 Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
11.2 Canellaceae + Winteraceae . . . . . . . . . . . . . . . . . . . . . . . 186
11.3 Capparaceae (incl. Cleomaceae) . . . . . . . . . . . . . . . . . . . 187
11.4 Caricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
11.5 Casuarinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
11.6 Celastraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
11.7 Chloranthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
11.8 Chrysobalanaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
11.9 Clethraceae + Cyrillaceae . . . . . . . . . . . . . . . . . . . . . . . . 200
11.10 Clusiaceae + Calophyllaceae + Bonnetiaceae +
Hypericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
11.11 Combretaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
11.12 Connaraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
11.13 Convolvulacaeae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
11.14 Cornaceae + Nyssaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 215
11.15 Corynocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
11.16 Cunoniaceae + Brunelliaceae . . . . . . . . . . . . . . . . . . . . . 217
11.17 Cycadaceae + Zamiaceae . . . . . . . . . . . . . . . . . . . . . . . . 218
11.18 Cyclanthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
Contents xiii
12 Dichapetalaceae to Dipterocarpaceae . . . . . . . . . . . . . . . . . . . . 221

12.1 Dichapetalaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
12.2 Dilleniaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
12.3 Dioscoreaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
12.4 Dipterocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
13 Ebenaceae to Euphorbiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 229
13.1 Ebenaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
13.2 Elaeocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
13.3 Ericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
13.4 Erythropalaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
13.5 Erythroxylaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
13.6 Euphorbiaceae + Peraceae . . . . . . . . . . . . . . . . . . . . . . . . 241
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
14 Fagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
14.1 Fagaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
15 Gentianaceae to Goupiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
15.1 Gentianaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
15.2 Gnetaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
15.3 Goodeniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
15.4 Goupiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259
16 Hamamelidaceae to Humiriaceae . . . . . . . . . . . . . . . . . . . . . . . 261
16.1 Hamamelidaceae (incl. Altingiaceae) . . . . . . . . . . . . . . . . 261
16.2 Hernandiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
16.3 Huaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
16.4 Humiriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266
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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
19.1 Lamiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
19.2 Lauraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
19.3 Lecythidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
19.4 Leguminosae—Introduction . . . . . . . . . . . . . . . . . . . . . . 285
19.5 Leguminosae–Cercideae–Cassieae–Detarieae–
Caesalpinieae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
19.6 Leguminosae–Mimoseae–Acacieae–Ingeae . . . . . . . . . . . 291
19.7 Leguminosae–Papilionoideae . . . . . . . . . . . . . . . . . . . . . . 297
19.8 Linaceae–Hugonioideae + Ixonanthaceae . . . . . . . . . . . . 302
xiv Contents
19.9 Loganiaceae + Gelsemiaceae . . . . . . . . . . . . . . . . . . . . . . 303

19.10 Lythraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
20 Magnoliaceae to Myrtaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . 309
20.1 Magnoliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 309
20.2 Malpighiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
20.3 Malvaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
20.4 Marcgraviaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320
20.5 Melastomataceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321
20.6 Meliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
20.7 Menispermaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
20.8 Monimiaceae s.l. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
20.9 Montiniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
20.10 Moraceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
20.11 Moringaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335
20.12 Muntingiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
20.13 Myricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
20.14 Myristicaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
20.15 Myrtaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342
21 Nyctaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
21.1 Nyctaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
22 Ochnaceae to Oxalidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
22.1 Ochnaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
22.2 Olacaceae (incl. Schoepfiaceae) . . . . . . . . . . . . . . . . . . . . 351
22.3 Oleaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353
22.4 Opiliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355
22.5 Oxalidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 358
23 Pandaceae to Putranjivaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 359
23.1 Pandaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
23.2 Pandanaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 360
23.3 Passifloraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
23.4 Pentaphylacaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 363
23.5 Phyllanthaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365
23.6 Physenaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368
23.7 Phytolaccaceae (incl. Petiveriaceae) . . . . . . . . . . . . . . . . 369
23.8 Picrodendraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370
23.9 Pinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
23.10 Piperaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
23.11 Pittosporaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
23.12 Poaceae (Bambuseae) . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
23.13 Podocarpaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
23.14 Polygalaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
23.15 Polygonaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
Contents xv
23.16 Primulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380

23.17 Proteaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
23.18 Putranjivaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388
24 Rhabdodendraceae to Rutaceae . . . . . . . . . . . . . . . . . . . . . . . . 389
24.1 Rhabdodendraceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389
24.2 Rhamnaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390
24.3 Rhizophoraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392
24.4 Rosaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
24.5 Rubiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397
24.6 Rutaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407
25 Sabiaceae to Symplocaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
25.1 Sabiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
25.2 Salicaceae (incl. Lacistemataceae) . . . . . . . . . . . . . . . . . . 410
25.3 Santalaceae + Loranthaceae . . . . . . . . . . . . . . . . . . . . . . . 414
25.4 Sapindaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416
25.5 Sapotaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420
25.6 Schisandraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 424
25.7 Simaroubaceae + Picramniaceae . . . . . . . . . . . . . . . . . . . 424
25.8 Smilacaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 426
25.9 Solanaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
25.10 Styracaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430
25.11 Surianaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430
25.12 Symplocaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
26 Tetramelaceae to Torricelliaceae . . . . . . . . . . . . . . . . . . . . . . . 433
26.1 Tetramelaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433
26.2 Theaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
26.3 Thymelaeaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
26.4 Torricelliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
27 Ulmaceae s.l. to Urticaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439
27.1 Ulmaceae + Cannabaceae . . . . . . . . . . . . . . . . . . . . . . . . 439
27.2 Urticaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441
Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
28 Verbenaceae to Vochysiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . 445
28.1 Verbenaceae + Callicarpa . . . . . . . . . . . . . . . . . . . . . . . . 445
28.2 Violaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447
28.3 Vitaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 448
28.4 Vochysiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 453
29 Zygophyllaceae s.l. (incl. Balanitaceae) . . . . . . . . . . . . . . . . . . 455
29.1 Zygophyllaceae (incl. Balanitaceae) . . . . . . . . . . . . . . . . 455
xvi Contents
30 Herbaceous Versus Arborescent Habit in the Families

and Orders of Flowering Plants—Evolutionary Trends . .... 459
30.1 A Diagrammatic Representation of Angiosperm
Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 459
30.2 References to Phylogenetic Studies . . . . . . . . . . . . . . . . . 464
30.3 Discussion and Hypothesis . . . . . . . . . . . . . . . . . . . . . . . 465
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 469
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
Introduction
1
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 3

R. Keller, A Field Guide to Tropical Plant Families,
https://doi.org/10.1007/978-3-031-05942-1_1
4 1 Introduction
1.3 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 con-
cerning vegetative traits. When appropriate, the family description ends with a short anecdote. Photos
References 5
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 adap-
tations 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.
How to Use This Book
2
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 Euphorbiaceae-
Phyllanthaceae.
• 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 alter-
native 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).

https://doi.org/10.1007/978-3-031-05942-1_2
8 2 How to Use This Book
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.
2 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 Cale-
donia; 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.
10 2 How to Use This Book
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 Aquifoli-
aceae 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 inter-
esting 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. Mabberley, D.J. 2017. Mabberley’s Plant-Book, 4th ed, 1102 pp. Cambridge: Cambridge University Press.
2. Hallé, F., and R. Keller. 2019. Mais d’où viennent les plantes?, 180 pp. Actes Sud.
3. Hallé, F. 2004. Architectures de plantes, 162 pp. Montpellier: JPC Editions.
4. 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 11
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.
Architectural Concepts and Key
to the Models 3
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).

https://doi.org/10.1007/978-3-031-05942-1_3
14 3 Architectural Concepts and Key to the Models
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 indi-
vidual. Axillary buds (b) are lateral axes too, but very small and with prophylls reduced to scale-
leaves. 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

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 reorien-
tation 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, plagio-
down) 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 consid-
ered. 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.
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 trans-
formation 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 Gym-
nosperms 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 combi-
nations 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, RAUH-
MASSART and CHAMPAGNAT-OLDEMAN for which continua represent better the reality.
• Architectural models are more easily identifiable in young trees, before reiteration occurs. Reit-
eration 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 bran-
ches 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 syn-
chronisation 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 pla-
giotropic 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
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
Fig. 3.11 Architectural models: CORNER, HOLTTUM, CHAMBERLAIN, SCHOUTE, TOMLINSON, LEEUWEN-
BERG and KORIBA
Fig. 3.12 Architectural models: CHAMPAGNAT, OLDEMAN, MANGENOT, TROLL, PRÉVOST and NOZERAN
Fig. 3.13 Architectural models: MASSART, RAUH, SCARRONE, AUBRÉVILLE and FAGERLIND
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
Key to the Groups of Families
of Trees and Shrubs 4
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, POA-
CEAE - 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, DIPTEROCAR-
PACEAE, EUPHORBIACEAE, LEGUMINOSAE, MORACEAE, PHYLLANTHACEAE, SAPO-
TACEAE, URTICACEAE
5b Leaves not stipulate.
Group 3: ANACARDIACEAE, APOCYNACEAE, BURSERACEAE, CALOPHYLLACEAE, CAR-
ICACEAE, ERYTHROPALACEAE, MYRISTICACEAE, PHYLLANTHACEAE, SAPOTACEAE
4b Leaves opposite or whorled (Fig. 4.1d).
Group 4: APOCYNACEAE, CELASTRACEAE, CLUSIACEAE s.l., EUPHORBIACEAE, MAL-
PIGHIACEAE, 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).

https://doi.org/10.1007/978-3-031-05942-1_4
30 4 Key to the Groups of Families of Trees and Shrubs
Group 5: CHRYSOBALANACEAE, DIPTEROCARPACEAE, HAMAMELIDACEAE, IRVIN-

GIACEAE, 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). 11
11a Presence of leaves with glands.
Group 7: CHRYSOBALANACEAE, DICHAPETALACEAE, DIPTEROCARPACEAE, PASSI-
FLORACEAE, PHYLLANTHACEAE, ROSACEAE, SALICACEAE
11b Leaves without glands. 12
12a Leaf margin entire (Fig. 4.2c).
Group 8: CAPPARACEAE, CELASTRACEAE, CHRYSOBALANACEAE, DICHAPETALA-
CEAE, DIPTEROCARPACEAE, ERYTHROXYLACEAE, EUPHORBIACEAE, FAGACEAE,
HAMAMELIDACEAE, HUACEAE, LEGUMINOSAE, MALVACEAE, OCHNACEAE, PAN-
DACEAE, 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, LACISTEMATA-
CEAE, LECYTHIDACEAE, LEGUMINOSAE, OCHNACEAE, PUTRANJIVACEAE, RHAM-
NACEAE, 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, ROSA-
CEAE, 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, ERY-

THROPALACEAE, 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, CHRYSOBALANA-

CEAE, CLUSIACEAE, CORNACEAE, ERICACEAE, ERYTHROPALACEAE, HUMIRIACEAE,
ICACINACEAE, IXONANTHACEAE, LECYTHIDACEAE MARCGRAVIACEAE, OLACA-
CEAE, 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, POLY-
GONACEAE, URTICACEAE, VITACEAE
17b Scars not annular and petiolar bases different. 18
18a Inner bark with a network of fibres.
Group 15: ACHARIACEAE, BIXACEAE, DIPTEROCARPACEAE, ELAEOCARPACEAE,
LECYTHIDACEAE, MALVACEAE, ROSACEAE, URTICACEAE

19a Petiole distally swollen into a pulvinus (Fig. 4.3c).
Group 16: ACHARIACEAE, CAPPARACEAE, CELASTRACEAE, ELAEOCARPACEAE,
EUPHORBIACEAE, MEDUSANDRACEAE
19b Petiole different. 20
20a Presence of leaves with glands (Fig. 4.3d).
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, IXO-
NANTHACEAE, LECYTHIDACEAE, LINACEAE, OCHNACEAE, RHAMNACEAE, SALICA-
CEAE, VIOLACEAE
16b Leaves not stipulate. 21
21a Leaf venation palmate or tripliveined (Fig. 4.3e). 22
22a Petiole distally enlarged into a pulvinus.
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, CAPPAR-

ACEAE, CELASTRACEAE, COMBRETACEAE, CONNARACEAE, LAURACEAE, OLACA-
CEAE, 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, PAS-
SIFLORACEAE, 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, PRIMU-
LACEAE, RHABDODENDRACEAE, RUTACEAE, SAPINDACEAE, SCHISANDRACEAE,
VERBENACEAE, WINTERACEAE
24b Leaves without glands, translucent/dark dots or stellate hairs. 26
26a Branches plagiotropic by apposition (Fig. 4.4c).
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 fur-
rowed (Fig. 4.4d).
Group 25: BORAGINACEAE, ICACINACEAE, LECYTHIDACEAE, PROTEACEAE, SABIA-

CEAE, 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
29a Petiole distally swollen or pulvinate.
Group 27: ACHARIACEAE, CAPPARACEAE, ELAEOCARPACEAE, EUPHORBIACEAE,
LEPIDOBOTRYACEAE, OXALIDACEAE, PHYLLANTHACEAE
30a Veins II or III secant on lamina margin (Fig. 4.4f).
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, HAMAMELIDA-
CEAE, IXONANTHACEAE, LAURACEAE, LECYTHIDACEAE, NYSSACEAE, PENTAPHY-
LACACEAE, PITTOSPORACEAE, PRIMULACEAE, SIMAROUBACEAE, THEACEAE
31b Branches without series of shorter internodes or branches
with frequent ap. m. ab. (Fig. 4.5b), these denoted by discontin-
ued pith.
Group 30: BORAGINACEAE, CANELLACEAE, CAPPARACEAE, CELASTRACEAE,
COMBRETACEAE, ERICACEAE, HUMIRIACEAE, ICACINACEAE, IXONANTHACEAE,
LAURACEAE, LEGUMINOSAE (phyllodinous Acacia), MYRICACEAE, OPILIACEAE,
PENTAPHYLACACEAE, PHYSENACEAE, PHYTOLACCACEAE, SABIACEAE, SALICA-
CEAE, SANTALACEAE, SOLANACEAE, SYMPLOCACEAE, THEACEAE
6b Leaves opposite or whorled (Fig. 4.5c). 32
33a Leaf venation palmate or tripliveined.
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, ERYTHROX-
YLACEAE, 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, MAL-
PIGHIACEAE, PICRODENDRACEAE, RHAMNACEAE, RHIZOPHORACEAE, VIOLACEAE,
VOCHYSIACEAE
35b Leaf margin toothed or crenulate.
Group 34: CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, OCHNACEAE,
RHIZOPHORACEAE, VIOLACEAE
36a Nodes with interpetiolar ridge (Fig. 4.5f). 37
37a Three or five veins starting from leaf base or slightly above it.
Group 35: LAMIACEAE, LOGANIACEAE s.l., OLEACEAE, MELASTOMATACEAE, VER-
BENACEAE, 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, LOR-
ANTHACEAE, LYTHRACEAE, MELASTOMATACEAE, PINACEAE
39b Leaves and venation different.
Group 38: ACANTHACEAE, ASTERACEAE, CLUSIACEAE s.l., CRYPTERONIACEAE, GEN-
TIANACEAE, LAMIACEAE, LOGANIACEAE s.l., LYTHRACEAE, VERBENACEAE
36b Nodes without interpetiolar ridge. 40
40a Three or five veins starting from leaf base or slightly above it (Fig. 4.6c).
Group 39: BUXACEAE, GENTIANACEAE, LAMIACEAE, LAURACEAE, MELA-

STOMATACEAE, MYRTACEAE, SAPINDACEAE, SCROPHULARIACEAE
41a Leaves with small dots, peltate trichomes, stellate hairs or glands.
Group 40: ASTERACEAE, BIGNONIACEAE, COMBRETACEAE, EBENACEAE, MAL-
PIGHIACEAE, 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. 44
44a Petiole distally pulvinate (Fig. 4.6f).
Group 43: BIGNONIACEAE, LAMIACEAE
45a Leaf margin entire.
Group 44: ANACARDIACEAE, BUXACEAE, ICACINACEAE, LAMIACEAE, LAURACEAE,
MALPIGHIACEAE, MELASTOMATACEAE, MONIMIACEAE, MONTINIACEAE, MYRTA-
CEAE, NYCTAGINACEAE, OLEACEAE, PICRODENDRACEAE, PROTEACEAE, SANTA-
LACEAE, THYMELAEACEAE, VERBENACEAE
Fig.
4.6
45b Leaf margin toothed or crenulate.

Group 45: CELASTRACEAE, LAMIACEAE, MONIMIACEAE, OLEACEAE, PROTEACEAE
2b Leaves compound (Fig. 4.7a). 46
46a Leaf arrangement alternate. 47
48a Leaves pinnate (Fig. 4.7b).
Group 46: BURSERACEAE, LEGUMINOSAE, ROSACEAE, RUTACEAE, SAPINDACEAE,
SIMAROUBACEAE, STAPHYLEACEAE, VITACEAE
48b Leaves palmate or trifoliolate (Fig. 4.7c).
Group 47: ARALIACEAE, BIXACEAE, CAPPARACEAE, CARYOCARACEAE, EUPHORBIA-

CEAE, LEGUMINOSAE, MALVACEAE, PHYLLANTHACEAE, URTICACEAE, VITACEAE
49a Leaves pinnate or bifoliolate (Fig. 4.7d).
Group 48: ARALIACEAE, BERBERIDACEAE, CONNARACEAE, JUGLANDACEAE, LEGU-
MINOSAE, MORINGACEAE, OXALIDACEAE, PICRAMNIACEAE, PROTEACEAE, SABIA-
CEAE, SAPINDALES (MELIACEAE, RUTACEAE, etc.), ZYGOPHYLLACEAE
49b Leaves palmate or trifoliolate.
Group 49: ARALIACEAE, BIGNONIACEAE, CARICACEAE, CONNARACEAE, OXALIDA-
CEAE, PICRODENDRACEAE, RUTACEAE, SAPINDALES (SAPINDACEAE, etc.)
46b Leaf arrangement opposite or whorled. 50
50a Leaves stipulate.
Group 50: BRUNELLIACEAE, CARYOCARACEAE, CUNONIACEAE, LEGUMINOSAE, OCH-
NACEAE, STAPHYLEACEAE, ZYGOPHYLLACEAE
51a Leaves pinnate with more than three leaflets/pinnae (Fig. 4.7e).
Group 51: BIGNONIACEAE, LAMIACEAE, MELIACEAE, OLEACEAE, RUTACEAE,
SAPINDACEAE, VIBURNACEAE
51b Leaves palmate or trifoliolate.
Group 52: BIGNONIACEAE, LAMIACEAE, PICRODENDRACEAE, RUTACEAE, SAPINDA-
CEAE
Fig.
4.7
Keys to Families of Trees and Shrubs
5
Group 1—Foliar base sheathing the stem or its base decurrent (Fig. 5.1a); leaves simple or com-
pound; venation indistinct or parallel. (Gymnosperms or Monocots).
ARACEAE, ARAUCARIACEAE, ARECACEAE, ASPARAGACEAE, CUPRESSACEAE, CYCA-
DACEAE+ZAMIACEAE, CYCLANTHACEAE, PANDANACEAE, PINACEAE, POACEAE-
BAMBUSEAE, PODOCARPACEAE
1a Leaves simple. 2
2a Alternate leaf arrangement in trunk and branches. 3
3a Leaves needle-like, in groups of 2, 3 or 5. (Fig. 5.1b) 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)

https://doi.org/10.1007/978-3-031-05942-1_5
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. 11
11a Not in AF. (MASSART). Leaves flat, with parallels veins
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)
5 Keys to Families of Trees and Shrubs 39
14b AM. Foliar sheath not tubular. 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 polymor-
phous 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
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
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
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)
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
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. 5

5a Venation brochidodromous (Fig. 5.8a). MAGNOLIACEAE
e.g.: Magnolia (AM, AS)
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, MUNTINGIA-
CEAE, 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)
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.)
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
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, PASSI-
FLORACEAE, 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, DIP-
TEROCARPACEAE, ERYTHROXYLACEAE, EUPHORBIACEAE, FAGACEAE, HAMAMELI-
DACEAE, HUACEAE, LEGUMINOSAE, MALVACEAE, OCHNACEAE, PANDACEAE,
PHYLLANTHACEAE, PUTRANJIVACEAE, RHAMNACEAE, SALICACEAE, ULMACEAE s.l.,
VIOLACEAE
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
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
19a AF. Stem apices or buds with stellate hairs (Fig. 5.13f).
Venation densely reticulate. Bark or leaves with smell of garlic. 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)
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
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)
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 ?).
Caraipa (AM)
6c Shrub. Leaves not coriaceous. (Crushed leaves giving out a curious smell).
SOLANACEAE
e.g. Cestrum (AM)
leaf arrangement; leaves simple, not stipulate; leaves and stems without glands or glandular tri-
chomes; stem apices or young leaves pubescent (s.t. indumentum of minute scaly hairs).
ARISTOLOCHIACEAE, CONNARACEAE, EBENACEAE, ERICACEAE, ERY-
THROPALACEAE, ICACINACEAE s.l., LAURACEAE, MELIACEAE, MYRISTICACEAE,
OLACACEAE, PENTAPHYLACACEAE, PERACEAE, SANTALACEAE, SOLANACEAE,
STYRACACEAE, THEACEAE
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)
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)
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.
e.g. Leptaulus (AF, MA), Gomphandra (AS, OC), Gonocaryum (AS)
leaf arrangement; leaves simple, not stipulate; leaves and stems without glands or glandular tri-
chomes; stem apices and young leaves glabrous.
ANISOPHYLLEACEAE, CANELLACEAE, CELASTRACEAE, CHRYSOBALANACEAE,
CORNACEAE, ERICACEAE, ERYTHROPALACEAE, HUMIRIACEAE, ICACINACEAE, IXO-
NANTHACEAE, 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
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. PUTRANJIVACEAE

e.g. Putranjiva (AS)
6b Twigs lenticellate, brittle. (L. toothed). SALICACEAE
e.g. Scolopia (Paleo)
5b Leaf margin not hyaline. 7
7a Leaf venation pinnate Fig. 5.20a. (See 8a to 8c).
8a TROLL. Twigs lenticellate. (Mainly AM). CHRYSOBALANACEAE

e.g. Licania (AM)
8b Twigs scarcely or not at all lenticellate. Leaves coriaceous. (NOZERAN: Citronella).
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)
11b Periderm suberizing early (i.e. on the last or next to last GU). 13
13a AM. Lenticels forming longitudinal rows Fig. 5.20f. Leaves subsessile.
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
6a Leaves with glands. (Glands on lamina base Fig. 5.22a). 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, LECYTHIDA-
CEAE, 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). ROSACEAE
Eryobotrya (AS, s.t. invasive)
7b AS. Growth not strongly rhythmic. DIPTEROCARPACEAE
e.g. Vatica (AS)
7c Growth weakly rhythmic. (Shrub). URTICACEAE
Oreocnide (AS)
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, EUPHORBIA-
CEAE, 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)
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)
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. 2
2a Petiole with glands Fig. 5.24d. (Liana). 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, DIPTER-
OCARPACEAE, 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
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)
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.)
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)
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.
Fig.
5.27
ACHARIACEAE, ANISOPHYLLEACEAE, BORAGINACEAE, CARICACEAE, ERICACEAE,

EUPHORBIACEAE, HERNANDIACEAE, LAURACEAE, PIPERACEAE, TETRAMELACEAE
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 cor-
date 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
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 two-
ranked, 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). 2
2a Leaves with translucent dots (essential oils). RUTACEAE
Aeglopsis (AF), Atalantia (AS)
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)
3b Plant different. (Leaves small). 4

4a Venation clearly visible. (Branches plagiotropic by apposition). 5
5a Petiole not distally pulvinate. (L. with glands). 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
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, PASSI-
FLORACEAE, 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, VERBENA-
CEAE, 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)
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, COM-
BRETACEAE, GOODENIACEAE, LAURACEAE, MELIACEAE, RUTACEAE, SAPOTACEAE
1a Bark or leaves giving out an aromatic or spicy smell. 2

2a Petiole distally swollen (or bearing an articulation) (Fig. 5.33d). RUTACEAE
e.g. Esenbeckia (AM)
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)
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, PROTEA-

CEAE, 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)
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 supra-
tripliveined; 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)
Fig.
5.35
1b Bark and leaves not aromatic. 2

2a Small tree bearing deeply incised leaves. TORRICELLIACEAE
e.g. Aralidium (AS)
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, LEPIDOBO-
TRYACEAE, 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)
2a Leaf margin entire with midrib ending in a mucro or stellate hairs. (Stellate hairs: Ste-
riphoma, 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
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)
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, PENTAPHY-
LACACEAE, 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)
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)
7a Lamina cuneate at base and long red petiole. (RAUH). DAPHNIPHYLLACEAE
Daphniphyllum (AS, AU, cult.)
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 tri-
chomes). PENTAPHYLACACEAE
e.g. Ternstroemia (AM, AS)
10e Series of short internodes. Large leaves with long petioles. Numerous V II con-
nected 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)
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)
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)
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, LEGUMI-
NOSAE (phyllodized Australian Acacia), MYRICACEAE, OPILIACEAE, PENTAPHYLACACEAE,
PHYSENACEAE, SABIACEAE, SALICACEAE, SANTALACEAE, SOLANACEAE, SYMPLO-
CACEAE, THEACEAE
1a Leaf margin toothed or crenulate Fig. 5.40d. 2

2a MA. Large leaves with distinct hyaline margin and secant venation. 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. crenu-
late, 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)
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: War-
burgia). 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)
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
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.
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)
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. 2
2a Venation scalariform Fig. 5.43d. (ROUX). 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
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)
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),
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
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. Intern-
odes distinctly angular. LYTHRACEAE
Duabanga (AS)
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
simple, stipulate; leaf margin toothed; petiolar bases not connected by stipular scars; leaf venation
pinnate.
CELASTRACEAE, ELAEOCARPACEAE, EUPHORBIACEAE, OCHNACEAE, RHIZOPHOR-
ACEAE, 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)
2b Buds and young leaves without appressed hairs. 3

3a Leaves with glands. (PETIT Fig. 5.46c). EUPHORBIACEAE
e.g. Tetrorchidium (AF)
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, glab-
rous. (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
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
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, LOR-
ANTHACEAE, 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)
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.)
7a Branches monopodial. (Venation camptodromous). LYTHRACEAE
e.g. Woodfordia (Paleo)
7b Branches sympodial. (Venation camptodromous). CRYPTERONIACEAE
Crypteronia (AS)
8a External wood furrowed Fig. 5.49d. ASTERACEAE
e.g. Calea, Verbesina (AM)
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)
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
with interpetiolar ridge; leaves simple, not stipulate; venation tripliveined or supra-3–5-veined.
BUXACEAE, GNETACEAE, LAMIACEAE, LAURACEAE, MELASTOMATACEAE, MYRTA-
CEAE, 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)
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
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)
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)
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
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)
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).
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
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.
Mesua (AS)
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)
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;
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
ANACARDIACEAE, BUXACEAE, ICACINACEAE, LAMIACEAE, LAURACEAE, MAL-
PIGHIACEAE, 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 campto-
dromous). LAURACEAE
e.g. Beilschmiedia (AM), Caryodaphnopsis (AM, AS)
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)
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).
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
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, SIMAR-
OUBACEAE, 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)
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
Fig.
5.57
3a Spines in stipular position Fig. 5.58a. RUTACEAE

Harrisonia (AF-E, AS)
3b No spines or spines different. 4
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)
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
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
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 net-
work 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
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 3-
foliolate.
ARALIACEAE, BIGNONIACEAE, CARICACEAE, CONNARACEAE, OXALIDACEAE,
PICRODENDRACEAE, RUTACEAE, SAPINDALES (SAPINDACEAE)
1a Petiolar base sheathing Fig. 5.61e. ARALIACEAE
e.g. Cussonia (AF, MA, cult.), Schefflera
2a Leaves with translucent dots Fig. 5.61f. (Aromatic smell). RUTACEAE
e.g. Cusparia (AM), Vepris (AF, MA), Aegle (AS)
Fig.
5.61

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)
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. 2
2a Leaflets toothed with its V II ending in a tooth. 3
3a Large interpetiolar stipular scars. STAPHYLEACEAE
e.g. Turpinia (AM, AS)
3b Stipules different (thin). 4
4a AM. Leaves pubescent. 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)
1b Plant not aromatic. Leaves without translucent dots. 2

2a Young internodes quadrangular. Rachis winged (w) Fig. 5.63d. 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)
4c AF. Leaves palmately compound or trifoliolate Fig. 5.64e. Leaflets with an abscission
zone. PICRODENDRACEAE
Oldfieldia (AF)
Fig.
5.64
Supplementary Keys
6
6.1 Key A—Tribes and Main Genera of the Euphorbiaceae s.l.
PHY—PHYLLANTHACEAE, ACA—Acalyphoideae, CRO—Crotonoideae, EUP—Euphor-

bioideae. 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

https://doi.org/10.1007/978-3-031-05942-1_6
96 6 Supplementary Keys
9a Petiole distally pulvinate Fig. 6.1f. 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. (Phyllomor-
phic 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
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
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)
33b Glands at the base and the underside of the lamina (glands disposed
between the veins) Fig. 6.5a. ACA—Alchorneae
e.g. Alchornea
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)
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)
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 com-
pound, its underside with stellate or peltate hairs. CRO—Ricinodendreae
Ricinodendron (AF)
52b Characters different. (GUs with apical meristem abortion). 53
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).
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
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 round-
shaped, 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)
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 pla-
giotropic. (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
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.)

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 rhyth-
mically 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.
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)
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
Fig.
6.14
15a V. brochidodromous (Fig. 6.15a). Rhythmic growth noticeable due to short

internodes, or brittle monopodial branches, or resinous smell. Leaves imparip-
innate. (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, Schle-
ichera, 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.)
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)
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
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
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
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)
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)
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)
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
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)
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. 46

46a Branches plagiotropic (TROLL, Fig. 6.24a). 47
47a Some GUs without apical meristem abortion. 48
48a Leaves with glands. (Young internodes angular). 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)
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 zizag-
ging 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 <5mm wide. MIM—Mimoseae
e.g. Pseudopiptadenia (AM)
53c Leaflets stipellate. MIM—Ingeae
e.g. Albizia (Paleo)
43b Axillary buds solitary (Fig. 6.25b), or ramified system of buds (Fig. 6.25c). 54
54a Buds always solitary. 55
55a Rachis with glands (Fig. 6.25d), (glands can be vestigial in Erythrophleum) or
strong rhythmic growth associated with short internodes. (See 56a to 56d).
56a Leaflets small, opposite, many (>100). MIM—Parkieae
Pentaclethra (AF), Parkia
56b Leaflets large, opposite, not many (<30). Existence of GU in monopodial

series. Branches orthotropic. MIM—Mimoseae
Xylia (AF, MA, AS)
56c Leaflets alternate, internodes short: Serianthes, leaflets few, opposite and
quite large: Zygia. MIM—Ingeae
Zygia (AM), Serianthes (AS, OC)
56d Paleo. Rachis with minute or vestigial glands. Leaflets alternate, strong
rhythmic growth (Fig. 6.25e). Leaflets fewer than 100. (TROLL).
MIM—Dimorphandreae
Erythrophleum (e.g. AS)
55b Rachis without glands. Growth different. (Branches erect or decumbent).
(Short shoots present, rachis modified into a spine: Parkinsonia, spiny climber:
Mezoneuron).
CSL—Caesalpinieae
e.g. Delonix (AF, MA, AS, ornam.), Mezoneuron (Paleo)
Parkinsonia, Schizolobium (AM, ornam.), Caesalpinia
54b Buds disposed in a linear longitudinal series (Fig. 6.25f). 57
57a Leaflets very numerous, midrib dividing the leaflet in two very unequal parts.
Fig.
6.25
MIM—Parkieae
Pentaclethra (AM)
57b Leaflets numerous or few. Leaflet divided by its midrib in two subequal parts.
(See 58a to 58d).
58a Growth weakly rhythmic. (Pinnae opposite, spiny weakly prostrate: Cae-
salpinia (in AM), Mezoneuron. CSL—Caesalpinieae
e.g. Bussea (AF, MA), Gymnocladus (AS, North AM)
Mezoneuron (Paleo), Caesalpinia, Peltophorum
58b Strong rhythmic growth and occurrence of short internodes. (Pinnae or
leaflets alternate f. s. spp.). CSL—Dimorphandreae
e.g. Dimorphandra (AM), Pachyelasma (AF)
58c Leaflets few, large, opposite. MIM—Mimoseae
e.g. Xylia (AF, MA, AS)
58d Paleo Leaflets alternate, strong rhythmic growth: leaflets less than 100.
(Rachis with minute or vestigial glands). MIM—Dimorphandreae
Burkea (AF), Erythrophleum (e.g. AS)
6.5 Key E—Some Genera of the Rubiaceae 119
6.5 Key E—Some Genera of the Rubiaceae
1a Trunk with continuous and monopodial branching (ROUX Fig. 6.26a), PETIT (Fig. 6.26b),
ATTIMS, STONE (Fig. 6.26c). 2
2a Inflorescences lateral (ROUX, ATTIMS). 3
3a Stipules obtuse (ob), (Fig. 6.27a), triangular (tr), (Fig. 6.27b) or lanceolate. 4
4a Short shoots modified into thorns (Fig. 6.27c). Canthium (AF, MA), Catunaregam (Paleo).
4b No such shoots. 5
5a Stipules keeled. Timonius (MA, AS, OC)
5b Stipules not keeled. (ROUX, helicoidal anisoclady: Lasianthus).
Faramea (AM), Coffea (AF, MA, cult.), Lasianthus, Psydrax (Paleo), Ixora
5c Phyllomorphic twigs. Zuccarinia macrophylla (Malaysia)
3b Stipules notched (nd), bilobate or free (fr) (4 stipules for 2 opposite leaves). Randia (AM)
2b Inflorescences terminal (PETIT, STONE, CHAMPAGNAT). 6
6a Stipules obtuse (ob), triangular (tr) or lanceolate. Posoqueria (AM), Tarenna (Paleo).
6b Stipules annular (an), (Fig. 6.27d). Morinda (Paleo).
6c Stipules notched (nd) (Fig. 6.27e), bilobate or free (fr), (Fig. 6.27f), (4 stipules for 2 opposite
leaves). (Shrub or liana).
Mussaenda (Paleo, cult.)
1b Trunk with distinct rhythmic branching or sympodial trunk. 7
7a Trunk sympodial (inflorescences terminal, LEEUWENBERG (Fig. 6.26d), KORIBA). 8
8a Stipules obtuse (ob), triangular (tr) or lanceolate. (KORIBA, SCARRONE). Gardenia
8b Stipules notched (nd), bilobate or free (fr) (4 stipules for 2 opposite leaves). Psychotria
7b Trunk monopodial (inflorescences lateral, MASSART (Fig. 6.26e), RAUH, FAGERLIND
(Fig. 6.26f), SCARRONE). 9
9a Branches with lateral flowering (MASSART, AUBRÉVILLE, RAUH).
9b Stipules obtuse (ob), triangular (tr) or lanceolate.
Hoffmannia (AM), Guettarda (AM, AS, OC)
9c Stipules notched (nd), bilobate or free (fr) (4 stipules for 2 opposite leaves).
9d Mangrove.Scyphiphora hydrophyllacea (AS, AU, NC)
Fig. 6.26
Fig. 6.27
References
1. Webster, G.L. 1994. Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Annals of the Missouri
Botanical Garden 1: 33–144.
2. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3rd ed., 1022 pp. Cambridge: Cambridge University Press,
Cambridge.
Keys to the Groups of Families of
Lianas and Woody Epiphytes 7
1a Latex or coloured exudate (Fig. 7.1a)

Group 53: APOCYNACEAE, CELASTRACEAE, CLUSIACEAE, CONVOLVULACEAE,
EUPHORBIACEAE, GNETACEAE, LEGUMINOSAE, SAPINDACEAE
1b No latex or coloured exudate 2
2a Leaf arrangement alternate (Fig. 7.1b). Leaves simple (Fig. 7.1c) 3
3a Leaf venation pinnate (Fig. 7.2a) or parallel (Fig. 7.2a)
Group 54: ANCISTROCLADACEAE, ANNONACEAE, ARACEAE, CACTACEAE, CELAS-
TRACEAE, CONVOLVULACEAE, DICHAPETALACEAE, DILLENIACEAE, DIONCO-
PHYLLACEAE, ICACINACEAE, LEGUMINOSAE, LINACEAE, MARCGRAVIACEAE,
PANDANACEAE, PHYTOLACCACEAE, POLYGALACEAE, POLYGONACEAE, RHAMNA-
CEAE, SABIACEAE, SOLANACEAE, THYMELAEACEAE, VIOLACEAE
3b venation palmate (Fig. 7.2c), tripliveined (Fig. 7.2d) or campylodromous (Fig. 7.2e).
Group 55: ARISTOLOCHIACEAE, CUCURBITACEAE, DIOSCOREACEAE, EUPHORBIA-
CEAE, HERNANDIACEAE, LEGUMINOSAE (Bauhinia pp.), MALVACEAE, MENISPERMA-
CEAE, PASSIFLORACEAE, RHAMNACEAE, SMILACACEAE, VITACEAE
2b Leaf arrangement opposite (Fig. 7.1d) or leaves compound (Fig. 7.1e) 4
4a Leaves simple (Fig. 7.1c) (s.t. gum exudate in bark)
Group 56: ACANTHACEAE, ASTERACEAE, BIGNONIACEAE, CELASTRACEAE, COM-
BRETACEAE, DIOSCOREACEAE, GNETACEAE, ICACINACEAE, LAMIACEAE, LOGA-
NIACEAE, MALPIGHIACEAE, NYCTAGINACEAE, OLEACEAE, RHAMNACEAE,
RUBIACEAE, THYMELAEACEAE, TRIGONIACEAE, VERBENACEAE
4b Leaves compound (Fig. 7.1e)
Group 57: ARALIACEAE, ARECACEAE, BIGNONIACEAE, CONNARACEAE, CON-
VOLVULACEAE, CYCLANTHACEAE, DIOSCOREACEAE, HERNANDIACEAE (Illigera),
LEGUMINOSAE, MENISPERMACEAE, OLEACEAE, PASSIFLORACEAE, RANUNCULA-
CEAE, RUTACEAE, SAPINDACEAE, VITACEAE
https://doi.org/10.1007/978-3-031-05942-1_7
122 7 Keys to the Groups of Families of Lianas and Woody Epiphytes
Fig.
7.1
Fig.
7.2
Keys to Families of Lianas
and Woody Epiphytes 8
Group 53—Lianas with latex or coloured exudate
1a Some twining stems (Fig. 8.1a). 2

2a Opposite or whorled leaf arrangement (Fig. 8.1b). (See 3a to 3c):
3a Base of internodes swollen. Laticiferous threads in the leaves (Fig. 8.1c). (Latex yellowish).
GNETACEAE
e.g. Gnetum urens (AM)
3b Latex white or pale yellow. (Most of the ASCLEPIADACEAE are herbs with glandular
leaves, their woody species can produce a very corky rhytidome). APOCYNACEAE
e.g. Forsteronia (AM), Landolphia (AM, AF, MA), Parsonsia (AS, AU, OC)
3c Leaves without glands. (Red exudate in bark). CELASTRACEAE
e.g. Prionostemma (AM), Salacia (e.g.AF)
2b Alternate leaf arrangement. 4
4a Stipules present. 5
5a Latex white or coloured. (Stipules, lamina with basal glands, rhytidome or periderm
with thin longitudinal slits). EUPHORBIACEAE
e.g. Manniophyton (AF)
5b Leaves or leaflets with pinnate venation.
Petiolules with an abscission joint (Fig. 8.1d). Leaves trifoliolate or pinnate. Red
exudate in bark. (Lenticels becoming transversely elongated). LEGUMINOSAE
e.g. Machaerium (AM), Mucuna
4b Stipules absent. (See 6a to 6c). 6
6a Latex white, milky (Fig. 8.1e). Venation pinnate or palmate. Petiole not pulvinate.
Hairs appressed or hairs simple, stellate, erect or spherical. (Young stems soft, com-
pressible, woody or herbaceous liana). CONVOLVULACEAE
e.g. Ipomoea
6b Latex white. Leaves cordate and petiole pulvinate at both ends.
“ICACINACEAE”-CARDIOPTERIDACEAE
Cardiopteris (AS)
https://doi.org/10.1007/978-3-031-05942-1_8
124 8 Keys to Families of Lianas and Woody Epiphytes
6c Red exudate in bark (Fig. 8.1f). Modules of trunk distally twining. CONNARACEAE
e.g. Agelaea, Manotes (AF), Connarus
1b No twining stems. 7
7a Presence of tendrils. 8
Fig.
8.1
8a Margin of leaves or leaflets not toothed. Leaf margin entire, bilobate or bifoliolate.
Tendrils circinnate or bifid Fig. 8.2a. (Bark with red exudate). LEGUMINOSAE
e.g. Bauhinia
8b Leaflets toothed. Tendrils circinnate. SAPINDACEAE
e.g. Paullinia (AM)
7b Plant without tendrils. 9
9a AM. Leaves plicate (bifidous) Fig. 8.2b. Milky exudate. CYCLANTHACEAE
e.g. Evodianthus, Thoracocarpus (AM)
9b Leaves compound. Plant weakly prostrate? Young internodes angular or grooved. Latex
white, milky. SAPINDACEAE
e.g. Paullinia (AM)
Group 54—Lianas without latex or coloured exudate; alternate leaf arrangement; venation pinnate or
parallel
1a Plant with green stems and very reduced leaves Fig. 8.2c. 2
2a AM, AF, MA. Leaves reduced into hairs or spines forming areoles. Epiphyte with angular
or cylindrical green stems. CACTACEAE
e.g. Epiphyllum, Rhipsalis (mainly AM, ornam.)
2b Paleo. Twining plant. Leaves reduced to scales subtending leaf-like linear cladodes.
ASPARAGACEAE
Asparagus (e.g. AU)
1b Plants with well-developed leaves. 3
3a Bark with a network of fibres (Fig. 8.2d). Bark aromatic (with pleasant smell). Stipules
absent.
Twining plant or climbing by means of hooks (Artabotrys) Fig. 8.2e or with prehensile
twigs, Rauwenhoffia, Uvaria). ANNONACEAE
e.g. Artabotrys (AF, AS), Melodorum (AS, AU), Uvaria (Paleo)
3b Bark different or leaves stipulate. 4
4a Some twining stems. 5
8 Keys to Families of Lianas and Woody Epiphytes 125
5a Stipules absent, s.t. stipule-like petiolar scars. Rhytidome scaly. Concentric rings of
phloem and conspicuous rays (Fig. 8.2f). Petiole canaliculate.
Scaly hairs: Tetracera, tick hairs: Davilla, Doliocarpus (petioles and stems scabrous),
thin hairs: Pinzona. DILLENIACEAE
e.g. Hibbertia (mainly AU, OC)
Tetracera, Davilla, Doliocarpus, Pinzona (AM).
Fig.
8.2
5b Plant different. (Stipules present or absent). 6

6a Ochrea present, its annular scar readily observable (Fig. 8.3a). (Short twigs, young
leaf revolute, xylem lobate). POLYGONACEAE
e.g. Coccoloba (AM)
6b Stems without ochrea or annular petiolar scar. 7
7a Veinlets III and IV easily observed (Fig. 8.3b) or indumentum very hairy. 8
8a Stipules present. 9
9a Leaves toothed. 10
10a AS, MA. Teeth small or minute. Leaves without glands. (Laticiferous
threads in bark). CELASTRACEAE
e.g. Celastrus (MA, AS)
10b AM. Teeth small or minute. L. without glands. (Leaf underside shiny).
VIOLACEAE
Anchietea, Corynostylis (AM)
9b Leaf margin entire.
Apices with oblique or appressed hairs (Fig. 8.3c). (Twining feature
sometimes not evident to observe). (Underside of lamina with glands).
DICHAPETALACEAE
Dichapetalum (e.g. AM, AF)
8b Absence of stipules. 11
11a Leaves or bark aromatic.
AS. Vertical serial buds (Fig. 8.3d). (Node with one large appressed
prophyll, extremities of stems with young leaves in a zigzag line).
SCHISANDRACEAE
e.g. Schisandra, Kadsura (AS)
11b Leaves and bark not aromatic. 12
12a AF, MA. Base of lamina with glands (Fig. 8.3e).
MALPIGHIACEAE
Acridocarpus (AF, MA)
12b Leaves without glands. See (13a to 13c). 13

13a Leaves not coriaceous. Hairs appressed, stellate or spherical (hand
lens!). (Young stems compressible, intrusion of phloem in the wood,
Fig. 8.3f). CONVOLVULACEAE
e.g. Dicranostyles, Maripa (AM), Neuropeltis (AF, AS)
13b Leaves coriaceous. (Petiole faintly swollen distally, Venation
embossed). ICACINACEAE
e.g. Desmostachys (AF, MA), Neostachyanthus (AF)
Pyrenacantha (AF), Sarcostigma (AS)
13c AS. Leaves glabrous, smooth. (Petiolar base geniculate).
SABIACEAE
Sabia (AS)
Fig.
8.3
7b Veinlets III-IV almost invisible. Leaves glabrous or not very hairy.

Stipules absent. Small glandular grooves or glands, here and there, on the under-
side of the lamina. POLYGALACEAE
e.g. Moutabea (AM)
4b Absence of twining stems. 14
14a Some twigs prehensile (Fig. 8.4a) or axes bearing hooks (Fig. 8.4b) or tendrils
(Fig. 8.4c). (Inflorescence ending in a tendril: Antigonon, POLYGONACEAE). 15
15a Twigs prehensile by thigmonastism. 16
16a Stipules absent. Leaves simple, entire. Venation pinnate. (Glands in stipular
position, f. s, spp.). POLYGALACEAE
e.g. Bredemeyera (AM), Securidaca (mainly AF)
16b Stipules small. Venation camptodromous, scalariform. No glands.
RHAMNACEAE
e.g. Ventilago (AS)
17a Twigs modified into tendrils. 18
18a Leaves with glands. (Tendril in the axil of a leaf Fig. 8.4d).
PASSIFLORACEAE
e.g. Passiflora (AM)
19a AS. Tendril at the base of a leafy twig (tendril more or less perpendicular
to the plane defined by the trunk and the twig). LOPHOPYXIDACEAE
Lophopyxis (AS)
19b AF. Short twig ending in a tendril. Young leaves revolute. Ochrea minute.
Afrobrunnichia (AF), POLYGONACEAE
17b Hooks or prehensile petioles. 20
20a Hooks. (See 21a to 21e). 21
21a AF, AS. Leafy shoots consisting of short internodes and long shoots
bearing terminal hooks (Fig. 8.4e). ANCISTROCLADACEAE
Ancistrocladus (AF, AS)
21b AF. Bifid hook at leaf apex (Fig. 8.4f). Strong heterophylly.
DIONCOPHYLLACEAE
e.g. Dioncophyllum (AF)
21c Paleo. Short twigs bearing a basal pair of hooks: Hugonia. Hooks solitary:
Indorouchera. (L. stipulate). LINACEAE
Hugonia (Paleo), Indorouchera (AS)
21d Paleo. Hook inserted in the base of a twig. Stipules absent. OLACACEAE
Anacolosa (mainly AS)
Fig.
8.4
21e AM. lateral inflorescences ending in a hook (Fig. 8.5a), (or a thick woody
tendril). PASSIFLORACEAE
Ancistrothyrsus (AM)
20b No hooks. (Petiole prehensile). 22
22a Plant weakly woody. SOLANACEAE
e.g. Solanum (e.g. AM)
22b Plant woody. (Laticiferous threads in the leaves). (MANGENOT).
ICACINACEAE
e.g. Desmostachys (AF, MA), Pyrenacantha (AF)
14b No prehensile twigs. Neither hooks nor tendrils. (Plant supporting itself by its
branches or spines, petiolar bases or adventitious roots or plant an epiphyte). 23
23a Stipules (hairy or fringed f. s. spp.) or the two stipule-like prophylls modified into
spines (Fig. 8.5b). 24
24a Ochrea (very short f. s. spp.) leaving an annular scar. Young leaf folding
revolute (Fig. 8.5c). (Xylem lobate). POLYGONACEAE
e.g. Coccoloba (AM)
25a Young trunks with lenticels in longitudinal rows (Fig. 8.5d). Apices with
appressed hairs. (Young internodes angular). DICHAPETALACEAE
Dichapetalum
25b Lenticels different or non-existent or no appressed hairs. (See 26a to 26d).

26a Venation scalariform and camptodromous (Fig. 8.5e). Stipules present.
RHAMNACEAE
e.g. Ventilago (mainly AS)
26a Underside of the lamina with well-marked midrib ending in a small mucro
or an indentation (Fig. 8.5f). Some twigs with two-ranked leaf arrangement.
(stipules modified into spines f. s. spp.). CAPPARACEAE
e.g. Capparis
26c Underside of lamina different. (Lenticels becoming transversely elongated.
LEGUMINOSAE
e.g. Dalbergia
26d AM, in deciduous forests. Leaf arrangement spiral, but becoming two-
ranked at the end of the twigs. (CHAMPAGNAT, MANGENOT, strong odour
of garlic, stipules modified into spines, f. s. spp.). PHYTOLACCACEAE
e.g. Seguieria (AM)
23b Stipules absent (s.t. axillary spines). 27
27a AS, AU, OC. Venation parallel or nearly so.
Fig.
8.5
Plant climbing by means of adventitious roots, (PETIT, CHAMBERLAIN).

Spirotristichous leaf arrangement (Fig. 8.6a). PANDANACEAE
Freycinetia (AS, AU, OC)
28a Indumentum scaly. (Bark fibrous, Twigs oriented backwards).
ELAEAGNACEAE
e.g. Elaeagnus (e.g. AS)
28b Hairs different or none. 29
29a AF. Liana bearing zigzagging plagiotropic branches (MANGENOT).
ICACINACEAE-METTENIUSACEAE
Rhaphiostylis (AF)
30a Leaves coriaceous. Venation not distinctly reticulate. (See 31a to 31c).
31a Rhytidome scaly (Fig. 8.6b). Basal V II grouped. ERICACEAE
e.g. Satyria (AM), Agapetes (AS, AU, OC)
31b AM. Petiole short. (Strong heterophylly between understorey-
leaves and canopy-leaves, lenticels in longitudinal rows (Fig. 8.6c),
underside of lamina with glandular striae (Fig. 8.6d).
MARCGRAVIACEAE
e.g. Marcgravia, Norantea (AM)
31c MA, AS. Lianas with stems leaning on their support. Petiole with
articulation. OXALIDACEAE
Dapania (MA, AS)
30b Leaves not coriaceous or venation distinctly reticulate. 32
32a Stems bearing adventicious roots.
Main shoot with very long internodes. Annular prophyll. Petiolar
base sheathing the stem. (Roots modified into tendrils, f. s.
spp. (Fig. 8.6e). ARACEAE
e.g. Heteropsis (AM)
32b No such roots.
Leaves thick. (Hemiepiphyte). L. grouped in clusters: Markea.
SOLANACEAE
e.g. Solandra, Markea (AM)
32c AF. Leaf margin entire with a intramarginal vein or a fimbrial vein
(Fig. 8.6f). THYMELAEACEAE
Dicranolepis (AF)
Fig.
8.6
Group 55—Lianas without latex or coloured exudate, alternate leaf arrangement; leaves simple;
venation palmate, tripliveined or campylodromous
1a Some stems twining (Fig. 8.7a). 2

2a Presence of stipules. 3
3a MA, AS. Bark with a network of fibres (Fig. 8.7b). Petiole distally swollen or very
short. (Stems angular). MALVACEAE
Byttneria (MA, AS)
3b Bark without a network of fibres. Fissured periderm. (Base of lamina with glands).
EUPHORBIACEAE
e.g. Omphalea
2b Stipules absent. (Prophylls simulating stipules: Aristolochia spp). 4
4a Base of petiole decurrent to the stem. 5
5a Leaves or bark aromatic. (Herb, nodes swollen (Fig. 8.7c), prophylls appressed to the
stem, f. s. spp. (Fig. 8.7d)). ARISTOLOCHIACEAE
e.g. Aristolochia (mainly AM)
5b Venation campylodromous, DIOSCOREACEAE
Dioscorea
4b Petiolar base not decurrent. 6
6a Petiole swollen at both ends (Fig. 8.7e). Young twigs soon becoming woody.
(Section of trunk with radiate concentric zones). (Leaves peltate, e.g. Cissampelos).
MENISPERMACEAE
Numerous genera
7a Long (prehensile?) petiole able to bend to reorientate the lamina.
ICACINACEAE
e.g. Pyrenacantha (AF, AS), Phytocrene (AS)
8a Leaf margin entire or lobate but margin not toothed. Underside of lamina with
appressed, stellate or spherical hairs (use a hand lens). (Woody plant with com-
pressible young stem or herb). (Latex?). CONVOLVULACEAE
e.g. Ipomoea
8b Leaves toothed. (Hairs simple). ICACINACEAE
e.g. Pyrenacantha (AF), Natsiatum (AS)
1b Absence of twining stems. 9
9a Twigs prehensile (Fig. 8.7f) or presence of tendrils or of hooks. 10
10a Twigs prehensile, becoming woody. 11
11a Stipules present. Twigs ending in a hook: Bandereia, twig-tendril: Bauhinia.
Fig.
8.7
LEGUMINOSAE
e.g. Bandereia (AF), Bauhinia
11b Stipules absent. Twigs hook-like. Bark aromatic. HERNANDIACEAE
Sparattanthelium (AM)
10b No prehensile woody twigs. 12
12a Presence of tendrils. 13
13a Axillary tendrils. 14
14a Stipules present. (See 15a to 15c). 15
15a Petiole or underside of lamina with glands. (L. diversely lobate). Swollen
spines or stems: Adenia. Tendrils (Fig. 8.8a). PASSIFLORACEAE
e.g. Passiflora (AM, AS, AU), Adenia (Paleo)
15b Leaf margin entire, without glands. (Tendrils simple or bifid, (Fig. 8.8b)
lenticels becoming transversely swollen, leaves not lobed). LEGUMINOSAE
Bauhinia
15c Leaves toothed, not lobed. Venation scalariform (Fig. 8.8c). (Tendrils s.t.
circinate). RHAMNACEAE
e.g. Helinus (AF, MA), Gouania
14b AS. Stipules absent. Petiole without glands. Leaves cordate.
ERYTHROPALACEAE
Erythropalum (AS)
13b Tendrils inserted otherwise. 16
16a Tendrils oppositifoliate (Fig. 8.8d). Stipules present. (Rays of phloem con-
spicuous, L. toothed, teeth glandular). VITACEAE
e.g. Tetrastigma (AS, AU), Cissus
16b Tendrils not oppositifoliate. 17
17a Two tendrils inserted on the petiole (Fig. 8.8e), (reduced into stipules, or
leaves spiny, f. s. spp.). SMILACACEAE
Smilax
17b Herbaceous vine. Tendril (or short twig bearing several tendrils) forming a
right angle with the stem and the petiole. CUCURBITACEAE
Numerous genera
12b Plant without tendrils (plant bearing hooks). 18
18a Stipules present. (Tendrils becoming woody, hook-like (Fig. 8.8f), venation
scalariform, f. s. spp.). RHAMNACEAE
e.g. Gouania (mainly AM)
Fig.
8.8
18b AM. Stipules absent. Leafless short shoots modified into hooks.
HERNANDIACEAE
Sparattanthelium (AM)
Group 56—Lianas without latex or coloured exudate (s.t. gum); opposite or whorled leaf
arrangement
1a AF, MA, AS. Trunk sympodial, each module ending in a tendril. (Xylem in blocks,
Fig. 8.9a). ICACINACEAE
Iodes (AF, MA, AS)
2a Trunk twining, (but twigs might not be). 3
3a Internodes swollen at the base. 4
4a Leaves glabrous. Venation “polygonal”, without blind veinlets (Fig. 8.9b). (Lati-
ciferous threads in the leaves (Fig. 8.9c), woody liana). GNETACEAE
Gnetum
4b Venation different. L. without laticiferous threads. Weakly woody. 5
5a AM. Venation pinnate. ACANTHACEAE

Mendoncia (AM, AF, MA)
5b AF, MA, AS. Stems not quadrangular. Leaves tripliveined (entire, toothed or lobate).
ACANTHACEAE
e.g. Thunbergia (AF, AS)
3b Base of internodes not swollen. 6
6a Interpetiolar ridge (Fig. 8.9d). 7
7a Stipules (s.t. inserted on the petiole). 8
8a Stipules interpetiolar (Fig. 8.9e). RUBIACEAE
e.g. Manettia (AM), Atractogyne, Sherbournia (AF), Paederia
8b Stipules free (Fig. 8.9f) or, if fused, not extending from one petiole to the other. 9
9a AM. Indumentum of erect or oblique hairs. (Numerous lenticels).
TRIGONIACEAE
Trigonia (AM)
9b Mainly AM. Hairs appressed. (Numerous lenticels, stipules small, L. with
glands: e.g. Banisteriopsis, Stigmaphyllon, or not, e.g. Tetrapteris, interpetiolar
stipules f. s. spp.). MALPIGHIACEAE
e.g. Hiptage (MA, AS, OC)
7b Stipules absent. 10
Fig.
8.9
10a External wood with long parallel furrows or ridges (Fig. 8.10a). Leaves without
glands. ASTERACEAE
e.g. Mikania (AM)
10b External wood not furrowed or leaves with glands. 11
11a Stems, leaves or buds hairy. 12
12a Buds with erect hairs. OLEACEAE
e.g. Jasminum bifarium (AS)
12b Buds with appressed hairs (Fig. 8.10b). MALPIGHIACEAE
e.g. Flabellaria (AF), Heteropteris (AM)
11b Plant glabrous. (Leaf margin entire). (See 13a to 13c). 13
13a AS. Stems distinctly quadrangular. Leaf margin entire, tripliveined.
OLEACEAE
Myxopyrum (AS)
13b AS. Young internodes angular. Leaves not tripliveined (GELSEMIACEAE:

Gelsemium). LOGANIACEAE
e.g. Gardneria (AS)
13c AS. Young internodes cylindrical. Leaves supratripliveined (Fig. 8.10c).
LOGANIACEE
Strychnos (e.g. AS)
6b No interpetiolar ridge. 14
14a Venation campylodromous (Fig. 8.10d) or supratripliveined. (Leaves lobate,
prickly stems, f. s. spp.). DIOSCOREACEAE and RIPOGONACEAE
Ripogonum (NG, AU, OC), Dioscorea
15a Leaves tripliveined (Fig. 8.10e) or venation palmate. 16
16a L. tripliveined or venation palmate. L. opposite or 3-whorled. (Stems quad-
rangular). LAMIACEAE
e.g. Clerodendrum
16b Venation pinnate. 17
17a Rhytidome becoming scaly. (Wood with strands of phloem (Fig. 8.10f),
bark with disagreeable smell, underside of L. with coloured spots, leaves with
glands, f. s. spp.). COMBRETACEAE
e.g. Combretum
17b Rhytidome different. 18
Fig.
8.10
18a Petiole neither distally swollen nor articulate. 19

19a Mainly AM. Minute stipules or hairs appressed. (Leaves with
glands). MALPIGHIACEAE
e.g. Banisteriopsis (AM), Flabellaria (AF), Stigmaphyllum (AM, AS)
19b Stipules absent. No appressed hairs. 20
20a Venation distinctly reticulate, brochidodromous (L. 3-whorled
f. s. spp., petiole modified into a thorn (Fig. 8.11a): Clerodendron).
LAMIACEAE-VERBENACEAE
e.g. Aegiphila, Petrea (AM)
20b Leaf arrangement opposite, subopposite or occasionally alternate.
(Venation camptodromous (Fig. 8.11b), blade with basal glands in the
American Combretum, bark with disagreeable smell f. s. spp.).
COMBRETACEAE
e.g. Combretum
18b Petiole distally swollen. (Leaves 3-whorled). OLEACEAE

e.g. Jasminum (AF, AS)
2b Trunk not twining. 20
20a Existence of twining or prehensile (thigmonastic) twigs (Fig. 8.11c). 21
21a Stipules small or minute (young stems have to be observed). 22
22a Apices with appressed hairs. (Leaves with glands).
Some twining twigs. (Interpetiolar ridge). MALPIGHIACEAE
e.g. Banisteriopsis (AM)
22b No appressed hairs. Leaves without glands.
(Thigmonastic twigs, xylem lobate (Fig. 8.11d), in blocks or in rings Fig. 8.11e).
(Laticiferous threads in leaves, L. subopposite, orange-coloured layer under the
rhytidome, f. s. spp.). CELASTRACEAE
incl. Hippocrateoideae, e.g. Salacia
21b Stipules absent.
Small twining twigs or twigs forming hooks. (When small leaflets at the base of a
tendril, maybe a BIGNONIACEAE). Fibrous bark, branches do not break easily).
THYMELAEACEAE
e.g. Craterosiphon (AF), Enkleia (AS)
20b Absence of twining or prehensile twigs. 23
23a Plant climbing by means of hooks (Fig. 8.11f). (See 24a to 24c).
24a Hooks subtended by a leaf or a scale-leaf. Leaves supratripliveined. (Wood with
strands of phloem). LOGANIACEAE
Strychnos
Fig.
8.11
24b Hooks subtended by a laminar leaf (Fig. 8.12a). Venation pinnate. RUBIACEAE
e.g. Canthium (Paleo), Uncaria
24c Old petiolar base hook-like. Stems quadrangular. L. tripliveined or venation
palmate (Fig. 8.12b). (Leaves 3-whorled f. s. spp.). LAMIACEAE
e.g. Clerodendrum
23b Plant without hooks. 25
25a Stipules or stipular expansions on petioles. 26
26a Interpetiolar stipules. Epiphyte, e.g. Hillia (AM). (If appressed hairs, L. with
glands, and stipules minute: MALPIGHIACEAE). RUBIACEAE
e.g. Sabicea (AM), Canthium (Paleo)
26b Stipules small. Shoot short modified into spines (Fig. 8.12c). RHAMNACEAE
Scutia (AM, MA)
25b Stipules absent, or stipular expansions. 27

27a Rhytidome becoming scaly. L. with glands or glandular trichomes. (Bark with
disagreeable smell f. s. spp.). COMBRETACEAE
Combretum
27b Rhytidome different. 28
28a Venation brochidodromous. Periderm more or less shiny.
Stem almost cylindrical, not quadrangular. Lenticels protruding.
SCHLEGELIACEAE (* BIGNONIACEAE)
Schlegelia (AM)
28b Venation or periderm different. 29
29a Paleo. Stem not spiny. 30
30a Leaves tripliveined or venation pinnate (Fig. 8.12d). (Petiole with an
articulation f. s. spp.). OLEACEAE
Jasminum (AF, AS)
30b Plant climbing by means of divaricate (zigzagging) twigs. (Interpetiolar
ridge). LOGANIACEAE
Strychnos
29b Stems spiny or thorny. 31
31a L. supratripliveined (Fig. 8.12e), somewhat coriaceous. Petiole not
articulate. (Wood with strands of phloem (Fig. 8.12f). LOGANIACEAE
Strychnos
31b Venation pinnate. Leaves thin and soft. (Anisophylly).
NYCTAGINACEAE
e.g. Pisonia aculeata (AU).
Fig.
8.12
Group 57—Lianas without latex or coloured exudate; leaves compound
1a Opposite or whorled leaf arrangement (or seeming “alternate” due to a pronounced aniso-
phylly in a few spp.). 2
2a Leaves with a terminal tendril (Fig. 8.13a) or a scar at the apex of the rachis.
Leaves modified into trifid hooks for Dolichandra (AM). (Trunk sections showing 4 poles
of phloem). BIGNONIACEAE
Numerous American genera: e.g. Arrabidaea, Adenocalymma, Dolichandra
2b Leaves without tendril or apical scar (L. imparipinnate or trifoliolate). 3
3a Petiole prehensile. AF, AU, North temp.. RANUNCULACEAE

Clematis (AF, AU)
3b Petiole not prehensile. 4
4a L. trifoliolate (L. articulate). Jasminum (Paleo), OLEACEAE
4b Twining stems and leaves imparipinnate. BIGNONIACEAE
Pandorea (AS, AU)
1b Spiral or two-ranked leaf arrangement. 5
5a Trunk or its extremity twining (Fig. 8.13b). (Trunk twisted but plant not twining for some
LEGUMINOSAE).
6
6a Leaves palmate. 7
7a Section of trunk or branches with radiate concentric zones. MENISPERMACEAE
e.g. Disciphania (AM), Burasaia (MA)
7b Trunk and branches different. 8
8a Paleo. Petioles prehensile (Fig. 8.13c). Petiolar base decurrent. HERNANDIACEAE
Illigera (mainly AS)
8b Petioles different. Petiolar base not decurrent. (Xylem lobate Fig. 8.13d), young
stem with well-developed pith). CONVOLVULACEAE
e.g. Merremia
6b Leaves (bi)pinnate or trifoliolate (Fig. 8.13e). 9
9a Stipules present. (Venation protruding). LEGUMINOSAE
(See key D), e.g. Derris (AM), Dioclea (mainly AM), Mucuna
9b Stipules absent. 10
10a Leaves pinnate or trifoliolate. 11
11a Trunk sympodial. (Basal secondary veins ascending, embossed venation,
xylem in rings, f. s. spp. Fig. 8.13f). (L. trifoliolate: e.g. Pseudoconnarus, AM).
Fig.
8.13
CONNARACEAE
e.g. Agelaea (Paleo), Connarus
11b Leaves trifoliolate. Herbaceous shoots arising from a tuber. (Stem prickly
Fig. 8.14a), venation Dicot-like!). DIOSCOREACEAE
e.g. Dioscorea dregeana (S AF, cult.)
5b Trunk or stems not twining (but could be prehensile). 12
12a Some twigs prehensile, but not like hooks or tendrils. (Xylem in rings or simple).
LEGUMINOSAE
e.g. Millettia (Paleo)
12b Hooks or tendrils present, or other climbing system. 13
13a Tendrils are axillary or oppositifoliate, not leaf appendices. 14
14a Tendrils circinnate (Fig. 8.14b) or bifid. 15
15a Leaf with more than 3 leaflets. (Leaflets toothed). 16
16a Petiole with glands. PASSIFLORACEAE
e.g. Passiflora
16b Petiole without glands. SAPINDACEAE
e.g. Paullinia, Serjania (AM).
15b Leaves bifoliolate or bilobate. (Young trunk flattened). LEGUMINOSAE
e.g. Bauhinia
14b Tendrils oppositifoliate (Fig. 8.14c). (Rays of phloem). VITACEAE
e.g. Rhoicissus (AF), Tetrastigma (AS, AU), Cissus, Cyphostemma
13b Tendrils or hooks are leaf parts, or plant climbing with other devices. 17
17a Hooks or tendrils present. 18
18a Rachis of leaf ending in a tendril (Fig. 8.14d). 19
19a Petiolule pulvinate. L. bipinnate (Fig. 8.14e). LEGUMINOSAE
See Key D, Entada,
19b AM. Petiolule not pulvinate. L. pinnate. POLEMONIACEAE
Cobaea (AM)
18b No terminal tendrils. 20
20a Plant climbing by the means of woody tendrils. LEGUMINOSAE
Mimosoideae: e.g. Pseudoprosopis (AF)
20b Paleo. Petiole twining (Fig. 8.14f) or modified in a hook. Leaves trifoliolate.
Petiolar base decurrent. HERNANDIACEAE
Illigera (mainly AF, AS)
Fig.
8.14
17b Plant without tendrils or hooks. (Weakly prostrate, climbing by means of hairs,
spines, roots, petiolar bases or plant leaning on its branches). 21
21a Venation parallel (Fig. 8.15a). (MONOCOTS). 22
22a Grapnel-climber. Support provided by a cirrus (extended leaf axis) or a

flagellum (a modified inflorescence axis fused to the internode and leaf sheath).
ARECACEAE
e.g. Desmoncus (AM), Ancistrophyllum (AF), Calamus (AS)
22b Leaves simple, distinctly bilobate.
AM. Plant climbing by means of adventitious roots. CYCLANTHACEAE
e.g. Asplundia, Evodianthus (AM)
21b Venation different. (See 23a to 23c).
23a Sheathing petiolar base (Fig. 8.15b) or bark aromatic. (Clamp-roots or
bristly hairs and L. pinnate or trifoliolate). ARALIACEAE
e.g. Aralia (AS), Cephalaralia (AU)
23b Petiole not sheathing. Smell different. (Leaves compound (bifoliolate
Fig. 8.15c): Bauhinia), rarely simple (bilobate: Bauhinia), leaves or stems spiny
Fig. 8.15d: Acacia, Mezoneuron). LEGUMINOSAE
e.g. Machaerium (AM), Mezoneuron (AS, AU, OC)
See Key D, Acacia, Bauhinia,
23c (L. trifoliolate). Strong short shoots modified into spines: Luvunga. Epi-
dermal spines: e.g. Toddalia. Translucent dots (Fig. 8.15e) or aromatic smell.
Petiole not sheathing. RUTACEAE
e.g. Luvunga, Toddalia (AS)
Fig.
8.15
Part III
Description of the Families
Acanthaceae to Asteraceae
9
9.1 Acanthaceae: 36-37-38-56
200 genera and more than 3800 species, order Lamiales. Coloured sap absent, plants not aromatic.
Internodes typically swollen at its base (Fig. 9.2) (but not in Avicennia or Hygrophila).
Prophylls modified into spines in Barleria (Fig. 9.4). Sectorial anisoclady (Fig. 9.2), i.e. the
longest twigs are disposed in the same quarter of a cylinder. Opposite phyllotaxy, rarely spiral
(Acanthus). Leaves simple and entire (Fig. 9.2) (rarely with spiny teeth: Acanthus spp.). Stipules
absent. Venation camptodromous (Fig. 9.5). Inflorescences in spikes, racemes or 1-flowered.
Flowers often subtended by bracts (Fig. 9.1), which may be the showier part of the inflorescence.
Corolla tubular, with strong (Fig. 9.4) or weak (Fig. 9.5 Ruellia) bilateral symmetry, or almost radial
(Fig. 9.3, Avicennia). 4–5 petals and 2–4 stamens, corollas diversely coloured according to the
species. Ovary superior with axile placentation. Fruits capsules (Fig. 9.3), s.t. with explosive
opening (e.g. Asystasia gangetica).
Fig. 9.1 Aphelandra pulcherrima, spike of flowers with conspicuous bracts (b), Venezuela. Avicennia marina, back
mangrove, Bali
Trees Avicennia (Fig. 9.1, mangroves). Shrubs Barleria (Fig. 9.4, Pantrop) Bravaisia (Fig. 9.4,
AM), Justicia (Pantrop), Pseuderanthemum carruthersii (Fig. 9.5, OC, ornam.). Twining lianas
(Mendoncia, AM; Fig. 9.2 Thunbergia, Paleo). Very numerous herbs (Aphelandra, Eranthemum,
Hygrophila, Justicia, Ruellia, etc.), Acanthus spp. (Fig. 9.3) being prickly plants of salt marshes and
mangroves.
https://doi.org/10.1007/978-3-031-05942-1_9
142 9 Acanthaceae to Asteraceae
Architectural models:
Similar families: TOMLINSON: Asystasia gangetica, Peristrophe
LAMIACEAE share more or less the same angustifolia, Ruellia simplex.
architectural models and bilateral, coloured LEEUWENBERG: Pseuderanthemum carruthersii,
flowers. Lepidagathis formosensis.
ATTIMS: Avicennia germinans.
! Several species (e.g. Strobilanthes callosus, India) have a gregarious flowering, they burst into
simultaneous blooming and cover entire hill ranges and plants die off after fruiting [1].
! Sectorial anisoclady in Acanthaceae was already observed as early as the middle of the nine-
teenth century by Wichura [2].
Fig. 9.2 Swollen internode bases (!). Sectorial anisoclady, longest lateral stems in the same quadrant. Leaves,
a cordate, Thunbergia, b sagittate, Thunbergia, c usual type, i.e. camptodromous venation
Fig. 9.3 Acanthus ebracteatus, mangrove, Bali. Aphelandra scabra, unripe capsules, Belize. Avicennia marina,
flowers regular, Bali
9.2 Achariaceae: 15-16-18-20-27 143
Fig. 9.4 Barleria sp., spines (sp), petals almost free, Kerala. Bravaisia berlandieriana, swollen internode bases,
Belize. Bravaisia berlandieriana, flower with bilateral symmetry, Belize
Fig. 9.5 Eranthemum nervosum, camptodromous venation, cult. Pseuderanthemum carruthersii, 4-petals corolla,
cult., Bali. Ruellia tuberosa, 5-lobed corolla, natur. (orig. AM), Bali
9.2 Achariaceae: 15-16-18-20-27
32 genera and 170 species, order Malpighiales. Coloured sap absent and plants not aromatic, but
s.t. bark with a smell of bitter almonds (Caloncoba, Lindackeria, Pangium). Inner bark not fibrous,
except in Pangium edule, s.t. with sclerenchymatic inclusions (Caloncoba). Twigs profusely lenti-
cellate (Fig. 9.7 Gynocardia, Pangium) Stipules absent. Leaves simple, alternate (s.t. also in pla-
giotropic branches, Ryparosa). Lamina entire or toothed, s.t. cordate, the petiole often long and
swollen distally (Lindackeria, Fig. 9.8 Ryparosa), or petiole very short (Phyllobotryon). Venation
camptodromous (Lindackeria, Fig. 9.8 Ryparosa) or palmate (Acharia, Caloncoba, Pangium).
Inflorescences variable, but epiphyllous (Fig. 9.8) in Phyllobotryon (AF) or s.t. cauliflorous (Fig. 9.8
Carpotroche, Kiggelaria). Flowers small, actinomorphic, unisexual. Stamens usually numerous
(Fig. 9.6) or in number of five (Hydnocarpus). Ovary superior with parietal placentation. Fruits
coriaceous berries (Kiggelaria Africana, Fig. 9.8 Pangium edule), tuberculate capsules (Fig. 9.8
Lindackeria) or prickly capsules (Caloncoba). Seeds usually with coloured aril.
Fig. 9.6 Caloncoba echinata, flowers with numerous stamens, Kebun Raya, Java
Trees (Hydnocarpus, Ryparosa, Paleo; Kiggelaria, AF-S; Pangium edule, Malaysia), shrubs
(Carpotroche, AM). Herbs (Acharia tragodes, South Africa).
CORNER (Fig. 9.8): Phyllobotryon (AF).
Similar families: RAUH: Caloncoba (AF), Lindackeria spp.? (AM).
Leaves of several EUPHORBIACEAE and almost all
AUBRÉVILLE (Fig. 9.8): Pangium edule (AS).
MALVACEAE also have petioles swollen distally.
ROUX: Caloncoba cf. echinata, Hydnocarpus spp.,
Ryparosa javanica (AS).
CHAMPAGNAT: Carpotroche spp. (AM),
Gynocardia odorata (Assam, Burma).
Predominantly in the Old World tropics. Except for timber (Kiggelaria africana, Natal maho-
gany), the family is of little interest for forestry and agronomy. Pangium edule (‘kepayang’) has large
poisonous fruit (hydrocyanic acid) which can be made edible by fermentation (Malaysia). The oil
extracted from the seeds of several species of Caloncoba, Carpotroche and Hydnocarpus was used
for healing leprosy, or other skin diseases. Antiseptic juice: Pangium edule.
Fig. 9.7 Gynocardia odorata, trunk with protruding lenticels, Kebun Raya, Java. Phyllobotryon sp., CORNER’s
model, shortly petiolate leaves, Cameroon. Ryparosa javanica, L. clustered distally on plagiotropic branches, Kebun
Raya, Java
9.3 Actinidiaceae: 24-28 145
Fig. 9.8 Pangium edule, AUBRÉVILLE’s model, extremity of a branch with a hanging fruit. Lindackeria paludosa,
branches orthotropic, leaves long-petiolate, petiole pulvinate (pv) at both ends, tuberculate capsule (Cp). Phyllobotryon
sp., CORNER’s model, fallen litter accumulates in leaf axils, epiphyllous flowers are inserted on main vein (V).
Kiggelaria africana, dangling fruits, ca. 20 cm long. Carpotroche platycarapa, trunciflory. Ryparosa javanica,
plagiotropic branch with spirally inserted leaves, petiole pulvinate (pu) at both ends
! Before APG studies, this family was part of the “Flacourtiaceae miscellany”. Achariaceae rep-
resent one of the few examples of an arborescent tropical family where herbs evolved only in the
Southern Hemisphere (i.e. Acharia tragodes).
9.3 Actinidiaceae: 24-28
3 genera and 360 species. Order Ericales. Trees and shrubs, lianas in Actinidia. Coloured sap absent.
Plants not aromatic. Spiral arrangement of leaves. Stipules absent. Leaves simple, usually toothed
and with secant venation. Stems and leaves strongly pubescent, with thick hairs (Fig. 9.10).
Inflorescences cymose (Fig. 9.9) (more exactly, thyrses of scorpioid cymes), lateral, s.t. reduced to a
single flower in Asiatic Saurauia. Flowers white or pink, with free petals and numerous stamens
(Fig. 9.9). Fruits berries with axile placentation (Fig. 9.10), more rarely capsules.
A small South American and Asiatic family, placed near THEACEAE, both are absent in Africa
(Laurasian origin?). Species grow mostly in montane forests. Saurauia has a discontinuous
distribution (AM and AS), but Actinidia is restricted to Indomalesia. Actinidia sinensis var. deliciosa
is widely cultivated for its fruits (kiwis).
In past times [3] Saurauia was placed in Dilleniaceae, a family currently placed in the separate
order Dilleniales.
Similar families: Architectural models:

DILLENIACEAE also have showy flowers AUBRÉVILLE (Fig. 9.10): Saurauia spp. (AS)
with numerous stamens and pubescent leaves OLDEMAN: Actinidia sinensis.
with thick hairs.
Fig. 9.9 Actinidia sinensis var. deliciosa, flowers with radial symmetry and numerous stamens, in cultivation.
Saurauia sp., leaves with thick rust-coloured hairs, Seram, Indonesia. Saurauia sp., cymose infloresence,
Fig. 9.10 Saurauia sp., Pubescence of thick hairs, part of a branch plagiotropic by apposition, Thailand. Saurauia
madrensis, berry with axile placentation (AM-C)
9.4 Anacardiaceae: 3-24-28-29-44-C 147
9.4 Anacardiaceae: 3-24-28-29-44-C
82 genera and 950 species, order Sapindales. Bark often fissured (Fig. 9.12). Bark with resiniferous
vessels secreting a kind of latex (Fig. 9.13) which can be toxic or vesicant (Comocladia, Lithraea,
Metopium, Rhus) or a resinous gum (Anacardium, Buchaniana lanzan, Schinopsis). Big buttressed
trees (Darcontomelon). Exudate s.t. turning brown or black (Fig. 9.13 Gluta, Rhus spp.). Spiral
arrangement of leaves (opposite in Bouea). Leaves imparipinnate, winged rachis in Lithraea, ter-
minal leaflet (Dracontomelon, Tapirira), trifoliolate (African Rhus, Sorindeia) or simple (Anac-
ardium, Mangifera). Stipules absent. Rhythmic branching, leaves s.t. crowded towards the end of
GUs (Fig. 9.13) (Dracontomelon, Gluta, Thyrsodium). Leaflets entire, s.t. toothed (Rhus). Venation
often with secondary veins abruptly bend near lamina margin (Fig. 9.12 Anacardium, AM, Par-
ishia, Indomalesia) or join to form an intramarginal vein (Spondias) or secant (Astronium, Fig. 9.12
Ozoroa). Inflorescences paniculate, cymose (Fig. 9.14). Flowers small with 5 petals basally connate,
5 or 10 stamens inserted on the margin of a glandular disk. Carpels free (Buchaniana), fused
(Dracontomelon, Spondias, Tapirira) or solitary (Gluta, Mangifera). Fruits drupes (Fig. 9.11), s.t. 5-
winged (Swintonia Fig. 9.16), in Anacardium the fruit is borne on a fleshy peduncle (Fig. 9.11).
Fig. 9.11 Anacardium occidentale, drupes inserted on fleshy peduncles, Orinoco Delta, Venezuela
Similar families:
CORNER: Trichoscypha (AF).
BURSERACEAE (pulvinate
LEEUWENBERG: Rhus typhina
petiolules).
(AM-N, ornam.)
MELIACEAE (venation
usually camptodromous). SCARRONE: Anacardium
occidentale, Mangifera indica.
Trees Buchanania, Gluta, Mangifera, Parishia (AS); Metopium (AM-C), Rhus (AM, AS); Sor-
indeia (AF, MA), very tall trees in Asia: Dracontomelon dao, Swintonia spp. can reach more than
50 m in height, shrubs (Ozoroa, AF).
Fig. 9.12 a Secondary veins typically bend, b intramarginal vein, Spondias, c Venation secant (e.g. Ozoroa). d Bouea
macrophylla, leaves opposite, rhythmic growth, cataphylls (c)
Distribution pantropical, with a bulk of species in Indomalayan forests. Timber: Gluta, Parishia
(Indomalesia). Fruit trees: mango tree (Mangifera indica), ‘monbin’ (Spondias mombin, AM),
‘pomme de Cythère’ Spondias dulcis, AS-OC), plum mango (Bouea macrophylla, Malaysia), ‘buah
rao’ (Dracontomelon daho); cashew tree, (Anacardium occidentale, AM), ‘marula’ (Sclerocarya
birrea, AF-S). Source of black laquer: Gluta usitata (AS). Schinus molle (Fig. 9.15), native to
Andean valleys, is frequently cultivated.
! Caution with Anacardiaceae! Several, usually harmless, edible fruits of Anacardiaceae, such as
mangos, can cause severe anaphylactic reactions to sensitive people. Woodcutters have to protect
themselves against dermatitis (vesicant sap of Metopium, Parishia, etc.).
Fig. 9.13 Campnosperma auriculata, leaves simple, Sumatra. Dracontomelon dao, clusters of pinnately compound
leaves. Gluta usitata, latex turning black, Thailand
9.5 Ancistrocladaceae: 54 149
Fig. 9.14 Mangifera indica, simple leaves, terminal and cymose inflorescences, Belize. Metopium brownei,
translucent resinous gum, Belize. Pseudospondias microcarpa, latex viscous and whitish, Kebun Raya, orig. tropical
Africa
Fig. 9.15 Schinus molle, cult. Scelocarya birrea, fissured bark and glaucous leaflet, Botswana
Fig. 9.16 Spondias caerulea, deciduous fruiting tree, Venezuela. Spondias mombin, stout trunk apex, with leaves
clustered (here petiolar bases), typical for many Anacardiaceae. Swintonia schwenkii, 5-winged drupe
9.5 Ancistrocladaceae: 54
Monogeneric family: Ancistrocladus, 27 species. Order Caryophyllales. Paleotropical lianas. Bran-

ches sympodial ending in a hook (Fig. 9.17). Leaves spiral. Flowers small, in spikes or panicles.
Fruits nuts embedded in an accrescent calyx. Some medicinal (alkaloids).
Fig. 9.17 Ancistrocladus tectorius, Sakaerat, Thailand. Liana bearing hooks. Leaves clustered, sympodial branch
9.6 Anisophylleaceae: 13-20
4 genera and 71 species. Order Cucurbitales. Trees and shrubs, s.t. accumulating aluminium. Leaves
appearing two-ranked for several species, because of a peculiar arrangement of smaller leaves on
the upper side of the twig (Fig. 9.18), the larger leaves being disposed in the same plane (especially in
Anisophyllea disticha, AS). Leaves simple, entire. Venation pinnate, pseudopalmate (Fig. 9.18).
Corolla usually with 4-petals. Ovary superior with pendulous ovules. Fruits are drupes.
Fruit trees: Anisophyllea laurina (Monkey apple, AF-W). MASSART’s model is frequent in
Anisophyllea.
! The name of this family is obviously based upon the marked variation in leaf size on the same
shoot (anisophylly).
a b
c d
Fig. 9.18 Anisophyllea sp., the smaller leaves are inserted above the stem, Cameroon. b. A. cinnamomoides,
pseudopalmate venation. c. Anisophyllea fallax, drupes with pendulus ovules. d. Anisophyllea sp. MASSART’s model,
Sumatra
9.7 Annonaceae: 10-54 151
9.7 Annonaceae: 10-54
107 genera and 2450 species, order Magnoliales. Sectioned bark (and s.t. leaves too) gives out a
pleasant smell and displays a network of fibres visible to the naked eye (Fig. 9.19). Stems s.t. with
septate pith. Lianas climbing with prehensile twigs (Fig. 9.24 Uvaria). Trunk or branches often
lenticellate (Fig. 9.22). Branches usually with frequent meristem abortions (Fig. 9.19). Leaf
arrangement two-ranked (Fig. 9.19) (exceptionally spiral for the American genus Tetrameranthus).
Leaves simple, entire. Stipules absent. Young leaf-folding conduplicate (Fig. 9.23). The character of
a raised (Unonopsis) or sunken (Annona, Guatteria, Xylopia) main vein on the upper side of the blade
might be useful for the identification of many genera. Inflorescences s.t. oppositifoliate. Trunciflory
or Cauliflory in Anaxagorea (AM), Goniothalamus (AS), Sapranthus (AM-C), Stelechocarpus
(Fig. 9.23, AS). Flowers usually solitary, somewhat large or medium-sized, with 2 or 3 cycles of
tepals (Fig. 9.22), stamens numerous. If flowers tiny, then pollinisated by thrips (Bocageopsis, AM).
Fruits are formed by stipitate free carpels (Fig. 9.23) (Anaxagorea, Desmos, Guatteria, Hexalobus,
Unonopsis, Uvaria) or by carpels fused into a syncarp (Fig. 9.20) because of the development of a
fleshy receptacle (Annona, Guatteria), or s.t. a fleshy capsule (Fig. 9.24 Xylopia) (Fig. 9.21).
Trees (Polyalthia, Xylopia, etc.) or woody lianas (Artabotrys, Paleo), Desmos (AS, OC), Uvaria
(Paleo). Leaves reaching 70 cm in length for Piptostigma calophyllum (AF)! [4].
Fig. 9.19 a. Typical fibrous bark. Anaxagorea javanica, sympodial branch with frequent meristem abortions (X).
b. Inflorescences one-flowered, oppositifoliate, flower (fl) with 3 internal tepals and 3 external tepals, Thailand
Fruit trees: ‘guanábano’, ‘corosol’ or ‘chirimoya’ (Annona cherimola, A. muricata, A. squamosa)

or source of perfumes: ‘ilang-ilang’ (Cananga odorata). Most genera are endemic to only one
continental mass, Xylopia being pantropical, Polyalthia paleotropical. Anaxagorea has an AM-AS
discontinuous distribution and is thought to have a Laurasian origin [5]. Anaxagorea is sister group of
all other extant genera of the family [6, 7].
! Annonaceae form a very old group, the origin of which dates back to an epoch where Laurasia
and Gondwana were still not very remote. Interestingly, Annonaceae have experimented with dif-
ferent forms of TROLL’s models (Fig. 9.25). The family is distributed in all evergreen forests and are
represented by small trees of the understorey, rarely by tall trees.
! Flowers of the 100 Myr old Annonaceae are pollinated by the most ancient insects: beetles or
Coleoptera [8].
! Wood of Annonaceae is characterized by very low fibre content and no visible gelatinous layer.
As a consequence, uprighting of tree stems (a mechanism characteristic of TROLL’s model) is caused
by tensile forces in bark, not in wood [9, 10].

MYRISTICACAE: but bark producing an ROUX: Unonopsis (AM), Cananga, Polyalthia (AS),
astringent phenolic exudate, usually red Cleistopholis (AF), Isolona (AF, MA), Uvaria (Paleo),
coloured. Uvariopsis (AF), Xylopia (Pantrop).
LAURACEAE: have different architectural PETIT: Duguetia, Guatteria (AM), Artabotrys (Paleo),
models and young stems angular or grooved. TROLL (Fig. 9.20): Anaxagorea (AM), Cleistopholis (AF),
Rollinia spp. (AM), Monodora (AF), Goniothalamus (AS).
Fig. 9.20 Annona muricata, syncarp, in cultivation, Venezuela. Annona sp., small tree conforming to TROLL’s model,
Belize. Anaxagorea sp., the fruit is a fascicle of stipitate carpels, Venezuela
Fig. 9.21 Anaxagorea javanica, corolla with 2 whorls of 3 tepals, Thailand. Desmos sp., flower, Bali. Desmos
chinensis, liana, fruit with lomentaceous carpels, Bali
9.7 Annonaceae: 10-54 153
Fig. 9.22 Goniothalamus laoticus, lenticels, bark with network of fibres, Thailand. Guatteria sp., corolla with three
whorls of 3 tepals, Venezuela. Polyalthia sp., the fruit is a fascicle of stipitate carpels, Madagascar
Fig. 9.23 Rollinia exsucca, young leaves conduplicate, French Guiana. Stelechocarpus burahol, trunciflory, Bali.
Unonopsis sp., the fruit is a fascicle of stipitate carpels, French Guiana
Fig. 9.24 Uvaria sp., twigs prehensile, Madagascar. Xylopia frutescens, fleshy capsules, French Guiana. Xylopia cf.
aromatica, ROUX’s model, lateral twigs develop in the axils of fallen scale-leaves, Venezuela
Fig. 9.25 Variation in TROLL’s model. Redrawn from [11]. a Tree consisting of a stacking of plagiotropic branches,
branches becoming erect after leaf fall (typical TROLL’s model, e.g. Anaxagorea acuminata). b Trunk sympodial
consisting of a stacking of erect modules, branches plagiotropic, abortions of trunk apical meristems (x), (e.g. Rollinia
spp.). c Trunk with monopodial branching, trunk apex plagiotropic but becoming erect during trunk growth (e.g.
Annona paludosa, Anaxagorea dolichocarpa, Cleistopholis patens)
9.8 Apocynaceae: 3-4-53
345 genera and ca. 4700 species. Order Gentianales. Latex white, in bark (Fig. 9.27) and leaves, s.t.
in wood (Fig. 9.26 Whrightia), latex very rarely yellow, opalescent (Pachypodium) or almost absent
(Vinca). Fleshy stems in Plumeria (Fig. 9.28f). Spines in Carissa (Fig. 9.31), or trunk thorny
(Lacmellea panamensis). Leaf arrangement opposite, s.t. whorled (Alstonia, Dyera) or spiral
(Fig. 9.31 Aspidosperma, Himatanthus, Cerbera, Plumeria), looking superficially two-ranked
(Fig. 9.28d Geissospermum). Stems lenticellate (Fig. 9.28e), very corky for some Asclepi-
adoideae. Leaves simple, entire, without glands, often discoloured (Fig. 9.33). Stipules absent.
Apical buds often protected by opposite petiolar bases (Fig. 9.29 Alstonia, Fig. 9.28-g Tabernae-
montana). Lamina usually discoloured, distinctly paler on underside. Plant most often entirely
Fig. 9.26 White latex in wood (Wrightia sp.), Fig. 9.27 Latex, interpetiolar ridge and venation not densely
Thailand reticulate, Stemmadenia donnell-smithii, Belize
9.8 Apocynaceae: 3-4-53 155
glabrous (pubescent in Wrightia pubescens). Venation not densely reticulate (Fig. 9.30), s.t. with an
intramarginal vein (Fig. 9.28f Plumeria). Flowers contortate (Fig. 9.31), with 5 petals and 5 stamens,
often scented. Fruits berries (Fig. 9.32), drupes or consisting of two almost free follicles (Fig. 9.32).
Arillate seeds (Fig. 9.33 Tabernaemontana). Winged seeds in Aspidosperma, a genus exhibiting
primitive traits.
Tall and small trees (Alstonia, Aspidosperma, Dyera), numerous shrubs (Cerbera, Tabernae-
montana, etc.), s.t. fleshy and/or spiny pachycaulous (Pachypodium, MA, AF-S). Numerous twining
lianas (Fig. 9.28e), (AM: Allamanda, Forsteronia, Landolphia, Mesechites, Prestonia; Paleo:
Cynanchum, Strophanthus). Herbs (e.g. Asclepias). Fleshy herbs and small lianas belong as a rule to
Asclepiadoideae, a group frequent in highly seasonal subtropics. In addition to tall trees harvested for
wood (Alstonia, Aspidosperma, Dyera), Apocynaceae are sources of many toxic substances used by
the pharmaceutical industry. Cardiotonic glycosides: ouabain and strophantin for treating heart dis-
eases and as a precursor of cortisone (Strophanthus).
LEEUWENBERG (dominant), (Fig. 9.28a): Alstonia,
Cerbera, Landolphia, Nerium, Plumeria, Rauvolfia.
Similar families: KORIBA (Fig. 9.28b): Alstoniamacrophylla (Malaysia),
SAPOTACEAE have bark producing abundant Cameraria latifolia (12), Tabernaemontana apoda.
white latex. However, their twigs do not bear PRÉVOST (Fig. 9.28c,d): Alstonia scholaris
lenticels. (Malaysia), Alstonia boonei (AF), Carissa macrocarpa (AF-
CLUSIACEAE–CALOPHYLLACEAE usually S), Funtumia elastica (AF), Geissospermum sp. (AM).
exude a yellow or an orange latex or resin. MASSART (Fig. 9.30): Aspidospermaspp., Lacmellea
aculeata (Guianas), Lacmellea panamensis (Panama).
CHAMPAGNAT: Allamanda cathartica (AM).
TROLL: several species of Holarrhena and Wrightia (Paleo).
Alkaloids: reserpine, rauwolfine in Rauvolfia spp., vinblastine anticancer and ajmalicine anti-
hypertensive in Catharanthus roseus.
Arrow poisons extracted from Pachypodium) or antidotes against snake venom (Strophanthus
eminii). Few species are agronomically important. Edible fruits: Carissa macrocarpa (Natal
prune), Hancornia speciosa, (Brazilian mangaba), Landolphia kirkii (‘mukanga’ in Zimbabwe).
Numerous ornamental species (Adenium, Allamanda, Plumeria or ‘frangipanier’, Thevetia), often
cultivated in “tropical” greenhouses (Asclepiadoideae: Hoya, Stapelia, etc.). There have been
unsuccessful attempts to produce rubber from the latex of several species (e.g. Funtumia
elastica).
! The subfamily Asclepiadoideae contains many leafless South African stem succulents, s.t.
threatened endemic species searched for by plant hunters [12]. The carrion flowers Stapelia
spp. are usually characterised by their foul-smelling flowers reminiscent of the odour of rotting
meat.
Fig. 9.28 Trilogy of models in Alstonia. a Leeuwenberg; b Koriba; c Prévost. d PREVOST’s model in
Geissospermum (French Guiana). e Typical: twining liana with lenticellate stems. f Fleshy stem (Plumeria). g Apical
bud embeded in petiolar bases (Tabernaemontana)
Fig. 9.29 Allamanda cathartica, flowers contortate, in cultivation, Brazil. Alstonia scholaris, apical bud embedded in
petiolar bases, Thailand. Asclepias fruticosa, spiny fruit, Botswana
Fig. 9.30 Aspidosperma megalocarpon, Belize, MASSART’s model. Venation not densely reticulate. Cameraria
latifolia, modular and sympodial branches, Belize
9.8 Apocynaceae: 3-4-53 157
Fig. 9.31 Carissa sp., twigs with bifurcate spines, Yucatan. Cerbera manghas, spiral leaf arrangement, Thailand.
Cerbera manghas, contortate corolla, Thailand
Fig. 9.32 Matelea sp., tuberculate corky fruits, Venezuela. Mesechites sp., long tubular corolla base, French Guiana.
Pentalinon andrieuxii, pairs of follicles joined at their extremities, Belize
Fig. 9.33 Rauvolfia alternifolia, fruits 2-foliculate, Flores, Indonesia. Rhabdadenia biflora, prostrate stem, leaves
discoloured, Belize. Tabernaemontana cymosa, arillate orange-coloured seeds, Venezuela
9.9 Aquifoliaceae: 18
A single genus: Ilex, but with 600 species! Order Aquifoliales. Plants without latex and not aromatic.
Spiral arrangement of leaves (Fig. 9.35). Leaves simple and entire (rarely spiny toothed). Stipules
existing but much reduced. Venation brochidodromous, not densely reticulate (Fig. 9.35). Plants
dioecious. Inflorescences lateral (Fig. 9.35). Flowers with radial symmetry and 4–9 petals. Stamens
alternate with carpels. Fruits drupes enclosing as many stones as carpels.
Similar families: Only two architectural models

CELASTRACEAE when these exhibit an MASSART: Ilex perado, Ilex cornuta.
alternate leaf arrangement. RAUH: Ilex paraguaiensis, Ilex mitis, Ilex sebertii, Ilex sp.
The European Holly (Ilex aquifolium) conforms to one or the
other models, depending of the situation.
Trees, usually small, or shrubs. Species are found predominantly in savannas or forests at low
altitudes of all land masses. Leaves of Ilex paraguayensis supply ‘mate’, a tea-like beverage con-
taining caffeine. The family is of little interest for forestry and pharmaceuticals, however Ilex vom-
itoria is used in Aerindian rituals.
! An amazing family encompassing hundreds of species in a single genus, the species resemble
each other and conform to the RAUH-MASSART continuum (Fig. 9.34). Ilex species are distributed
over both hemispheres, in America, Africa, Australasia, and reach the Hawaiian Islands. Never
change a winning team!
Fig. 9.34 Architecture in Aquifoliaceae Document available for free access on Editor’s website [13] 1. Ilex
paraguaiensis, RAUH’s model, Paraguay-Argentina. 2. Ilex perado, MASSART-RAUH’s model, reiteration (R),
Azores Islands. 3. Ilex sebertii, RAUH’s model, New Caledonia. 4. Ilex mitis, RAUH’s model, old and young tree,
Madagascar and AF-S. 5. Ilex sp., RAUH’s model, Bulolo forest, New Guinea
9.10 Araliaceae: 19-26-47-48-49-57 159
Fig. 9.35 Ilex belizensis, spiral arrangement of leaves, Cockscomb, Belize. Ilex cf. oliveriana, lateral inflorescence,
Gran Sabana, Venezuela. Ilex sp., leaf faintly crenulate and venation not densely reticulate (typical)
9.10 Araliaceae: 19-26-47-48-49-57
38 genera and more than 1600 species, of which about 600 in Schefflera. Order Apiales. Latex and
resin absent. Bark or leaves emanate a strange smell s.t. reminiscent of ivy or carrot. Stems s.t. spiny
(Fig. 9.37). Strong rhythmic growth. Architecture almost always modular and sympodial
(Fig. 9.38). Spiny stems in Aralia and Kalopanax. Spiral arrangement of leaves (but two-ranked in
the prostrate stage of ivy). Leaves usually compound palmate (Fig. 9.39), pinnate (Fig. 9.37) or
simple (Fig. 9.38), s.t. complex in their form (Fig. 9.42). Sheathing petiolar bases (Fig. 9.41),
leaving large scars after leaf fall (Fig. 9.40). Stipules absent. Inflorescences are terminal umbels
Fig. 9.36 Didymopanax morototoni, monopodial phase long Fig. 9.37 Aralia spinosa, monocaulous
lasting, then tree becoming sympodial, French Guiana spiny treelet, in cultivation
(Fig. 9.39) or panicles (Fig. 9.41). Flowers actinomorphic with 4–5, or 10 petals. Stamens variable in
numbers. Ovary superior. Fruits drupes enclosing as many stones as carpels, a berry or a schizocarp
(Harmsiopanax, AS, Papua) (Fig. 9.36).
Trees (Gastonia, AF, AS; Polyscias, Paleo; Schefflera), usually small. Didymopanax morototoni
(AM-C, N AM-S) reaches 40 m in height. Shrubs (Cussonia, AF, MA) and a few epiphytes (e.g.
Schefflera venulosa, AS). Only Hydrocotyle (cosmopolitan) and Panax (AM-N, AS-E) are
herbaceous.
Araliaceae grow predominantly in lowland or mountain forests of all land masses. Tetrapanax
papyrifer (Southern China, Taiwan) is the rice-paper tree (pith used to make cigarette paper). A few
Araliaceae are medicinal (Panax ginseng, extratropical).
Similar families: TORRICELLIACEAE (Aralidium), Malaysia, Thailand.
HOLTTUM: Harmsiopanax ingens
(New Guinea).
TOMLINSON: Aralia glabra (Japan).
CHAMBERLAIN: Meryta coriacea
(NC), Gastonia duplicata (Mayotte),
Heptapleurum calyptratum, Trevesia
glomerulata (AS).
LEEUWENBERG (Fig. 9.36):
Cussonia barteri, Polyscias fulva (AF),
Didymopanax morototoni (AM).
Fig. 9.38 Sympodial architecture, leaves simple and

tripliveined, e.g. Dendropanax sp
Fig. 9.39 Brassaia actinophylla, inflorescence a panicle of long spikes, BG Singapore. Dendropanax arboreus, L.
simple, entire or lobate, according to their location on the GU, Belize. Osmoxylon lineare, leaves trifid, cult., Bali
9.10 Araliaceae: 19-26-47-48-49-57 161
Fig. 9.40 Polyscias sp., Madagascar. Pinnate leaves and large petiolar scars. Didymopanax morototoni, leaves
compound palmate, Belize
Fig. 9.41 Schefflera quinquestylorum, inflorescence a terminal raceme of spikes, Venezuela. Didymopanax
morototoni, sheathing petiolar bases, Belize. Trevesia palmata, leaves palmatifid, Thailand
Fig. 9.42 Compound leaves. a palmate-pinnatifid; b pinnate; c palmate. Kalopanax pictus, spiny stem, in cultivation
! Araliaceae and Apiaceae (Umbelliferae) are twin families exhibiting almost the same traits except
for their distribution: Araliaceae are mainly tropical while Apiaceae are in medium and high latitudes.
It is probably only by convention that both families are not united.
! More than 400 species of Schefflera occur in tropical America, especially in the very moun-
tainous areas of the northern Andes, many of these species do not yet have scientific names and have
rarely been seen by scientists [14].
9.11 Araucariaceae: 1-42
3 genera and 32 species. Bark scaly (Fig. 9.44). Spiral arrangement of leaves (Fig. 9.43 Araucaria) or
opposite in branches (Fig. 9.44 Agathis). Internodes angular. Petiole base decurrent on the stem.
Venation parallel, often indistinct. Large female cones (Fig. 9.44), their seeds taking two or even three
years to ripen. Agathis (Kauri pines) are trees of the rainforests in Indonesia, New Guinea, Queensland, NC
and NZ. Agathis labillardieri up to 60 m in Papua, for manila copal. Araucaria have a rather subtropical
distribution (S AM-S, NC), several species of the genus are cultivated (A. araucana, A. cunninghamii).
Similar families:
MASSART (Fig. 9.43):
Podocarpaceae, but their twigs are never Agathis spp., Araucaria spp.,
arranged in a same plane. Wollemia nobilis (New South
Wales).
RAUH: Agathis dammara,
Araucaria cunninghamii.
Fig. 9.43 Araucaria angustifolia, MASSART’s model, Serra do Sudeste, Brazil. Araucaria cunninghamii, RAUH’s
model, in cultivation, Ambon, Indonesia
9.12 Arecaceae: 1-57 (Monocotyledons) 163
Fig. 9.44 Agathis damara, scaly bark, Kebun Raya, Java. Agathis sp., plagiotropic twigs with opposite leaf
arrangement, Seram, Indonesia. Wollemia nobilis, trunk with rhythmic growth and branches bearing terminal female
cones (black axis of cone visible when scales are shed), in cultivation, Montreux, Switzerland
! During the Jurassic and Cretaceous periods, Araucariaceae was distributed almost worldwide
(fossils of Araucariaceae have been found in Triassic layers of both hemispheres, [15]). In the present,
this coniferous family is restricted mainly to the Southern hemisphere and has lost ground to Pinaceae
in the Northern hemisphere.
9.12 Arecaceae: 1-57 (Monocotyledons)
184 genera and 2550 species. Order Arecales. Monocaulous trees: Areca, Cocos, Phoenix (COR-
NER’s model) or monocarpic, flowering only one time in their life, then dying following profuse
fruting: Caryota urens, Corypha utan (Fig. 9.48), Arenga pinnata (HOLTTUM). Polycaulous trees
sprouting at their base: Acoelorrhaphe (Fig. 9.45), Euterpe, Metroxylon (Fig. 9.51), Raphia, Salacca
(TOMLINSON). Aerial ramification is very rare (Hyphaene). Climbers AM: Desmoncus (Fig. 9.49);
AS: Salacca (Fig. 9.52), with prickly flagella. Trunk or leaves often spiny (Fig. 9.49). Leaves pinnate
(Fig. 9.47c), costapalmate (Fig. 9.47h), palmate Fig. 9.47k, simple bifid Fig. 9.47m or even entire.
Fig. 9.45 Acoelorrhaphe wrightii, sprouting palm, Xopol, Belize. Areca catechu, Betel nut tree, Indonesia
Fig. 9.46 Gaussia principes, spathe and paniculate inflorescence, orig. West Indies
Fig. 9.47 Arecaceae. a Trunk with long persisting leaves before shedding (e.g. Pritchardia); b induplicate palmate leaf
(leaflets like ‘V’ in section, e.g. Rhapis, reduplicate leaflets are ‘L’ in section); c pinnate and induplicate leaf (Phoenix);
d pinnate reduplicate leaf (very frequent, e.g. Hyophorbe); e: pinnate leaf with reduplicate leaflets, these inserted in a
spiral arrangement on the rachis (e.g. Syagrus); f trunk and leaf rachis spiny (e.g. Calamus); g trunk smooth because of
precocious shedding of leaves (e.g. Coccothrinax); h costapalmate leaf (e.g. Sabal); j climbing palm (rattan, e.g.
Calamus); k compound palmate leaf (e.g. Licuala); m young Geonoma sp. conserving simple leaves throughout his life
9.12 Arecaceae: 1-57 (Monocotyledons) 165
Leaf forms show drastic variation from juvenile to adult stage of the tree. Inflorescences almost
always lateral (terminal in Corypha), in spikes or panicles (Fig. 9.46), and subtended by a large bract
(Fig. 9.46). Flowers numerous, tiny 3-merous flowers (Fig. 9.50). Fruits drupes (Fig. 9.49), the
‘coconut’ being a endocarp with seed inside. Figures 9.47 and 9.53 illustrate different types of trunk
and leaf scars, just as leaf forms, which are useful for identification at the genus-level.
Fig. 9.48 Chamaedorea ernesti-augusti, leaves bifid, Shipstern, Belize. Cocos nucifera (Coconut tree, ‘Cocotier’),
sheathing foliar bases decomposing into fibres. Corypha utan, monocarpic palm, Flores, Indonesia
Fig. 9.49 Desmoncus sp., spiny climber, San Ignacio, Belize. Euterpe precatoria, drooping infructescence of drupes,
Cockscomb, Belize. Euterpe precatoria, stilt-roots, Belize
Fig. 9.50 Hyphaene petersiana, prickly petioles, Chobe, Botswana. Latania sp., spike of flowers, Sekaerat, Thailand.
Licuala peltata, Khao Chong, Thailand
Palms are well represented in the understorey as small plants (Chamaedorea, Geonoma, Licuala).
Able to grow on damp soils, palms are frequent in shallow water (many American species bear stilt-
roots: Euterpe, Iriartea, Socratea exorrhiza). This family is unique and is immediately recognizable,
but see also Cyclanthaceae.
Palms are intimately associated with human groups, especially forest dwellers. Uses are innu-
merable. Basketwork makes abundant use of rattan harvested from the spiny stems of various
Calamus spp. The most emblematic species is certainly the coconut tree (Cocos nucifera), native to
Oceania. Other stars of the family: Areca catechu (Fig. 9.45. betel nut tree, Indomalesia), Arenga
pinnata (sugar palm in Asia), Attalea cohune (‘corozo’, Central America, for thatching), Bactris
gasipaes (‘pupunha’, Amazonia, for starch, protein, oil, and carotene), Borassus aethiopum (‘rônier’,
AF-W, for palm wine or ‘arrack’); Borassus flabelliefer (‘lontar’, AS), Ceroxylon quindiuense (wax
palm, Colombia), Euterpe edulis (‘Açai’, cabbage palm, Northern South America, for palm-heart
exploitation), Metroxylon sagu (Fig. 9.51, ‘sagu’, a source of starch, in Moluccas and Papuasia),
Phoenix dactylifera (date palm, North Africa), Salacca zalacca (Fig. 9.52 ‘salak’, Indomalesia). The
oil palm Elaeis guineensis, although of African origin is nowadays intensively cultivated in Malaysia
and Indonesia.
Fig. 9.51 Metroxylon sagu, spiny foliar sheaths, Seram, Indonesia. Metroxylon sagu, agroforest, Seram. Phoenix
reclinata, savanna, Botswana
Fig. 9.52 Pinanga sp., long tubular foliar sheaths, Khao Chong, Thailand. Salacca sp., so many spines! Bali. Fruits of
Salacca zalacca in a market of Sumatra
9.13 Aristolochiaceae: 12-55 167
Fig. 9.53 ARECACEAE. a pinnate leaves with praemorse (jaggedly toothed) leaflets (e.g. Arenga); b praemorse
leaflets for Caryota; c dried up foliar bases decomposing into fibres (Cocos); d inflorescence inserted below leaf cluster
(typical for Arecaceae); e foliar bases forming a cylinder (e.g. Hyophorbe, Veitchia); f L-fissured sheaths of senescent
leaves (e.g. Sabal); g trunk with persistent foliar bases (e.g. Attalea, Scheelea); h longitudinal cut of trunk showing
sclerenchymatous inclusions in the wood (e.g. Euterpe); j slender trunk of a dwarf palm (e.g. Chamaedorea), nodes are
well spaced out; k trunk with annular expansions (e.g. Attalea, Cocos); m foliar bases forming stumps after leaf fall
(e.g. Butia); n spiny trunk and leaves (e.g. Acrocomia, Bactris); p adventitious aerial roots are adapted to swampy
places (e.g. Iriartea, Socratea)
9.13 Aristolochiaceae: 12-55
8 genera and 660 species. Order Piperales. Herbs and twining lianas (Fig. 9.54), rarely shrubs
(Aristolochia arborea, Thottea, AS). Leaf arrangement usually spiral (Aristolochia), rarely two-
ranked (Fig. 9.54. A. arborea, Thottea). Leaves simple, often cordate. Venation palmate (Fig. 9.54
Aristolochia) or tripliveined (Thottea). Perianth plastic in shape, s.t. very large (Fig. 9.54), consisting
of three sepals fused into a long tube. Fruits capsules, less often indehiscent berries or schizocarps.
Fig. 9.54 Aristolochia arborea, shrub with plagiotropic twigs and two-ranked leaf arrangement, Belize. Aristolochia
gigantea, spotted perianth, BG LC, France. Aristolochia staehli, twining liana, French Guiana
Several poisonous plants (alkaloids and aristolochic acid), e.g. Aristolochia indica. Aristolochia
species have varied floral scent, usually attracting flies, smell often like carrion or like faeces, but also
fruit-like or inconspicuous [8].
Similar families:
Aristolochia arborea can be confusedwith Architectural models:
Piper (PIPERACEAE) or members of the MANGENOT: Thottea spp. and
ANNONACEAE. Aristolochia arborea
9.14 Asteraceae: 21-23-38-40-56
A giant family consisting of ca. 1570 genera and 25,000 species. Order Asterales. Plant without latex,
s.t. aromatic, rarely spiny (Fig. 9.55 Dasyphyllum). The external wood of shrubby or arborescent
species exhibits long parallel furrows (Fig. 9.59a Baccharis, Mikania). Arrangement of leaves spiral
(Fig. 9.55), opposite (Fig. 9.56), but never two-ranked or exceptionally spiromonostichous
(Fig. 9.60). Leaves simple, lobed or pinnatifid (Fig. 9.59). s.t. with glandular trichomes (Fig. 9.59c)
or translucent dots. Flowers small, tubular, densely grouped, often surrounded by ligulate florets
(Fig. 9.58) and by bracts looking like sepals, the whole set forming a capitulum (Fig. 9.55).
Inflorescences often panicles (Fig. 9.58) or corymbs (Fig. 9.56). Fruits are almost always cypselas,
exceptionally drupelets (Wulffia Fig. 9.58). The “pappus”, a tuft of hairs inserted on the cypsela of the
Dandelion, although so familiar to “extratropical” botanists, is uncommon in tropical Asteraceae (e.g.
Figure 9.57 Pluchea).
Rainforest trees are relatively infrequent (Dasyphyllum, Piptocoma, AM; Apodocephala, MA),
some herbaceous lianas (Fig. 9.57 Mikania, mainly AM, also AS). Mostly shrubs and woody
herbs, these being very common in open areas (Bidens, Sonchus, Verbesina, Viguiera, Wedelia,
Herbs and shrubs are immediately identifiable at the

family level (flowers clustered in capitula). Lianas
(Mikania) can be confused with Clerodendron
(LAMIACEAE).
9.14 Asteraceae: 21-23-38-40-56 169
Xanthium, etc.). Montanoa quadrangularis is a fast growing pioneer tree of N AM-S which reiterates
profusely [16] (used for renaturation on waste ground), hence its name: ‘arboloco’ or mad tree! The
Andean region is especially rich in Asteraceae (Baccharis, Barnadesia, Fig. 9.60 Gongylolepis, etc.,
pachycaulous ‘frailejones’ or Espeletia spp.).
! Considering the Asteraceae of open areas of Africa, Asia and Oceania, one wonders why so
many species are native to the New World (Fig. 9.58). But this does not necessarily support the
hypothesis that the earliest Asteraceae probably migrated from the Gondwanan continent [17].
Fig. 9.55 Calostephane divaricata, winged stems, Botswana. Dasyphyllum sp., short shoots, spiral arrangement of
leaves and spiny stems, Venezuela. Flaveria linearis, opposite leaves, Botswana (AM)
Fig. 9.56 Mikania sp., herbaceous liana, leaves opposite, Belize. Mikania sp., inflorescences are corymbs, Seram,
Indonesia. Neurolaena lobata, herb, leaves extremely bitter, Cockscomb, Belize
Fig. 9.57 Pluchea carolinensis, shrub, Belize. Monosis parishii, shrub with paniculate inflorescence, ‘Dipterocarp’ dry
forest, Thailand. Viguiera dentata, (orig. AM), Indonesia
Fig. 9.58 Wulffia (syn. Tilesia) baccata, herb with baccate fruit dispersed by birds, Venezuela. Xanthium strumarium,
herb bearing spiny “fruits” (i.e. prickly involucres enclosing female flowers), Botswana
Fig. 9.59 Leaf and stem characters. a Liana Mikania, A: external wood furrowed, B: opposite, long petiolate leaves, C:
swollen petiolar base; b Sympodial architecture in Vernonia, capitula (cap); c Underside of blade with glandular
trichomes (Vernonia, Paleo); d Decurrent petiolar bases (Baccharis, AM); e Spiny stem of Dasyphyllum; f Leaf with a
hyaline margin (hy m), Stifftia, Brazil; g Scabrous leaf, Verbesina, AM, A: leaf, B: rigid hairs on the blade underside, C:
hispidous main vein; h Leaf pinnate compound or deeply pinnatifid (e.g. Dahlia)
Fig. 9.60 a. Gongylolepis sp., Gran Sabana, Venezuela. b. A curious spiromonostichous phyllotaxy in an unidentified
specimen (Baccharis sp.?), Gran Sabana, Venezuela
References 171
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1278–1313.
15. Axmith, B.J., I.H. Escapa, and P. Huber. 2008. An araucarian conifer bract-scale complex from the Lower Jurassic
of Massachusetts: Implications for estimating phylogenetic and stratigraphic congruence in the Araucariaceae.
Paleontologica-Electronica article 11 (3): 13A.
16. Meier, W. 1998. Flora und Vegetation des Avila-Nationalparks (Venezuela - Küstenkordillere) unter besonderer
Berücksichtigung der Nebelwaldstufe. Dissertationes Botanicae, Band 296. Gebrüder Borntrager, Berlin.
17. Keeley, S.C., Z.H. Forsman, and R. Chan. 2007. A phylogeny of the “evil tribe” (Vernonieae: Compositae) reveals
Old/New World long distance dispersal: Support from separate and combined congruent datasets (trnL-F, ndhF,
ITS). Molecular Phylogenetics and Evolution 44: 89–103.
Bignoniaceae to Buxaceae
10
10.1 Bignoniaceae: 21-40-43-49-51-52-56-57
82 genera and 860 species. Order Lamiales. Plant without latex or resin. Twigs often lenticellate.
Strong rhythmic growth. Climbing lianas bear modified leaves whose rachis ends in a bi-or trifidous
tendril (Fig. 10.6c), exceptionally into a trifidous hook (Dolichandra unguis-catis). Trunk sections of
lianas often exhibit a cross of four (e.g. Adenocalymma), or a higher number (Fig. 10.2), of phloem
poles. Frequent heterophylly where leaves change form along one growth unit (Fig. 10.5). Leaves
opposite (Fig. 10.4), rarely whorled, (Deplanchea, AS, AU, OC). Leaves pinnately compound
(Fig. 10.7, mainly in lianas) or palmate (Fig. 10.4 Godmania, Fig. 10.6 Tabebuia). The Calabash tree
(Crescentia cujete) being an exception with its short twigs, spiral leaf arrangement and simple leaves
(Fig. 10.3). Another peculiar shrub: Parmentiera cereifera (AM), decumbent and with trifoliolate
leaves (Fig. 10.5). Petioles often suberized in their extremities (Fig. 10.6d). Stipules absent (stip-
uliform basal leaflets in Markhamia stipulacea). Inflorescences panicles or racemes, s.t. cauliflory or
even trunciflory (Fig. 10.6a Radermachera, Fig. 10.3 Crescentia, Parmentiera). Flowers large,
showy, usually with bilateral symmetry (Fig. 10.5). the corolla tubuliform with 5 petals. Pollination
often performed by moths, bats or birds. Ovary superior. Fruits bivalved capsules, or indehiscent in
Crescentieae (Fig. 10.3). Trees (Jacaranda, Kigelia africana, Markhamia, Spathodea campanulata,
Tabebuia). Shrubs or small trees (Crescentia, Parmentiera, Tecoma). Numerous Neotropical lianas
(Adenocalymma (incl. Memora), Arrabidaea, Bignonia, Dolichandra). A few herbs, but these are
extratropical (e.g. Argylia, Incarvillea).
Predominantly in lowland forests or in savannas, mainly in tropical America. Several trees are
cultivated for the spectacular appearance of their flowers or fruits, this feature being amplified by a
deciduous habit, mauve-coloured flowers of Jacaranda, explosive yellow blooming of Handroanthus
chrysantha (Fig. 10.1), long hanging fruits in Parmentiera cereifera (to 130 cm in length), amazing
spherical fruits of Crescentia cujete. Africa also has its spectacular species: Spathodea campanulata
(tulip tree) with large red flowers, Kigelia africana (sausage tree) with long fruits and brown–red
flowers, Newbouldia laevis (‘akoko’) used for fences. Ornamental lianas (AM: Bignonia, Pyrostegia,
Parmentiera; AS: Radermachera). Several timber trees (Tabebuia spp.). Roots of Oroxylum indicum
serve to make the ‘dasamoola’ (ten roots).
https://doi.org/10.1007/978-3-031-05942-1_10
174 10 Bignoniaceae to Buxaceae
Architectural models (trees):

CORNER: Colea fusca, C. hirsuta,
Ophiocolea floribunda (MA).
CHAMBERLAIN: Oroxylum indicum
(AS).
LEEUWENBERG: Deplanchea speciosa
Similar families: (NC), Jacaranda copaia (Guianas), Kigelia
LAMIACEAE: Vitex might be confused africana (AF) Phyllarthron
with Godmania, Peronema with madagascariensis (MA).
paripinnate leaved-trees, but the latter KORIBA: Tabebuia donnel-smithii (AM-
genera do not have lenticellated twigs. C) ; Stereospermum colais (AS).
SCARRONE: Jacaranda mimosifolia
(AM), Markhamia lutea, Newbouldia
laevis (AF).
CHAMPAGNAT: Crescentia cujete
(Calabash tree, AM), Parmentiera cereifera
(Candle tree, AM), Rhigozum obovatum
(AF-S).
! This family, endowed with beautiful flowers adapted to bird pollination, exemplifies the flow-
ering strategies established by Alwyn Gentry [1], i.e. ‘steady-state’, ‘cornucopia’, ‘big-bang’ and
‘multiple-bang’ flowerers. See Handroanthus chrysantha (Fig. 10.1), a big-bang flowerer.
! Handroanthus guayacan (syn. Tabebuia guyacan), ‘ipê’ in Brazil, has a very dense hardwood
with the same fire rating as concrete and steel, meaning it resists flames much longer than softer
woods, and is so dense that it doesn’t float in water.
Fig. 10.1 Handroanthus (syn. Tabebuia) chrysantha, a Fig. 10.2 Phloem poles or lobate xylem
flowerer of the Llanos, Venezuela
10.1 Bignoniaceae: 21-40-43-49-51-52-56-57 175
Fig. 10.3 Amphilophium porphyrotricha, spike of large tubular flowers, Venezuela. Crescentia cujete, calabash tree,
shoot bearing a cluster of leaves, Venezuela
Fig. 10.4 Fernandoa adenophylla, large petiolar scars, Thailand. Godmania aesculifolia, shrub with compound
palmate leaves, Venezuela. Jacaranda obtusifolia, deciduous tree with mauve-coloured flowers, Venezuela
Fig. 10.5 Markhamia sp., tree with pinnate compound leaves, Thailand. Markhamia stipulata, flowers adapted to bat
pollination, Thailand. Parmentiera cereifera, twigs lenticellate, Kebun Raya, Java
Fig. 10.6 a Radermachera ignea, BG Sirikit, Thailand. b Tabebuia sp., compound palmate leaves (cult., orig. AM),
Bali. c in lianas, the foliar apex is typically modified into a tendril. d petiolules suberized at both ends (an unidentified
specimen, Venezuela). e: heterophyllous liana, leaves bifoliolate to paripinnate, leaves with or without tendrils
10.2 Bixaceae: 2-15-47
4 genera and 21 species. Order Malvales. Bark fibrous (Fig. 10.8), s.t. exudating a pale orange gum.
Spiral arrangement of leaves. Leaves simple, entire (Fig. 10.8 Bixa), lobate (Fig. 10.9
Cochlospermum spp.) or compound palmate (Cochlospermum spp. Fig. 10.9). Petioles somewhat
long, s.t. distally pulvinate (Bixa). Indumentum of orange scaly hairs in Bixa (Fig. 10.8). Flowers
attractive, actinomorphic, with 5 petals and numerous stamens. Fruits capsules, spiny in Bixa
orellana and enclosing seeds with an orange-coloured testa (Fig. 10.7). Seeds hairy (kapok of
Cochlospermum). Small trees (Cochlospermum), shrubs (Bixa, Cochlospermum) or herbs
(Amoreuxia, AM-C).
Similar families: KORIBA (Fig. 10.8): Cochlospermum
vitifolium, ‘Bototo’, Venezuela.
MALVACEAE have a fibrous bark, showy
flowers, and petioles usually pulvinate. SCARRONE: Bixa orellana (AM),
Cochlospermum religiosum (AS).
10.2 Bixaceae: 2-15-47 177
Fig. 10.7 Bixa orellana, spiny capsules and arillate seeds
Fig. 10.8 Internal bark with a network of fibres. Cochlospermum vitifolium, KORIBA’s model, Venezuela. Bixa
orellana, scaly orange-coloured hairs on the underside of the lamina
Fig. 10.9 a, b Cochlospermum vitifolium, Belize. a Leaves compound palmate. b Bark fibrous, chlorophyllous layer
between rhytidome and inner bark. c Cochlospermum religiosum, Kebun Raya, Java
A pantropical family, mainly in savannas. Several species of Cochlospermum ornamental, but also
sources of gum or dyes and medicines (C. tinctorium AF-W). Seeds of Bixa orellana (‘annato’,
‘achiote’, ‘onoto’, ‘urucú’) provide an orange-coloured pigment which, mixed with carapa oil, was
used by native Amerindians to protect themselves from mosquito bites and also as body painting.
Nowadays, Bixa seeds are used to dye silk yellow or as food colouring agent. Cochlospermum
religiosum is often planted in India near temples for its intense yellow flowers [2].
! Bixaceae are close relatives of Cistaceae, a Mediterranean family. Both groups have leaves covered
by fasciculate or stellate hairs, attractive flowers with numerous stamens and fruits like capsules.
10.3 Boraginaceae (incl. Cordiaceae, Ehretiaceae): 20-21-24-25-30
121 genera and 2600 species. Order Boraginales. Latex or coloured sap absent. Inner bark s.t. with a
network of fibres or turning brown when cut (Fig. 10.11). Several species have short shoots, these s.t.
modified into thorns (Fig. 10.14 Rochefortia). Myrmecophily in Cordia nodosa (Fig. 10.12). Spiral
arrangement of leaves (Fig. 10.13). Branches usually sympodial (Fig. 10.11 e.g. PRÉVOST’s
model). Leaves simple, entire or rarely toothed (Fig. 10.14 Wigandia), compound only in extrat-
ropical herbs (Hydrophyllum, Phacelia). Numerous extratropical herbs (Echium, Myosotis, etc.).
Leaves usually pubescent, s.t. rough, on both faces (Fig. 10.12), when leaves are ‘glabrous’ pub-
escent buds have to be observed.
Fig. 10.10 Cordia sebestana. Belize

10.3 Boraginaceae (incl. Cordiaceae, Ehretiaceae) … 179

Species of SOLANACEAE (Lycium) and SALICACEAE (Xylosma) LEEUWENBERG (Fig. 10.13):
have also decumbent spiny branches with short shoots. Heliotropium argenteum (OC),
Cordia lutea (Galapagos), Cordia
africana (Ethiopia).
KORIBA: Cordia sebestena, WI,
Inflorescences terminal cymes (Fig. 10.13), s.t. con- naturalized in all tropical coasts.
densed heads (Fig. 10.14). Flowers with radial symmetry, PRÉVOST (Fig. 10.11): Cordia
5 petals and 5 stamens. Style gynobasic. Fruits drupes nodosa, C. collococa, Cordia
(Fig. 10.13 Cordioideae, Ehretioideae), s.t. samaras elliptica (AM).
(Fig. 10.11 Bourreria) or schizocarps of four drupelets FAGERLIND: Cordia alliodora
(Heliotropioideae). (AM).
Trees (Cordia (incl. Patagonula), Ehretia), s.t. deciduous SCARRONE: Patagonula
(Cordia sebestena). Shrubs (Cordia, Heliotropium, Wigan- americana (S AM-S).
dia) s.t. spiny Rochefortia (AM-C), Rochefortia represents a CHAMPAGNAT: Cordia
good example of the frequent combination of characters in curassavica (WI), C. cochinchinensis
(AS).
dry environments, i.e. «decumbent branches—short shoots—
TROLL’s model probably absent in
spines». Woody lianas (Cordia bantamensis). Herbs
Boraginaceae.
(Heliotropium indicum).
Boraginaceae is architecturally and biologically highly
diversified, and occurs in all environments, from the tropics to high latitudes. Fine timbers for
carpentry (AM): Cordia alliodora (‘Pardillo’), C. goeldiana (‘Frei-jorge’), Cordia americana
(‘guajivira’). Fruit trees: Cordia myxa (‘lasura’ in India), along the Hymalayan tract, cultivated in SE-
Asia. Numerous ornamental species (Fig. 10.10 Cordia sebestana, Echium spp.). Light tolerant trees
for afforestation of degraded areas: Cordia alliodora, Cordia americana.
! Inflorescence architecture is exclusively cymose in Boraginaceae [3]. The family provides one of
the best examples of a group ranging from the equator to alpine biomes (Eritrichium hemisphaericum
up to 5700 m a.s.l. in the Himalayas).
Fig. 10.11 Bourreria cumanensis, samaras, Venezuela. Cordia sp., bark section quickly turning brownish. Cordia sp.,
PRÉVOST’s model, Venezuela
Fig. 10.12 Cordia nodosa, hollow stem inhabited by ants, Venezuela. Sympodial branch, Cordia cf. sulcata,
Guadeloupe. Cordia sp., leaves pubescent on both faces, Venezuela
Fig. 10.13 Cordia sp., terminal inflorescence, Belize. Ehretia microphylla, leaves inserted on short twigs, fruits
drupaceous, Bali. Heliotropium sp., small shrub conforming to LEEUWENBERG’s model, Yucatan
Fig. 10.14 Rochefortia sp., branch with twigs modified into spines and bearing drupes, Belize. Tournefortia
cuspidata, terminal inflorescences in heads, Venezuela. Wigandia caracasana, serrulate leaves, sunken venation, BG
Geneva
10.4 Burseraceae: 3-46-C 181
10.4 Burseraceae: 3-46-C
19 genera and 675 species. Order Sapindales. Bark aromatic, source of translucent resin (Fig. 10.19)
or resinous gums. Trunk s.t. buttressed (Fig. 10.15) or with stilt-roots (Fig. 10.17), e.g. Canarium.
Trunk and branches s.t. thorny in African savannas (Fig. 10.17). Several species deciduous
(Fig. 10.16 Bursera simarouba). Strong rhythmic growth (Fig. 10.16). Spiral arrangement of leaves.
Leaves compound pinnate (Fig. 10.16), exceptionally simple (Fig. 10.18 Protium unifoliolatum).
Stipules absent, rarely present (Canarium spp., Indomalesia). Basal leaflets in stipular position
(Garuga spp.). Leaflets entire, s.t. toothed (Boswellia serrata, Bursera tomentosa, Crepidospermum
rhoifolium, Garuga spp., Protium africanum). Petiolules distally enlarged or pulvinate (Fig. 10.19
Canarium, Protium), s.t. only the terminal petiolule is pulvinate. Base of petiol(ul)e pulvinate and
suberized (Fig. 10.20 Tetragastris, Fig. 10.19 Protium). Venation brochidodromous (Fig. 10.19).
Indument of simple hairs, rarely stellate hairs (Dacryodes). Plant often dioecious. Inflorescences
lateral (Fig. 10.16). Flowers small with petals fused at their base, surrounding a glandular disk.
Ovary superior. Fruits drupes (Fig. 10.16) enclosing 1–5 stones or only one stone enclosing several
seeds. Protium spp. might have capsules, these red coloured and with seeds covered by a white aril
(Fig. 10.18), (zoochory) [4].

ANACARDIACEAE with imparipinnate leaves and resinous RAUH probably frequent, even dominant
exudate. Anacardiaceae and Burseraceae are two closely (e.g. Bursera simarouba).
linked families of the order Sapindales.
Fig. 10.15 Canarium sp., buttressed tree, Seram

Trees (AM: Bursera, Dacryodes, Protium, Tetragastris, Trattinnickia; AF: Aucoumea; Paleo:
Canarium, Santiria). Shrubs (AF: Boswelllia; AM-AF-MA: Commiphora), s.t. spiny.
Usually in rainforests, scarce in secondary forests but several genera are common in African
savannas (Boswellia, Commiphora). Timber: AF: Aucoumea klaineana (‘okoumé’), AS: Santiria
apiculata (‘kedondong’). Varnishes: Canarium strictum (black damar). Body care: Canarium
luzonicum (‘Manila elemi’). Fruit trees: Dacryodes edulis (African plum, ‘safoutier’), oil seeds:
Canarium commune (‘kenari’, Indomalayan). Myrrh resin extracted from Commiphora spp. has been
used throughout history as perfume, incense, medicine and, before the term was coined, “chewing
gum”.
! Several species growing in savannas have a photosynthetic bark. Having fewer bark layers during
the dry season (leaf-off condition) is an advantage for the plants as then their inner photosynthetic
bark is more exposed to light, enabling them to trap light energy [5].
Fig. 10.16 Bursera simarouba (all three illustrations). Deciduous twigs and fruits, Venezuela. Peeling bark, Shipstern,
Belize. Rhythmic growth, leaves imparipinnate, drupes, Venezuela
Fig. 10.17 Canarium sp., stilt roots, Seram, Indonesia. Commiphora pyracanthoides, spiny small tree, Botswana.
Dacryodes edulis, leaves compound pinnate, Cameroon
10.4 Burseraceae: 3-46-C 183
Fig. 10.18 Dacryodes edulis, flushes of young leaves, Campo forest, Cameroon. Typical pulvinate petiolules (p) of
several genera; serrulate leaflets and winged rachis; peeling bark (pb) in Bursera. Protium unifoliolatum, capsular fruits
with red pericarp, seeds covered by white arils, Venezuela
Fig. 10.19 Protium copal, petiolules pulvinate at both ends, Belize. Protium sp., resinous exudate in bark, French
Guiana. Tetragastris sp., petiolar bases pulvinate and suberized, Venezuela
Fig. 10.20 Tetragastris sp., petiolules suberized at the base, but not distally enlarged, Venezuela. Protium sp.,
suberized petiolar bases, Venezuela. Unidentified buttressed tree, Venezuela
10.5 Buxaceae: 39-44
A small family encompassing 7 genera and 115 species. Order Buxales. Internodes distinctly
angular. Opposite (Buxus) or spiral (Styloceras) leaf arrangement. Stipules absent. Leaves glabrous,
coriaceous and shiny (Fig. 10.21). Venation somewhat indistinct with two lateral ascending sec-
ondary veins. Flowers small in heads or spikes. Fruits dehiscent capsules or drupes.
Buxus (30 species) has a curious discontinuous distribution: EU, AF-S, MA, AM-C, with a center
of radiation in Cuba. Edible fruits, in Peruvian Amazonia: Styloceras laurifolia (‘naranjillo’).
Similar families:
MYRTACEAE, but bark ochraceous and internodes usually not angular.
CELASTRACEAE, however these have minute stipules.
Fig. 10.21 Buxus bartlettii, shiny coriaceous leaves, Belize
References
1. Gentry, A.H. 1972. Flowering phenology and diversity in tropical Bignoniaceae. Biotropica 6: 64–68.
2. Burkill, I.H. 1935. A Dictionary of the Economic Products of the Malay Peninsula, Vol. I and II, 2402 pp. Oxford:
University Press.
3. Buys, M.H., and H.H. Hilger. 2003. Boraginaceae cymes are exclusively scorpioid and not helicoid. Taxon 52:
719–724.
4. Nair, K.K.N. 2000. Non-wood Forest Products of Kerala, 450 pp. Peechi: Kerala Forest Research Institute.
5. Girma, A., A.K. Skidmore, C.A.J.M. Bie, F. de Bongers, and M. Schlerf. 2013. Photosynthetic bark: Use of
chlorophyll absorption continuum index to estimate Boswellia papyrifera bark chlorophyll content. International
Journal of applied Earth Observation and Geoinformation 23: 71–80.
Cactaceae to Cycadaceae
11
11.1 Cactaceae: 21
126 genera and 1500 species. Order Caryophyllales. Lacking leaves, Cactaceae look very peculiar,
even when tall, one hesitates to considerate them as trees, e.g. (Fig. 11.1), Cereus hexagonus
(Venezuela). However Pereskia species bear true leaves (Fig. 11.3) and are readily recognizable as
shrubs or small trees. Some species are climbing (Fig. 11.3). Stems spiny, except epiphytic Rhip-
salis. Cactus spines are produced from specialized structures called areoles, a kind of highly reduced
branch. Areoles are an identifying feature of cacti.
Fig. 11.1 Cereus hexagonus,
Venezuela
https://doi.org/10.1007/978-3-031-05942-1_11
186 11 Cactaceae to Cycadaceae
Fig. 11.2 Hylocereus undatus, (in cultivation, orig. AM), Indonesia. fleshy stems, horned fruits. Opuntia sp., paddle
spiny stems, Tulum, Mexico
Fig. 11.3 Pereskia grandiflora, BG FBV, Venezuela. Pereskia aculeata, in cultivation, Santa Catarina, Brazil.
Selenicereus testudo, climbing cactus, Xopol, Belize
Almost all Cactaceae are of New World origin and live in dry forests, savannas or “chaparral”, the
only non American species is Rhipsalis baccifera in AF-S, Madagascar and Sri Lanka. Rainforests do
not usually shelter “cactus trees”, however species of Hylocereus, Selenicereus (Fig. 11.3) and
Rhipsalis can be found in dry deciduous forests. Several edible fruits: Hylocereus undatus (Fig. 11.2
‘pitaya’ or dragon fruit), Opuntia spp. (Fig. 11.2 prickly pear, ‘nopal’).
! Before eating the fruits of Opuntia (‘tunas’ for Spanish-speakers), roll them in a damp cloth
which will remove most of the spines.
11.2 Canellaceae (C) + Winteraceae (W): 13-23-29-30
Two families encompassing 10 genera and 92 species. Order Canellales. Bark often aromatic or
tasting spicy (C: Cinnamodendron, Warburgia). Wood lacking vessels (conducting elements are
tracheids). Twigs lenticellate. Spiral arrangement of leaves (C: Canella, Cinnamodendron sect.
Cinnamodendron, Warburgia; W: Drimys, Fig. 11.4e or two-ranked arrangement (C: Cinnamod-
endron sect. Capsicodendron, W: or Fig. 11.4-a Cinnamosma, MA). Leaves simple, entire, s.t. with
translucent dots (C) or waxy underside of laminas (W: Drimys). Inflorescences lateral (C: Cin-
namodendron, Cinnamosma; W: Fig. 11.4d Drimys) or terminal (C: Canella), in racemes or panicles.
Flowers s.t. solitary in leaf axils, with 3 sepals and petals in varied numbers. Stamens fused into a
tube. Parietal placentation (C). Fruits capsules, berries (Fig. 11.4-b Canella) or heads of follicles
(Fig. 11.4c Drimys).
11.3 Capparaceae (incl. Cleomaceae) … 187
Fig. 11.4 a Cinnamosma macrocarpa, Montagne d’Ambre, Madagascar. b Canella winterana, sectioned fruits showing
black seeds, Guadeloupe. c–e Drimys brasiliensis, Santa Catarina, Brazil. c: lateral infructescences with spirally arranged
carpels (cp); d inflorescences in the axils of scale-leaves. e RAUH’s model, monopodial and rhythmic branching
Similar families: SCARRONE: Canella (West Indies),
Venation also indistinct in Takhtajania (MA).
ERYTHROPALACAE and RAUH (Fig. 11.4e ): Drimys (AM-S,
PENTAPHYLACACEAE (Ternstroemia). Malaysia, OC).
Aromatic bark in a few LAURACEAE.
MANGENOT: Cinnamosma (MA), probably
Cinnamodendron sect. Capsicodendron (WI).
TROLL: probably absent.
Trees or shrubs. Few uses. Medicine: Warburgia salutaris (AF-E). Spices: Cinnamodendron
corticosum (West Indies). Essential oils (saro oil): Cinnamosma fragrans (MA).
Both families are mainly distributed in the Southern hemisphere, their Gondwanan origin is
suggested by [1].
! A lianescent habit is reported for Tasmannia cordata (New Guinea) [2], however this species is
not a true liana because it does not use an active mechanism for climbing, it merely gets caught on its
support with the bases of its cordate leaves.
11.3 Capparaceae (incl. Cleomaceae): 8-16-21-22-27-29-30-47
32 genera and more than 750 species. Order Brassicales. Latex absent. Bark not aromatic, but
secreting mustard oil glucosides. Short shoots bearing leaf clusters (Boscia, Maerua). When leaf
arrangement is spiral, petioles are often very variable in length (Buchholzia, Capparis and Morisonia
exhibit rhythmic growth), two-ranked leaf arrangement (Fig. 11.9) in Capparis spp. Leaves stip-
ulate (stipules s.t. vestigial or absent) or spines in stipular position (Fig. 11.6a). Leaves simple
(Boscia, Capparis, Morisonia), compound trifoliolate (Fig. 11.6b Crateva, Maerua, Ritchiea) or
compound palmate (Cleome), lamina margin entire. When branches are erect, petioles are usually
distally swollen into a pulvinus (Fig. 11.8), (Buchholzia, Capparis spp., Morisonia). Main vein
ending in a mucro and/or an indentation (Fig. 11.9c). Indument of simple, stellate or peltate hairs
(Fig. 11.6c–e). Receptacles of flowers usually extended in an (andro)-gynophore. Flowers with
2–4–6 free petals and numerous stamens (Fig. 11.9). Ovary superior. Fruits fleshy or coriaceous
berries, these often stipitate (Fig. 11.8).
Similar families: CHAMBERLAIN: Euadenia eminens (AF).
LEEUWENBERG: Cleome anomala (AM).
Morisonia can be confused with several
EUPHORBIACEAE. Capparis, when RAUH: Capparis pittieri (AM).
bearing plagiotropic branches, can be CHAMPAGNAT: Boscia albitrunca, Capparis sp.,
confused with PHYLLANTHACEAE or Maerua sp. (AF).
SALICACEAE. TROLL (Fig. 11.8 ): Capparis cynophallophora
(West Indies).
ACOSTA: Cleome hirta (AF-E,S).
Small trees (Capparis, Crateva, Steriphoma), shrubs (Capparis, Euadenia), s.t. climbing (Capparis,
Maerua, Ritchiea) or herbs (Fig. 11.9) in Cleomaceae (Cleome, Podandrogyne).
A pantropical family, with most species occurring in African savannas (Fig. 11.5 Boscia angus-
tifolia, Buchholzia, Capparis, Maerua, Ritchiea). Few species are of economical importance
Fig. 11.5 Boscia angustifolia, decumbent branches, Botswana

11.3 Capparaceae (incl. Cleomaceae) … 189
Fig. 11.6 a Stipular spines (Capparis spp.). b Trifoliolate leaf (Crateva sp.). c–e Simple, stellate, and peltate hairs
(Capparis spinosa or Caper bush, is extratropical). Crateva religiosa is, in addition to its medicinal
properties, a symbolic plant in the Hindu religion.
! Boscia senegalensis, a shrub which may be eaten by cattle when grass is rare, has many names in
different regions of of the African savannas, ‘houassa’, ‘ndianda’, ‘gigilé’, etc.
! Extracts of Capparis decidua have strong anti-termite activity and are used in India for wood
protection in the soil against the Indian white termite [3].
! In southern Africa termites seem to support the presence of Capparis tomentosa (Fig. 11.7) while
preventing the establishment of other plant species.
Fig. 11.7 Boscia angustifolia, leaves clustered on short shoots, Botswana. Capparis tomentosa in symbiosis? with a
termite mound, Botswana
Fig. 11.8 Capparis cynophallophora, TROLL’s model, Guadeloupe. Capparis flexuosa, stipitate fruits, Guadeloupe.
Capparis frondosa, petioles distally swollen, stipitate fruits, Venezuela
Fig. 11.9 a Capparis pittieri, main vein ending in a mucro, Belize. b Capparis sp., two-ranked leaf arrangement, four
petals and numerous stamens, Venezuela. c Capparis sp., leaves two-ranked, spines in stipular position, Venezuela.
d Cleome hirta, ruderal herb, Botswana
11.4 Caricaceae: 3-20-49 191
11.4 Caricaceae: 3-20-49
A small, essentially New World, family of 6 genera and 34 species. Order Brassicales.
Small trees (Fig. 11.10 Carica, Vasconcellea), not ramified or sparsely branched (Jarilla a rhi-
zomatous herb). Dioecious (Fig. 11.11a, b Carica) or monoecious (Cylicomorpha, AF). Xylopodium
in Jacaratia corumbensis (Fig. 11.11e). Trunk spiny (Jacaratia pp., Cylicomorpha). Wood soft.
Latex whitish or opalescent. Spiral leaf arrangement. Leaves large, simple but deeply incised
(Fig. 11.11c Carica, Vasconcellea) or compound palmate (Fig. 11.11d Jacaratia). Flowers with
parts in five, solitary, axillary or in cymes. Fruits berries (Fig. 11.10).
Fig. 11.10 Fruiting Papaya tree
Carica papaya (AM), the papaya tree, is a small pachycaulous dioecious fruit tree cultivated
everywhere in the tropics. The wood of Jacaratia spinosa (wild mango) is soaked in honey or syrup
and served as a delicacy. Protease enzymes of Papaya fruits have the ability to tenderize meat. Leaves
for shrinkproof finishing method for wool.
Similar family: None! Architectural models: CORNER: Carica spp. RAUH: Vasconcellea spp. (AM)
The two species Cylicomorpha are the only African representatives of the family, they grow in
montane forests. Cylicomorpha is thought to be ancestral in Caricaceae.
Fig. 11.11 a–c Carica papaya. a Female flowers (f), b male flowers (m), (lateral flowering); c leaves palmatilobate
with pinnatifid divisions; d: Jacaratia spinosa, compound palmate leaves and berries; e: Jacaratia corumbensis,
xylopodium (orig. Bolivia, Brazil)
11.5 Casuarinaceae: 37
A small family of 4 genera and 95 species, all Australasian and Oceanian. Order Fagales. A strange
group placed in past times next to horsetails! But furrowed external wood (Fig. 11.13) reveals affinity
with oaks and chestnuts (Fagaceae). Roots bearing nitrogen-fixing bacterian nodules. Whorled leaf
arrangement. Flowers much reduced, the male ones with 1 stamen and the female ones with 2 carpels.
Leaves like small scales (Fig. 11.13). Infructescences are cone-like groups of drupeoles surrounded
by woody bracts (Fig. 11.13).
Similar family: Architectural models:

Tamaricaceae (Tamarix): slender twigs bearing scale- ATTIMS-RAUH (Fig. 11.12): exclusive.
leaves, extratropical.
Casuarina equisetifolia trees (filao or Australian pine) are often seen on seashores (Fig. 11.12),
but they are forest trees too. Uses: dune stabilizers, windbreakers, soil fertilizers, very calorific wood.
! Casuarina oligodon, fixing atmospheric nitrogen, have been used by highland dwellers of Papua
New Guinea in agroforestry for thousands of years [4], a demonstration of clear-sighted empirical
research.
11.5 Casuarinaceae: 37 193
Fig. 11.12 Casuarina equisetifolia, back-shore, Southern Brazil
Fig. 11.13 Casuarina equistifolia, Santa Catarina, Brazil. External wood furrowed, suberized twigs bearing scaly
chlorophyllous leaves and drooping twigs bearing fruits
11.6 Celastraceae: 4-8-13-16-18-21-30-33-34-45-53-54-56
96 genera and 1300 species. Order Celastrales. Plants without latex, s.t. red exudate in bark (Prionostemma,
AM). Bark not aromatic. Prehensile twigs (Fig. 11.15) in many lianas. Branches thorny (Fig. 11.18
Gymnosporia, African dry tropics and subtropics). Thin orange-coloured layer (Fig. 11.17) between dead
bark and inner bark (Lophopetalum beccarianum). Inner bark often pink coloured (Fig. 11.17). Climbing
species are twining (Celastrus) or have prehensile twigs (Fig. 11.15) (Hippocrateoideae and Salacioideae).
Xylem of liana in blocks or lobate (Fig. 11.16). Leaves opposite (Fig. 11.19) (Euonymus, Hippocratea,
Fig. 11.14 Salacia sp.,

flowering liana, Cameroon
Fig. 11.15 Prehensile twigs
Fig. 11.16 Xylem in blocks

(a) or lobate (b)
11.6 Celastraceae: 4-8-13-16-18-21-30-33-34-45-53-54-56 195
Lophopetalum, Rhacoma, Salacia, Zinowiewia), spiral (Fig. 11.20) (Bhesa, Celastrus, Gymnosporia)
or two-ranked (Maytenus, AM; Fig. 11.20 Siphonodon, AS). Stipules much reduced (absent in
Gymnosporia and Siphonodon). Torn leaves of several species show laticiferous threads. Leaves simple,
their margins entire, rarely weakly toothed (Catha edulis, Maytenus, AM). Petiole distally pulvinate in
Bhesa (Fig. 11.20). Venation not densely reticulate, s.t. indistinct for coriaceous leaves in savannas. Stems
and leaves glabrous (Fig. 11.18). Flowers small, 3–5 petals (Fig. 11.14, here a cauliflorous Salacia),
stamens alternate with carpels, disks various, often present and conspicuous. Ovary half inferior. Fruits
drupes (Fig. 11.19), (Crossopetalum, Salacia), samaras (Plenckia) or capsules (Euonymus).
Similar families:
MASSART: Euonymus spp.; Maytenus spp.
ICACINACEAE: but their apices often pubescent.
RAUH: Crossopetalum rhacoma (AM-C).
AQUIFOLIACEAE: also glabrous leaves, but these ROUX: Gymnosporia sp., (AF), Salacia (Pantrop).
usually faintly crenulate.
TROLL? Bhesa cf. paniculata (AS).
Fig. 11.17 Inner bark pink-coloured, distinctly delimited from external wood (Gymnosporia senegalenssis),
Botswana. Orange-coloured phelloderm and pink inner bark of a unidentified Celastraceae, Madagascar. Pink-coloured
inner bark (Maytenus elliptica), Guadeloupe
Fig. 11.18 Catha edulis, leaves thinly toothed, cult. Cheiloclinium cognatum, Salacioideae, stem and leaves glabrous,
Venezuela. Gymnosporia senegalenssis, decumbent branches bearing thorns, Botswana
Fig. 11.19 Rhacoma crossopetalum, shrub with red drupes, Belize. Salacia sp., drupaceous fruits, Venezuela.
Salacioideae, cf. Tontelea hondurensis, precocious suberization of a twig, venation not thinly reticulate, Belize
Fig. 11.20 a Siphonodon celastrineus, leaves alternate, plagiotropic branch, Thailand. b Bhesa cf. paniculata, spiral
leaf arrangement, leaves with long, pulvinate petioles, Sumatra (author’s field book).
Trees (AS: Bhesa, Euonymus, Kokoona, Lophopetalum; AM: Maytenus, Zinowieva). Lianas, s.t.
with very thick trunks (AM: Cheiloclinium, Hippocratea, Prionostemma, Pantrop: Salacia). Shrubs
(Crossopetalum, Gymnosporia, Mystroxylon aethiopicum), s.t. spiny (Gymnosporia, AF). Herbs
(Stackhousia, AU). Celastraceae grow in all tropical rainforests (Salacia, 200 spp.) and savannas
(Crossopetalum, Gymnosporia). Few species are economically important, even though Catha edulis,
‘khat’, is used (too often) as a stimulant in East Africa and Madagascar. Arrow-poisons: Lophope-
talum spp. (Moluccas, Philippine).
! Bark of Lophopetalum beccariana serves forest dwellers of Sumatra to light fires, even with
damp wood.
! Circumscription of Celastraceae is still subject debate: Bhesa (Fig. 11.20) is included in this
family but is currently treated in the Centroplacaceae, Malpighiales. Siphonodon (Fig. 11.20) might
be better placed in a distinct family.
11.7 Chloranthaceae: 32
4 genera and 67 species. Order Chloranthales. Aromatic shrubs and herbs. Base of internodes swollen
(Fig. 11.21). Opposite leaf arrangement. Leaves simple and toothed (Fig. 11.21). Petioles connate
basally. Stipules interpetiolar or annular (Fig. 11.21). Flowers minute, grouped in dense
11.7 Chloranthaceae: 32 197
Fig. 11.21 Sarcandra sp.,

base of internodes swollen,
stipules annular.
Hedyosmum sp., toothed
opposite leaves
Fig. 11.22 a Ascarina philippinensis, leaves almost entire, New Guinea, herbarium specimen, CBG H. b Chloranthus
japonicus, herb conforming to TOMLINSON’s model, typical ‘ranunculoid’ venation. c, d Hedyosmum sp., STONE’s
model; c in French Guiana, d diagramatic representation of the model
inflorescences (Fig. 11.22). Stamens variable in numbers. Ovary superior or half inferior, carpels
uniovulate. Fruits berries or drupes.
Shrubs or small trees Ascarina (MA, AS, OC), Hedyosmum (AM), Sarcandra (AS). Herbs
Hedyosmum (AM, AS).
Chloranthaceae grow in the understorey of dense rainforests. Few uses are known, leaves of
Chloranthus served in Java as a ‘tea’ before the cultivation of Camellia sinensis.

RUBIACEAE, but not aromatic and leaves TOMLINSON (Fig. 11.2 2): Chloranthus (AS).
entire. LEEUWENBERG: Chloranthus, Sarcandra (AS)
MONIMIACEAE, but much more woody. STONE (Fig. 11.22 ): Ascarina (MA, AS, OC),
Hedyosmum (AM).
Leaves of the Chloranthaceae have a peculiar venation (broken arcs, Fig. 11.22) which is also
found in Nymphaea, Monimiaceae and Ranunculaceae, all these taxa being representatives of earlier
Angiosperms. Hedyosmum is the only American genus of the family (44 American species and a
single Asiatic one). Fossil pollens have been recorded in Lower Cretaceous layers (as early as 112
million years ago) [5].
11.8 Chrysobalanaceae: 2-5-7-8-13
20 genera and 525 species. Order Malpighiales. Several beautiful trees exhibiting crown shyness
(Fig. 11.23). Individuals without latex and not aromatic, s.t. red aqueous exudate (Fig. 11.28). Bark
very hard due to the inclusion of silica bodies. Inner bark often pink- or red-coloured (Figs. 11.27,
11.28). Trunk or branches distinctly lenticellate (Fig. 11.27). Sympodial architecture with apex
abortions (Fig. 11.24) or terminal flowering (Fig. 11.27). Rhythmic growth with series of scale-
leaves. Leaves two-ranked (Fig. 11.25) but a few species have a spiral leaf arrangement in all
branches (Fig. 11.24 Chrysobalanus icaco). Stipules present, entire or fringed and glandular
(Fig. 11.24 Hirtella). Venation usually camptodromous (Fig. 11.25 Couepia, Licania). Stems and
leaves with a tomentum of short hairs, s.t. long spread hairs (Hirtella) or underside of laminas whitish
(Fig. 11.27 Licania), or rust-coloured (Fig. 11.26 Couepia). Base of laminas often glandular
(Fig. 11.24). Inflorescence terminal, in racemes (Fig. 11.27) or panicles (Fig. 11.28). Flowers small,
5 free petals and numerous stamens. Carpels often moved away from the center in relation to the
receptacle. Ovary 2–3 carpellate. Fruits 1-stone drupes.
Fig. 11.23 Parinari sp., crown shyness, Bochinche, Venezuela
Fig. 11.24 Chrysobalanaceae. a: abortion (+) of an apical meristem; b: intrapetiolar stipules (Licania spp.); c stipules
glandular (Hirtella spp.); d monopodial branching, leaves glandular, a-b prophylls (Chrysobalanus icaco); e–f pairs of
glands (gl) on junction of lamina with petiole (e: Parinari, f: Licania)
11.8 Chrysobalanaceae: 2-5-7-8-13 199
Fig. 11.25 A unidentified sapling with two ranked leaf arrangement, venation camptodromous, French Guiana
Fig. 11.26 Couepia sp., two-ranked leaf arrangement, rust-coloured indument, French Guiana. Chrysobalanus icaco,
shrub on seashore, Yucatan. Chrysobalanus icaco, flowers, Tulum, Mexico
Fig. 11.27 Hirtella sp., terminal inflorescence, Venezuela. Licania hypoleuca, bark lenticellate, inner bark pink-
coloured, Belize. Licania hypoleuca, leaves discoloured, Belize
Fig. 11.28 Licania sp., inner bark producing a red aqueous exudate, Venezuela. Licania sp., terminal panicles,
Venezuela. Parinari cf. rodolphii, strong rhythmic growth revealed by scaly leaves, young leaf-folding revolute

DICHAPETALACEAE also have TROLL (strongly dominant): Couepia, Licania, Parinari.
hairy, stipulate, and glandular leaves. SCARRONE? : Chrysobalanus icaco.
Trees (Couepia, Licania, AM; Parinari, AM-AF; Maranthes, mainly AF). Shrubs Chrysobalanus
(Pantrop), s.t. decumbent (Dactyladenia, AF). Geoxylic (Parinari capensis AF-C), Hirtella (mainly
AM). No lianas and no herbs.
Almost all native to America, exceptions are Parinari (Pantrop), Atuna (AS, OC). Chrysobalanus
icaco (Icaco coco plum) grows on both sides of Atlantic coastal regions, but is naturalized on many
shores of Asia and Oceania. Very frequent in Northern AM-S (on siliceous sands following erosion of
the Guiana Shield). Bad for carpentry because of a high content of silica, a property that sawyers do
not appreciate (consider the vernacular names of several species: Devil’s wood, ‘Bois de fer’,
‘Hierrillo’, ‘Mancharo caspi’). Fruit trees (Parinari spp. Guinea Plum tree, AF-C).
11.9 Clethraceae (CL) + Cyrillaceae (CY): 28-29
2 genera, 60 species (CL) and 2 genera, 2 species (CY). Order Ericales. Trees and shrubs. Spiral leaf
arrangement (Fig. 11.30). Stipules absent. Leaves simple, entire (Fig. 11.29 Clethra spp., Cyrilla),
toothed only in some AS-Clethra (Fig. 11.30). Venation camptodromous, s.t. with fimbrial veins
(Fig. 11.30 Clethra spp.) or different (Cyrilla). Indument: stellate hairs (Clethra spp.) or none (e.g.
Clethra delavayi, Cyrilla). Inflorescence racemose (Fig. 11.31), terminal (Clethra), lateral (Cyrilla).
Flowers with 5–7 free petals (poss. fused at its base). Ovary superior, axile placentation. Fruits
capsules (CL), drupes or 2–5 winged samaras (CY).

SCARRONE (Fig. 11.31): Clethra longispicata (AS).
SAPOTACEAE (but latex).
RAUH: Cyrilla sp.
LAURACEAE (but late suberization of stems).
ERICACEAE (but frequent apex abortions). AUBRÉVILLE (Fig. 11.30): Clethra cf. pringlei (AM-C).
11.9 Clethraceae (CL) + Cyrillaceae (CY): … 201
Fig. 11.29 Clethra cf. pringlei, camptodromous venation, Belize
Fig. 11.30 Clethra cf. pringlei, branch plagiotropic by apposition, Cockscomb, Belize. Clethra spp. a Cl. cuneata,
entire leaf, fimbrial vein, Bolivia; b Clethra delavayi, toothed leaf, venation ± brochidodromous, Burma, from
herbarium specimens, CBG H
Fig. 11.31 Clethra scabra, racemose infructescence, Southern Brazil. Clethra longispicata, SCARRONE’s model,
Flores, Indonesia. Cyrilla sp., entire leaves with cuneate laminas, Belize
The AM-AS distribution of Clethraceae reveals the Laurasian origin of this group. Africa, formerly
part of the Gondwanan continent would have not hosted this family. The same distribution pattern
exists also in several other groups (e.g. Cornaceae, Symplocaceae, Theaceae).
11.10 Clusiaceae + Calophyllaceae + Bonnetiaceae + Hypericaceae:

2-3-4-11-13-24-29-38-41-42
Four families encompassing 40 genera and 1850 species. Order Malpighiales. Exudate sticky,
resinous, pale yellow (Fig. 11.32 Caraipa densiflora), bright yellow or orange-coloured, (Guttiferae
is the ancient name for Clusiaceae but gutta-percha is the exudate of some Palaquium gutta, Sapo-
taceae). Hemiepiphyte species usually have coriaceous leaves and can develop long adventicious
roots (Fig. 11.34). Stilt-roots Symphonia (Fig. 11.36). Thorny trunk in Cratoxylum (Fig. 11.38).
Stems not lenticellate. Apical buds often embedded in leaf bases (Fig. 11.35). Immediate branching
frequent (Fig. 11.37, e.g. Calophyllum, Clusia). Opposite leaf arrangement (Calophyllaceae, Clusi-
aceae and Hypericaceae) or spiral (Fig. 11.40 Bonnetiaceae, Calophyllaceae: Caraipa, Kielmeyera,
Fig. 11.38 Mahurea, these taxa all AM). Stipules absent or minute (Fig. 11.36 Symphonia, AM and
MA, Garcinia). Leaves shortly petiolate or even sessile (Fig. 11.40 Archytaea). Opposite-leaved
species have young leaves of the same pair at first appressed then spreading (Fig. 11.33 Vismia
baccifera); species with spiral arrangement have young leaves folded in ‘V’ (Fig. 11.38). S.t.
numerous parallel secondary veins (Fig. 11.39) or venation indistinct (Clusia, AM). Leaves with
translucent glands or black dots (Fig. 11.40 Hypericum). Inflorescences terminal (Hypericum,
Fig. 11.39 Vismia pp.) or lateral (Fig. 11.39 Mesua (AS), Garcinia, Mammea (Pantrop), Mon-
trouziera (New Cal.). Flowers showy, with radial symmetry (Fig. 11.34), free sepals and petals, and
numerous stamens. Ovary with axile placentation. Fruits usually berries (Fig. 11.35 Clusia spp.,
Vismia) or capsules (Caraipa).
LEEUWENBERG: CLU: Montrouziera (NC).
FAGERLIND: CLU: Tovomita plumieri (AM).
RAUH: CAL: Calophyllum inophyllum (AS),
C.brasiliense (AM), Mammea americana (AM), CLU:
Similar families: Clusia sp., Pentadesma butyracea (AF).
APOCYNACEAE but they have lenticellate branches AUBRÉVILLE (only in Bonnetiaceae): Ploiarium (AS),
and often a pure white latex. Archytaea (AM).
ROUX: CLU: Garcinia edulis (Mexico), G. cymosa (NG),
G. livingstonei (AF); Hypericaceae: Vismia laxiflora
(AM).
MANGENOT: CAL: Mesua ferrea (AS).
MASSART: CLU: Platonia insignis, Symphonia
globulifera (AM).
TROLL (only in Hypericaceae): Vismia macrophylla (AM),
Hypericum androsaemum (EU).
Bonnetiaceae (Fig. 11.40 Archytaea, Bonnetia, Ploiarium), Calophyllaceae (Fig. 11.37 Calo-
phyllum), Fig. 11.38 Mahurea, Mammea, Mesua and Hypericaceae (Fig. 11.38 Cratoxylum,
Hypericum, Harungana, Fig. 11.39 Vismia) are currently treated as separate families.
Trees, s.t. very tall: CLU (Symphonia, AM, MA; Allanblackia, AF; Moronobea, Platonia,
AM; Garcinia), CAL (Calophyllum, Caraipa). Symphonia globulifera bears stilt-roots
(Fig. 11.36) and is a dominant component of swamp forests of Northern AM-S. Shrubs or
small trees: CLU (Tovomita, AM), HYP (Harungana, AF; Vismia, AM). Woody epiphytes CLU
(Clusia rosea). Herbs HYP (Hypericum). Clusia is perhaps the only woody genus endowed with
Crassulacean acid metabolism (CAM) [6], an adaptation common in Bromeliaceae and Cactaceae.
11.10 Clusiaceae + Calophyllaceae … 203
Fig. 11.32 Clusia sp., gutta pale yellow Fig. 11.33 Vismia baccifera, appressed young leaves,
Belize
Fig. 11.34 CLUSIACEAE. Clusia sp., hemi-epiphyte with adentitious-roots embracing the host trunk, Belize. Clusia
fockeana, showy flowers, Venezuela. CLUSIACEAE. Clusia sp., RAUH’s model, Belize
Fig. 11.35 CLUSIACEAE. Clusia sp., coriaceous berries with axile placentation, Belize. CLUSIACEAE. Garcinia
sp., apical meristem hidden between petiolar bases, Thailand. CLUSIACEAE. Moronobea coccinea, short shoot
bearing a terminal flower, French Guiana
Fig. 11.36 CLUSIACEAE. Platonia insignis, rhythmic growth, French Guiana. CLUSIACEAE. Symphonia
globulifera, stilt-roots and minute stipules, Belize
Fig. 11.37 CLU: Symphonia globulifera, flower bud, Belize. CALO: Calophyllum brasiliense, immediate branching
of the terminal growth unit, Venezuela. CALO: Caraipa densiflora, pale yellow gutta, French Guiana
Fig. 11.38 HYPERICACEAE: Cratoxylum cochinchinense, spiny trunk, Thailand. CALOPHYLLACEAE: Mahurea
palustris, young leaves ‘V’-folded, French Guiana. CALO: Mesua ferrea, flush of young leaves, Thailand
Genera are usually endemic to a single continental mass. Garcinia (CLU), Mammea (CAL) and
Symphonia (CLU) are pantropical. Calophyllum (CAL) has a discontinuous distribution pattern
(AM-AS).
Dugout canoes or ‘praos’ used in the earliest trans-pacific migrations were probably built with
trunks of the sea-shore tree Calophyllum inophyllum (‘Bitangor’ Malaysia). Timbers: Calophyl-
lum, Platonia, Mesua ferrea. Fruit trees: Garcinia mangostana (mangosteen, ‘manggis’),
11.11 Combretaceae: 21-22-24-30-40-41-56 205
Fig. 11.39 CALO: Mesua sp. lateral flowering, Thailand. HYP: Vismia baccifera, numerous secondary veins, terminal
flowering, French Guiana. HYP: Vismia sp., gutta turning red, Venezuela
Fig. 11.40 HYPERICACEAE: Hypericum montanum, lamina margins with dark dots, Switzerland. BONNETIA-
CEAE: Bonnetia sessilifolia, bush with showy flowers, Venezuela. BONNETIACEAE: Archytaea triflora, leaves
sessile and coriaceous, Venezuela
Mammea americana (apricot of San Domingo). Oil seeds: Allanblackia, Garcinia, Mammea,
Pentadesma butyracea (lamy butter, for cooking and candles). Medicinal: Hypericum spp. Oil for
soap-factories (seeds of Calophyllum inophyllum). Pigments for varnishes (Garcinia gummi-
gutta).
! Guttiferae is a good example of a group split into separate families by APG classifications and
showing consistency in the different ranges of architectural models existing in each family. However,
this does not occur in every large taxonomic group, because phylogenies are not expected within
architectural models I.3.1 [7].
11.11 Combretaceae: 21-22-24-30-40-41-56
14 genera and 500 species. Order Myrtales. Several Terminalia species exhibit crown-shyness.
Trunk buttresses frequent (Fig. 11.48). Cross-sections of lianas show spots of phloem scattered in
wood (Fig. 11.43). Climbers often have woody persistent petiolar bases (Fig. 11.45 Combretum).
Some decumbent-short shoot-thorny shrubs in savannas (Fig. 11.49). No latex, but in some cases
sectioned bark secreting gum (Fig. 11.49). Bark sloughing off in strips (Fig. 11.45) or scaly
(Fig. 11.47). Twigs or trunks not lenticellate. Branches often plagiotropic by apposition (Fig. 11.42
Bucida buceras, Fig. 11.49 Terminalia spp.). Trees usually evergreen, s.t. deciduous (Fig. 11.41).
Fig. 11.41 Terminalia randii, deciduous tree, Fig. 11.42 Bucida (syn. Terminalia) buceras, branches
Botswana plagiotropic by apposition, Venezuela
Leaf arrangement spiral (Buchenavia, Conocarpus, Lumnitzera, Terminalia) or opposite (Fig. 11.43
Anogeissus, Fig. 11.45 Combretum, Laguncularia), the two leaves of a same pair often subopposite
(Combretum, Guiera). Stipules absent. Leaves simple and entire. Indument of simple or stellate-
peltate hairs (Combretum), small “black dots” (Guiera), rarely absent (Lumnitzera). Venation often
camptodromous (Fig. 11.43), s.t. indistinct (Fig. 11.48). Leaf base often glandular (Conocarpus) or
glands at vein axils (Fig. 11.46 Conocarpus, Terminalia). Inflorescences in racemes (Fig. 11.46),
spikes (Fig. 11.44) or panicles. Flowers small, 4–5 free petals and 8–10 stamens (Fig. 11.47). Fruits
one-stone drupes, s.t. ribbed (Fig. 11.48) or winged (Fig. 11.44 Combretum).
Similar families: AUBRÉVILLE: (Fig. 11.49): Bucida buceras,
Buchenavia guianensis, Terminalia superba, T.
LYTHRACEAE (Lagerstroemia), with
ivorensis.
subopposite leaves and camptodromous venation.
ATTIMS: Laguncularia, Lumnitzera.
MALPIGHIACEAE with their many twining and
glandular species. CHAMPAGNAT: Combretum imberbe (AF), Guiera
senegalensis?
TROLL: Anogeissus acuminata, (AS).
Trees (Anogeissus, Buchenavia, Terminalia). Shrubs (Bucida, Conocarpus, Guiera, Laguncu-

laria, Lumnitzera, Combretum, Terminalia) s.t. subterranean (Combretum). Lianas (Combretum
spp.), no herbs or aquatic species. Frequent in African savannas (Fig. 11.41), also an important
component in mangroves of all tropical coasts (AM: Fig. 11.42 Bucida buceras, Fig. 11.46 Cono-
carpus, Fig. 11.48 Laguncularia, AF-AS-OC: Fig. 11.48 Lumnitzera (AF, AS, OC). Timbers, in AF:
Terminalia ivorensis (‘framiré’), T. superba (‘limba’), in AS: Anogeissus. Terminalia catappa
(‘ketapang’, Malabar almond, pagoda tree) is often cultivated as a shade tree. Several ornamentals:
especially Combretum indicum (syn. Quisqualis indica). Tanins (bark of Laguncularia racemosa,
drupes of Terminala chebula, India).
! In the Caribbean region, Conocarpus erectus presents a continuum of forms between the glabrous
and pubescent extremes, even in the same area (Fig. 11.46).
11.11 Combretaceae: 21-22-24-30-40-41-56 207
Fig. 11.43 a Camptodromous venation (e.g. Combretum spp.). b Twig plagiotropic by apposition, cp: cataphyll.
c Base of lamina glandular (e.g. Conocarpus). d Liana with phloem poles scattered in wood. e Anogeissus acuminata,
opposite leaf arrangement in a plagiotropic branch, Thailand
Fig. 11.44 Combretum sp., ‘bottle-brush’ inflorescence, French Guiana. Combretum sp., winged drupes, Venezuela.
Combretum sp., winged drupes, Botswana
Fig. 11.45 Combretum mossambicense, opposite leaf arrangement, Botswana. Combretum sp., bark sloughing off in
strips, camptodromous venation, Belize. Combretum sp., petiolar bases modified into hooks
Fig. 11.46 Conocarpus erectus, terminal infructescence, Venezuela. Conocarpus erectus, glabrous form, traces of
glands visible at vein axils (upper side of lamina), Belize. Conocarpus erectus, pubescent form, Belize
Fig. 11.47 Laguncularia racemosa, scaly bark, leaves with indistinct venation, mangrove, Tulum, Mexico.
Lumnitzera littorea, leaves coriaceous, inflorescence racemose, lateral, Bali
Fig. 11.48 Laguncularia racemosa, ribbed fruits, Santa Catarina, Brazil. Lumnitzera littorea, large tree in mangrove,
leaves glabrous, venation indistinct, Bali
11.12 Connaraceae: 12-21-48-49-53-57 209
Fig. 11.49 Terminalia catappa, inflorescence in spike, Southern Brazil. Terminalia amazonia, wounded trunk
secreting gum, Belize. Teminalia prunioides, thorny short shoots of a shrub, Botswana. Terminalia sp.,
AUBRÉVILLE’s model, flushes of leaves after leaf-fall, Seram, Indonesia
11.12 Connaraceae: 12-21-48-49-53-57
12 genera and 170 species. Order Oxalidales. Plants not aromatic, without latex, but s.t. cut bark
producing a reddish exudate (Manotes, Connarus). Cross-sections of lianas showing excentric blocks
(Fig. 11.51) or alternate rings of phloem and xylem (Fig. 11.51). Twigs lenticellate (Fig. 11.52).
Lianas with stems twining at their extremities (Fig. 11.51). Spiral arrangement of leaves, rarely
two-ranked arrangement (Connarus venezuelanus). Terminal short shoots in Burttia (Fig. 11.55).
Stipules absent. Leaves usually compound pinnate (Fig. 11.51) or trifoliolate (Fig. 11.52), rarely
simple and ‘unifoliolate’ (Fig. 11.55 Burttia, AF; Fig. 11.53 Connarus venezuelanus, AM; Manotes,
AF; Ellipanthus, Paleo). Petiolar base distinctly pulvinate (Fig. 11.52). Venation more or less
brochidodromous (Fig. 11.53, Fig. 11.54), densely reticulate (Fig. 11.54), sunken at upper side.
Flowers with 5 petals and as many stamens and staminodes. Ovary inferior, 5-carpels. Fruits con-
sisting of 1 to 5 follicles (Fig. 11.53) or drupes (Fig. 11.50).
Large woody lianas (Paleo: Agelaea, Cnestis, Ellipanthus, Pantrop: Connarus, Rourea), and
several small trees (Fig. 11.53 Connarus spp.).
Fig. 11.50 Agelaea sp., liana bearing infructescences, Madagascar
Fig. 11.51 a Prehensile trunk extremity of a liana, French Guiana. b CHAMBERLAIN’s model (Connarus, Rourea).
c Excentred xylem blocks of several species. d Phloem rings (darker) alternating with xylem rings, an unidentified liana,
Venezuela
11.12 Connaraceae: 12-21-48-49-53-57 211
Fig. 11.52 a, b Agelaea sp., transition from simple leaves to trifoliolate leaves, Indonesia. Agelaea sp., trifoliolate
leaves, Cameroon. c Agelaea borneensis, trunk strongly lenticellate, Indonesia. d Connarus fasciculatus, cauliflory,
clusters of follicular fruits, French Guiana
Fig. 11.53 Connarus fasciculatus, pulvinate petiolar bases and campto-brochidodromous venation (underside of a
leaflet), French Guiana. Connarus venezuelanus, small tree in savanna, leaves unifoliolate or trifoliolate, Venezuela
Fig. 11.54 Rourea glabra, infructescences reduced to a single follicle, Belize. An unidentified specimen, leaves
whitish on underside, venation brochidodromous, French Guiana
Fig. 11.55 Rourea mimosoides, trunk bearing a lateral short shoot with clustered compound leaves, Kebun Raya, Java.
Burttia prunoides, terminal short shoots (sh), lateral flowering (plagiotropy by apposition?), unifoliolate leaves with
articulate petiole (a), from a herbarium specimen, CBG H
A discreet family, mostly found in pristine forests, however several species are common in open
places (e.g. Agelaea vestita, Connarus ferrugineus, AS). Timber: Ellipanthus tomentosus (India and
South-East Asia). Several “non-wood products”: tanins, dyes and healing molecules. Seeds of
Connarus ferrugineus (Indomalesia) are poisonous [8]. Stems of Agelaea spp. were used in past times
in Sumatra to bind rafts and nets [9].

LEGUMINOSAE-PAPILIONOIDEAE lianas when their CORNER: Jollydora (AF-W).
stipules not visible (leaves pinnately compound and petiole CHAMBERLAIN (Fig. 11.51-b): Agelaea spp.
pulvinate at base for both families). (Paleo), Connarus, Rourea (Pantrop).
In CONNARACEAE only stem extremities are twining, AUBRÉVILLE? (Fig. 11.58) Burttia (AF).
whereas stems of PAPILIONOIDEAE are twining along TROLL’s model probably absent.
their entire length.
! Chewed fruits of several species of African Connarus are so astringent when chewed that one is
unable to whistle (personal observation!).
11.13 Convolvulacaeae: 53-54-57
55 genera and 1850 species. Order Solanales. Trunk cross-sections of lianas often with phloem
intrusion in wood (Fig. 11.60b Neuropeltis). Stems of herbaceous lianas always twining to right [10].
Leaves simple, often cordate (Fig. 11.59), palmatilobate (Ipomoea spp.), pinnately compound
(Fig. 11.59 Ipomoea spp.) or compound palmate (Fig. 11.60a Merremia). Venation pinnate
(Fig. 11.57 Dicranostyles, Erycibe) or vaguely palmate. Underside of leaves with an indumentum of
short hairs (Fig. 11.57), upper side glabrous. Flowers funnel-shaped (Fig. 11.59), less often disk-
shaped (Evolvulus), set of bracteoles surrounding the calyx (Fig. 11.56). Ovary superior, deeply 2–4
lobed. Fruits are usually dehiscent capsules (Fig. 11.56).
Herbaceous twining lianas with latex (Ipomoea, Merremia, Fig. 11.60-d Xenostegia, etc.) or
woody lianas without latex (Fig. 11.57), Dicranostyles, AM; Erycibe, AS,-AU; Neuropeltis, AF-
AS). Prostrate herbs or small shrubs (Fig. 11.58 Ipomoea pes-caprae, Fig. 11.58 Evolvulus sp.).
Small parasitic twining herbs (Cuscuta).
11.13 Convolvulacaeae: 53-54-57 213
Similar families: Architecture poorly known:

BORAGINACEAE and SOLANACEAE have leaves ACOSTA (Fig. 11.60c): Evolvulus cf. alsinoides
similar in shape, however, leaf laminas in these (herb of open sites).
families are usually pubescent on the upper side. BELL (Fig. 11.58): Ipomoea pes-caprae (prostrate
herb of back-shores).
Fig. 11.56 Argyreia

splendens, set of bracteoles,
Thailand
Fig. 11.57 Dicranostyles guianensis, Leaves with pinnate venation, pubescent apex and conduplicate leaf-folding,
forest liana, French Guiana. Erycibe sp., forest liana, Bali
Fig. 11.58 Evolvulus alsinodes, subshrub, Bali. Ipomoea pes-caprae, prostrate stems on sand, Flores, Indonesia
Fig. 11.59 Ipomoea quamoclit, flower funnel-shaped, leaves compound pinnate, La Llovizna, Venezuela. Ipomoea
sp., cordate leaves, La Escalera, Venezuela
Fig. 11.60 a Merremia macrocalyx, leaves compound palmate, Venezuela. b Neuropeltis sp., trunk cross-section of a
liana with phloem intrusions in wood. c Evolvulus cf. alsinodes, prostrate subshrub, Bali. d Xenostegia tridentata
subsp. angustifolia, herb twining to the right, Maun, Botswana
Edible tubers: Ipomoea batatas (Sweet potato, native to Central America but cultivated mostly in
sub-Saharan Africa). Ornamentals: Ipomoea spp. Some psychotropic substances (seeds of Ipomoea
purpurea). In past times, medicinal (Erycibe spp.) used chiefly in childbirth [9].
! The absence of latex in the woody species of this family is somewhat surprising, as is the case in
Bonamia, Erycibe and Maripa, three genera belonging to three different tribes (respectively, Cresseae,
Erycibeae and Maripeae).
11.14 Cornaceae (C) + Nyssaceae (N): 13-29 215
11.14 Cornaceae (C) + Nyssaceae (N): 13-29
2 genera and 80 species, order Cornales. Plants without latex and not aromatic. Leaf arrangement
opposite (C: Cornus, N: Mastixia kaniensis), alternate and spiral (Mastixia spp.) or two-ranked
(Fig. 11.62 Alangium). Leaves simple and entire. Venation palmate (Alangium platanifolium) or
leaves tripliveined (Alangium salviifolium). If venation pinnate, then camptodromous (Fig. 11.62).
Indument absent (leaves glabrous). Inflorescences umbels, corymbs or panicles (Fig. 11.61).
Flowers with radial symmetry, 4–5 carpels and as much stamens. Ovary superior. Fruits drupaceous.
Trees (C: Alangium, AF-AS-OC; N: Mastixia, AS), shrubs (Cornus, AM, AS) and a few lianas
(e.g. Alangium salviifolium).
Cornaceae and Nyssaceae have a rather discontinuous American-Asiatic distribution, being almost
absent from Africa.

THEACEAE: when leaves spiral, as Mastixia, FAGERLIND: Cornus florida (AM-N).
Nyssa.
MASSART: Alangium uniloculare, A. platanifolium,
RUBIACEAE, but their leaves are stipulate.
Nyssa sylvestre (AM-N).
ROUX: Alangium salviifolium (AS).
! Cornales is the most basal order in the Asteridae clade (APG IV). Relationships among families
within Cornales remain unclear [11]. This group which seems to have arisen from nowhere, about 125
Myr ago, will puzzle botanists for future decades.
Fig. 11.61 Cornus sanguinea (the European Dogwood), inflorescence corymbiform

Fig. 11.62 Mastixia kaniensis, leaves opposite, terminal cymose inflorescences, venation camptodromous, Papua,
CBG H specimen. Alangium uniloculare, Malaysia. Laminas asymmetrical. Branching plays a three-beat rhythm in
which every third node is ramified in a monopodial and plagiotropic branch (not all leaves are illustrated)
11.15 Corynocarpaceae: 18
A monogeneric family, 5 species, order Cucurbitales. Trees with bitter bark. No latex and not
aromatic. Spiral leaf arrangement (Fig. 11.63). Branches plagiotropic by apposition (Fig. 11.63).
Stipules intrapetiolar. Leaves leathery and entire. Inflorescences in terminal panicles. Flower parts in
five. Fruits drupes.
SW Pacific. Seeds of Corynocarpus laevigatus are poisonous, but when roasted, they are a staple
food of Maoris (‘Karaka’ nuts in New Zealand).
GOODENIACEAE also have short petioles and branches plagiotropic by apposition.
Fig. 11.63 Corynocarpus laevigatus, apposition growth, Kebun Raya, Java. Growth, FAGERLIND’s model, Kebun
Raya
11.16 Cunoniaceae (C) + Brunelliaceae (B): 32-50 217
11.16 Cunoniaceae (C) + Brunelliaceae (B): 32-50
Two closely related families encompassing 28 genera and 372 species, of which 155 in Weinmannia
(C). Order Oxalidales. Small evergreen trees and shrubs without latex, not aromatic. Inner bark or
wood s.t. red. Opposite leaf arrangement. Cunoniaceae have large, intra- or interpetiolar stipules
(Fig. 11.65), Brunelliaceae, small stipules. Leaves compound pinnate (Fig. 11.64) or trifoliolate, s.t.
simple. Leaves or leaflets usually toothed. Cunoniaceae are glabrous and Brunelliaceae, pubescent.
Inflorescences racemose in Cunoniaceae (Fig. 11.65), complicated multiramified cymes in Brunel-
liaceae (Fig. 11.65). Fruits capsules or follicles.
Brunelliaceae (monogeneric) grow in the understoreys of American rainforests. Cunoniaceae are
mainly a Southern hemisphere group (AU, New Cal). Few uses are known. Bark of Weinmannia (e.g.
W. denhamii, Vanuatu archipelago) produces a red dye.
Similar families:
RUBIACEAE, but leaves entire.
ATTIMS and RAUH are quite common.
RHIZOPHORACEAE, but with many
lenticels. TROLL probably absent
Fig. 11.64 Weinmannia sp., Flores, Indonesia
Fig. 11.65 Weinmannia sp., stipules large, interpetiolar, Madagascar. Cunonia balbisiana, simple leaves, inflores-
cences racemose. Brunellia sp., multiramified cymose inflorescence
11.17 Cycadaceae + Zamiaceae: 1 (Gymnosperms)
Two families totalizing 11 genera and 285 species, Cycadaceae being monogeneric. Order Cycadales
(Cycadopsida). Dioecious pachycaulous small palm-like (Fig. 11.68) trees with large pith
(Fig. 11.67). CYC: Leaves compound pinnate (Fig. 11.66). Base of leaflets decurrent on the rachis
(Fig. 11.66). Leaf-folding circinately involute. CYC: venation indistinct. ZAM: leaflets with parallel
venation (Fig. 11.67). Sexuality lateral in female plants, terminal in males. Sporophylls in deter-
minate cones covered by peltate scales (Fig. 11.66). Zamia “seeds”, ovules bright red when ripe
(Fig. 11.67).

At first sight, palm trees (Arecaceae)! But these have a very
CORNER: female plants. CHAMBERLAIN: male plants.
different type of leaf insertion (sheathing petiole).
Cycadaceae: AF-E, AS and AU. Zamiaceae: Tropical AF, AM and AU. Lepidozamia hopei
(Queensland), at 20 m the tallest Cycad [10]. Cycad seeds are well-known to be highly poisonous
[12] and must be carefully processed to remove toxins before they are edible. If they are not properly
processed they can cause liver damage and even death.
Cycadopsida are reported from the Triassic onwards, they belong to the Gymnospermae. Some-
times nicknamed dinosaur plants, cycads appeared on the earth about 300 million years ago and were
at their greatest diversity during the Triassic and Jurassic.
Fig. 11.66 Cycas sp., sapling, Thailand. Cycas rumphii, decurrent insertion of leaflets on foliar rachis. Leaflets with
strong midrib. Encephalartos horridus, scales removed to show orange-coloured ovules
Fig. 11.67 Encephalartos ghellinckii, trunk cross-section showing a large pith (private collection, France). Zamia
polymorpha, seeds, Shipstern, Belize. Zamia sp., leaflet, BG Utrecht University
11.18 Cyclanthaceae: 1-57 (Monocotyledons) 219
Fig. 11.68 Encephalartos longifolius, pachycaul, Villa Thuret, France. Zamia sp., BG Orinoco, Venezuela Zamia
wallisii, leaflets with parallel venation
11.18 Cyclanthaceae: 1-57 (Monocotyledons)
12 genera and 225 species. Order Pandanales. At first sight, Cyclanthaceae look like Palms, Araceae
or Pandanaceae. Their plicate leaves (Fig. 11.69) are reminiscent of Arecaceae, their inflorescences
are of Aracean style (Fig. 11.70) and their compound fruits look like those of a small Pandanus.
Trunk not woody. Leaves plicate, fan-shaped: Carludovica, bifidous: Asplundia, Evodianthus,
Thoracocarpus (Fig. 11.70) or simple (Ludovia). Flowers reduced with numerous stamens. Stami-
nate and pistillate flowers in separate, alternating cycles around the spadix (Cyclanthus) or in stag-
gered rows (Fig. 11.70). Filaments of stamens becoming very long and drooping (Fig. 11.70). Fruits
fleshy, berry-like, often a ‘compound fruit’.
Similar families: Large herbs or climbing epiphytes (Fig. 11.69).

All species American. Panama hats or ‘toquillas’ are
Young or juvenile Arecaceae.
made in Ecuador, not Panama!, using foliar fibres of
Carludovica palmata.
Fig. 11.69 Asplundia rigida, a climber with clamp-roots and plicate leaves, Guadeloupe
Fig. 11.70 Thoracocarpus bissectus, leaves bifid, St-Elie, French Guiana. Asplundia rigida, spike of staminate and
female flowers in staggered rows, Guadeloupe. Dicranopygium sp., inflorescence with drooping, senescent stamen
filaments, Venezuela
References
1. Müller, S., S. Karsten, J. Salazar, J. Naumann, M.A. Jaramillo, C. Neinhuis, S. Taylor, T.S. Feild, and S. Wanke.
2015. Intercontinental long-distance dispersal of Canellaceae from the New to the Old World revealed by a nuclear
single copy gene and chloroplast loci. Molecular Phylogenetics and Evolution 84: 205–219.
2. Feild, T.S., D.S. Chatelet, L. Balun, E.E. Schilling, and R. Evans. 2012. The evolution of angiosperm lianescence
without vessels - climbing mode and wood structure–function in Tasmannia cordata (Winteraceae). New
Phytologist 193: 229–240.
3. Upadhyay, R., G. Jaiswal, and S. Ahmad. 2010. Anti-termite efficacy of Capparis decidua and its combinatorial
mixtures for the control of Indian white termite Odontotermes obesus (Isoptera: Odontotermitidae) in Indian soil.
Journal of Applied Sciences and Environmental Management 14: 101–105.
4. Diamond, J. 2006. Effondrement, 870 pp. Paris: Gallimard.
5. Martínez, C., S. Madriñán, M.S. Zavada, and A. Jaramillo. 2013. Tracing the fossil pollen record of Hedyosmum
(Chloranthaceae), an old lineage with recent Neotropical diversification. Grana 52: 1–20.
6. Lüttge, U. 2008. Clusia: Holy Grail and enigma. Journal of Experimental Botany 59: 1503–1514.
7. Hallé, F., and Keller, R. 2019. Mais d’où viennent les plantes?, 180 pp. Arles: Actes Sud.
8. Wiart, C. 2006. Medicinal Plants of Asia and the Pacific, 336 pp. Boca Raton: CRC Press.
9. Burkill, I.H. 1935. A dictionary of the economic products of the Malay peninsula, vol. I and II, 2402 pp. Oxford:
University Press
10. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3rd ed., 1022 pp. Cambridge: Cambridge University Press.
11. Xiang, Q.Y.J., D.T. Thomas, and Q.P. Xiang. 2011. Resolving and dating the phylogeny of Cornales—Effects of
taxon sampling, data partitions, and fossil calibrations. Molecular Phylogenetics and Evolution 59: 123–138.
12. Grant, W.M. 1963. Toxicity of cycads. Economic Botany 17: 270–302.
Dichapetalaceae to Dipterocarpaceae
12
12.1 Dichapetalaceae: 7-8-54
3 genera and 170 species. Order Malpighiales. Plants without latex and not aromatic. Lianas present
intrusions of phloem in the wood (Fig. 12.2). Stems or trunks lenticellate, with lenticels disposed in
rows parallel to stem axis (Fig. 12.3a). Leaves two-ranked (Fig. 12.1). In Tapura and Stephano-
podium inflorescences are glomerulate and inserted on the petiole (Fig. 12.3b). Leaves stipulate,
simple and entire. Indument of appressed hairs on buds and apices. Venation brochidodromous,
very densely reticulate (Fig. 12.3). Glands scattered on the underside of lamina (Fig. 12.3), s.t. on the
Fig. 12.2 Phloem intrusions
upper side. Indument of short appressed hairs

on apices and buds. Flowers small with 4–5 free
petals. Fruits pubescent drupes (Fig. 12.3)
enclosing 1–3 seeds. Trees (Tapura), but mostly
lianas (Dichapetalum spp.), No herbs, no
Fig. 12.1 Dichapetalum sp., ROUX’s model, Cameroon aquatic plants.
https://doi.org/10.1007/978-3-031-05942-1_12
222 12 Dichapetalaceae to Dipterocarpaceae
Fig. 12.3 Dichapetalum sp., drupes, Venezuela. Dichapetalum sp., venation densely reticulate, glands scattered on
underside of lamina, Venezuela. a, b, c Tapura antillana, sapling, Guadeloupe. a Lenticels in rows. b Inflorescences
inserted on the petiole. c Architectural model in between ROUX’s and MANGENOT’s, (lateral branches represented in
the same plane), f: fallen leaves, c: scale-leaf, st: stipule

As a rule, no possible confusion with other ROUX (Fig. 12.3c): Tapura antillana, a small tree.
families. MANGENOT: frequent in lianas.
Dichapetalaceae are pantropical, frequent in Africa, and entirely dependent on forests. The self-
supporting stage can last a very long time. Several species are very toxic (fluoroacetic acid). No
important economical use reported.
12.2 Dilleniaceae: 14-26-28-54
11 genera and 425 species. Order Dilleniales. Trees, shrubs with scaly bark, but especially twining
lianas (Doliocarpus, Pinzona, Tetracera) with broad vessels in wood (water lianas), their stem cross-
sections (Fig. 12.7) showing concentric rings of phloem (Fig. 12.6). The only herbaceous genus is
extratropical: Acrotrema (Tasmania). Spiral arrangement of leaves (Fig. 12.4 Curatella americana,
French Guiana). Petiole canaliculate to winged, the wing-like stipules adnate to the petiole
(Fig. 12.5). Leaves rough, scabrous, without glands. Young leaf-folding conduplicate. Venation
secant on lamina margin (Fig. 12.4). The Australasian genus Hibbertia is atypical in that leaves are
12.2 Dilleniaceae: 14-26-28-54 223
Fig. 12.4 Curatella americana, secant venation Fig. 12.5 Dillenia sp., stipuliform petiolar expansions
small, ericoid and without secant venation. Flowers solitary or in racemose, or cymose inflores-
cences. Flowers usually yellow, or white, with spiral imbricated tepals and numerous stamens, calyx
cupular in Davilla (Fig. 12.6). Fruit s.t. surrounded by a fleshy calyx (Fig. 12.6 Dillenia).
The majority of Dilleniaceae are native to Australasia and tropical America, Tetracera is
pantropical. Curatella Americana (‘curata’, ‘chaparro’, ‘cajueiro bravo’) a typical small tree of South
American savannas impacted with periodic fires, has a deeply fissured bark.
! Water lianas are often used to demonstrate that one need never be thirsty in the forest, but
this involves cutting the trunk of a Tetracera or Pinzona species, and so destroying the plant!
Fig. 12.6 Cross-section of a liana, rays and concentric rings of phloem. Davilla rugosa, cupular calyx, Santa Catarina,
Brazil. Davilla kunthii, infructescence, Belize. Dillenia indica, fruits inserted in a fleshy calyx, Kebun Raya, Java

As a rule, no possible confusion with other families. OLDEMAN: likely for the lianas of this family.
SCARRONE: Dillenia spp. (AS).
Fig. 12.7 Dillenia sp., large yellow flower, BG Singapore. Pinzona coriacea, liana, trunk section, Belize. Tetracera
sp., twining liana, Sumatra
12.3 Dioscoreaceae: 55-56-57 (Monocotyledons)
3 genera and 630 species. Order Dioscoreales. Lianas, often spiny (spines in stipular position,
Fig. 12.9). Spiral or opposite leaf arrangement. Plants with starchy tubers rhizomes (up to 2 m in
length for Stenomeris). Leaves simple or trifoliolate (Fig. 12.9). Venation campylodromous
(Fig. 12.9). Inflorescences axillary, panicles, racemes or spikes. Flowers small with 6 tepals and 6
stamens. Ovary inferior. Fruits angular (Fig. 12.9) or winged capsules (Fig. 12.8), or berries (Tamus).
Fig. 12.8 Dioscorea sp., winged capsules, Thailand
Fig. 12.9 Dioscorea dregeana, trifoliolate leaves, Bali, Indonesia. Dioscorea sp., cordate leaf with campylodromous
venation and angular capsules, Southern Brazil. Dioscorea sp., spines in stipular position, Seram
12.4 Dipterocarpaceae: 2-5-7-8-15-18 225
Yams are cultivated everywhere in the tropics. Diosgenine, a precursor of progesterone and
cortisone is synthesized by numerous species of Dioscorea (oral contraceptives).
Disocorea is pantropical, Stenomeris Indomalaysian.
! The Dioscoreaceae resemble ‘Dicotyledons’, and can be confused with Convolvulaceae. What is
more, seedlings exhibit a second, but very reduced cotyledon [1].
12.4 Dipterocarpaceae: 2-5-7-8-15-18
16 genera and 535 species. Order Malvales. Plants without latex (but wood resinous in several
species) and leaves not aromatic. By means of artificial cuts (tapping) wood of several tapped tree
species produce resins or oleo-resins (dammar). Crown shyness can be impressive (Fig. 12.11,
Fig. 12.12). Growth can be strongly rhythmic or weakly rhythmic. Leaf arrangement spiral in all
stems, or two-ranked on branches (Fig. 12.15, e.g. Hopea), depending on the species and/or the
developmental stage. Stipules present, s.t. large and hood-like (Fig. 12.13 Dipterocarpus, Shorea).
Leaves simple, entire. Petiole s.t. slightly swollen or bent distally (Fig. 12.15) (Dryobalanops,
Hopea, Shorea). Venation often camptodromous, s.t. secant and scalariform (Fig. 12.10 Diptero-
carpus, Shorea), rarely brochidodromous (Fig. 12.14b Dipterocarpus pp.) or with a submarginal vein
(Fig. 12.14c, Cotylelobium, Dryobalanops). Leaves without glands except Monotes and Marquesia
with one gland at the base of the lamina (Fig. 12.14a). Indument of fasciculate hairs in Anisoptera,
Shorea (Fig. 12.16). Several species (Hopea) with underside of lamina bearing domatia in vein axils
(Fig. 12.16). Flowers with contorted corolla, 5 free petals and numerous stamens. Mass flowerers
pollinated by thrips. Ovary superior. Fruits indehiscent capsules rimmed by two, s.t. three, wings
(Fig. 12.10) which are in fact long persistent sepals (the family owes its name to this feature), or
without persistant sepals (Vateria).
Trees of Indomalesia, s.t. very high (Shorea, ‘meranti’,
Dipterocarpus, ‘keruing’, Dryobalanops, ‘kapur’, etc.). No
lianas, no herbs.’
ROUX (Fig. 12.12).
MASSART and RAUH
Similar family:
exist as a continuum within
MALVACEAE when species.
petioles are distally bent or Very often the young tree
when hairs are fasciculate. conforms to ROUX’s
model (Fig. 12.11), then
its branches will become
erect (sequential
reiteration).
Dipterocarpaceae may have had a Gondwanan origin and

Fig. 12.10 Dipterocarpus obtusifolius, then migrated to the north on the Indian raft [2]. Apart from
leaf and fruits, Thailand
Dipterocarpoideae, a subfamily encompassing almost all the
Fig. 12.11 Dipterocarpus aromatica, onset of the sequential reiteration and of subcrown shyness, FRIM, Malaysia.
Dipterocarpus turbinatus, onset of the sequential reiteration, BG Sirikit, Thailand
species, the Monotoideae (Monotes and Marquesia) are endemic to Africa. Pakaraimaea and
Pseudomonotes (Pakaraimaeoideae, now included in CISTACEAE) are endemics in Colombian
Amazonia and in a Guianan mountain range.
Oleo-resins (dammar) are used for the preparation of varnishes (Dipterocarpus indicus, white
dammar from Vateria indica), waxes or oil painting.
! Trees often reiterate sequentially, i.e. after a critical developmental stage depending on the
species or environmental conditions, upper branches grow orthotropically and acquire the function of
trunks. In Fig. 12.12, the process is at a more advanced stage than in (Fig. 12.11). The ‘colonial’
character is still more visible when crown shyness occurs (or better said: subcrown shyness).
! Groups of different species of Dipterocarpaceae are often gregarious, a feature that has permitted
profitable, but not sustainable management. It is possible that ectomycorrhiza allow this group to be
dominant in forests [3].
! To quote LaFrankie [4] about a peculiar phenological trait of dipterocarps: “Dipterocarps exhibit
supra-annual masting and sequential flowering among sympatric congeneric species. By some
physiological trick not yet known, dipterocarps of a given species are able to synchronize their
flowering with a peak period that lasts about five to seven days, each species with a peak that is
separated by about a week, and each species with at least a day or so in which it is alone in
reproduction.”
! The world’s tallest known tropical tree is a Dipterocarp in Borneo, Shorea faguetiana [5]. This
discovery is the first 100-m tropical tree and the world’s tallest known flowering plant.
12.4 Dipterocarpaceae: 2-5-7-8-15-18 227
Fig. 12.12 Dipterocarpus costatus, crown shyness, Thailand. Shorea macrophylla, ROUX’s model, Malaysia
Fig. 12.13 Dipterocarpus obtusifolius, stipules hood-like, leaving annular scars, venation secant, Thailand.
Dipterocarpus sp., flush of young leaves, branches orthotropic, RAUH’s model, Thailand
Fig. 12.14 a Monotes africana, a solitary gland (gl), basal on the lamina. b Brochidodromous venation in
Dipterocarpus sp. c Petiole slightly pulvinate (p) at apex and submarginal vein, Cotylelobium, Dryobalanops
Fig. 12.15 Hopea sp., two-ranked leaf arrangement, petiole slightly pulvinate distally, BG Sirikit, Thailand. Shorea
henryana, stipule enclosing the apical bud, Thailand
Fig. 12.16 Hopea sp., camptodromous venation and domatia at vein axils, Thailand. Fasciculate hairs (Anisoptera,
Shorea), Thailand
References
2. Dayanandan, S., P.S. Ashton, S.M. Williams, and R.B. Primack. 1999. Phylogeny of the tropical tree family
Dipterocarpaceae based on nucleotide sequences of the chloroplast rbcL gene. American Journal of Botany 86:
1182–1190.
3. Torti, S.D., P.D. Coley, and D.P. Janosh. 1997. Vesicular-arbuscular mycorrhizae in two tropical monodominant
trees. Journal of tropical ecology 13: 623–629.
4. LaFrankie, J.V. 2010. Trees of Tropical Asia: An illustrated guide to diversity, 748 pp. Philippines: Black Tree
Publication Inc.
5. Gagen M. 2019. The world’s tallest known tropical tree has been found - and climbed [https://www.
nationalgeographic.com/environment/2019/04/worlds-tallest-tropical-tree-discovered-climbed-borneo; 12.10.2021].
Ebenaceae to Euphorbiaceae
13
13.1 Ebenaceae: 11-12-40
3 genera and 575 species, of which 550 species for Diospyros. Order Ericales. Plants without latex
and not aromatic. The phelloderm forms a black layer between dead bark and internal bark
(Fig. 13.3). Heart wood of several tree species becoming blackish because of an accumulation of
anthraquinones. Trunk with strong rhythmic growth bearing tiers of plagiotropic branches
(Fig. 13.3). Frequent apex abortions (Fig. 13.6b). Branches with two-ranked leaf arrangement
(Fig. 13.6a), rarely with spiral or opposite arrangement (Euclea divinorum). Stipules absent. Leaves
simple, entire. Venation not densely reticulate (Fig. 13.4). Glands scattered on the underside of
leaves (Fig. 13.2), often in vein axils (minute glandular trichomes in Euclea). Apices and buds with
appressed hairs (Fig. 13.5). Inflorescences lateral, pauciflorous. Flowers unisexual, regular, with 4
free petals and numerous stamens. Ovary superior (Diospyros, Euclea) or inferior (Lissocarpa, NW
AM-S). Fruits berries (Fig. 13.5), often 4-seeded, seeds relatively large.
Fig. 13.1 Euclea divinorum,

Botswana
https://doi.org/10.1007/978-3-031-05942-1_13
230 13 Ebenaceae to Euphorbiaceae
Trees or shrubs (D. verae-crucis, AM-C, D. lycioides, AF-S), s.t geoxylic (Euclea crispa, AF).
Pantropical distribution, with a majority of species in the Paleotropics, especially in Africa.
Fig. 13.2 Diospyros

malabarica, scattered glands,
Thailand

PHYLLANTHACEAE when Diospyros glands are
MASSART (Fig. 13.3 ): almost exclusive in
not very visible.
POLYGALACEAE: Xanthophyllum also bear Diospyros.
scattered glands on the underside of leaves. RAUH (Fig. 13.1): Euclea divinorum (AF-S).
Ebenaceae grow mainly in dense forests, but also in savannas where they have a more fissured or
cracked bark (Fig. 13.3) than in dense rainforests (Fig. 13.5). Diospyros trees provide very valuable
wood (ebony), used for carving or making tool handles. Several fruit trees (D. mollis ‘maplua’,
Thailand), but Diospyros kaki is extratropical (China, Japan). Many “non-wood products”, such as
leaves of D. melanoxylon for cigarette paper manufacturing (Sri Lanka), black dye (roots and bark of
Euclea divinorum), medicinal plants (bark of D. mespiliformis, African savannas).
! Diospyros lotus, a relic of the Tertiary Era in the Caucasian mountain range is naturalized in
many urban parks in Southern Europe.
! In Namibia many tribes use the wood of the magic gwarri (Fig. 13.1. Euclea divinorum) for the
entrance gate of their corrals in order to repel evil spirits [1].
Fig. 13.3 Diospyros buxifolia, MASSART’s model, Bali. Diospyros inconstans, anthraquinones in phelloderm,
Venezuela. Diospyros mespiliformis, bark fissured, Botswana
13.2 Elaeocarpaceae: 15-16-18-27-34 231
Fig. 13.4 Diospyros verae-crucis, berries, Shipstern, Belize. Diospyros sp., trunk bearing small leaves, plagiotropic
branches bearing large leaves, Thailand. Diospyros sp., venation not densely reticulate, Thailand
Fig. 13.5 Diospyros sp., fruits inserted on trunk, bark smooth, Bali Barat National Park. Diospyros sp., leaves two-
ranked, indumentum of appressed hairs, Thailand. Diospyros sp., berry with acrescent sepals, Imataca, Venezuela
Fig. 13.6 Diospyros spp. a. Trunk leaves are smaller than branch leaves (A: side view, B: top view). b. Branch
monopodial but abortion (+) of apical meristem of each growth unit (GU), axillary leaf (ax-L) of the next to last growth
unit (GU). c. leaf with scattered glands
13.2 Elaeocarpaceae: 15-16-18-27-34
12 genera and 550 species. Order Oxalidales. Plants without latex and not aromatic. Buttressed
trunks in Sloanea (Fig. 13.8). Inner bark showing s.t. a network of fibres. Several Elaeocarpus
species are deciduous. Stiff and elastic branches in Sloanea. Rhythmic growth (Fig. 13.9). Leaves
usually spiral (Elaeocarpus, Sloanea) or opposite (Aceratium, Aristotelia, Sericolea). Stipules
present or absent. Leaves simple, entire or toothed. Elaeocarpus leaves often turning red before
shedding (Fig. 13.7). Petiole pulvinate at both ends (Fig. 13.10 Sloanea). Young leaf-folding slightly
involute (Fig. 13.9). Venation pinnate (Fig. 13.10), densely reticulate; secant on leaf margin in
Dubouzetia. Indument of simple, not stellate, hairs. Inflorescences lateral (Fig. 13.9), racemose or
cymose. Flowers regular, with 4–5 free petals (Fig. 13.9), fringed in Aceratium and Elaeocarpus.
Ovary superior. Fruits prickly capsules (Fig. 13.10 Sloanea) or blue drupes (Fig. 13.7 Elaeocarpus
spp.). The family owes its name to the oily endosperm of the seeds.
Trees (Paleotrop except AF: Elaeocarpus,

Dubouzetia; NG: Sericolea; Pantrop: Sloanea) or
shrubs (Vallea, AM-S). Pantropical distribution. No
lianas and no herbs.
Fig. 13.7 Elaeocarpus angustifolia, leaf with-

ering red, and blue drupes, Seram, Indonesia
Similar families:
EUPHORBIACEAE, but quite supple twigs, and leaf-
folding not involute (Sloanea has stift branches).
MALVACEAE, but stellate hairs and leaf-folding not
involute. RAUH: Sloanea (mainly AM).
PHYLLANTHACEAE (possible confusion in Asia): as AUBRÉVILLE: Elaeocarpus (MA, AS, AU, OC).
for Elaeocarpus, AUBRÉVILLE’s model in Baccaurea. TROLL: Aceratium dasyphyllum (Moluccas).
A pantropical family, but best represented in the Southern Hemisphere, where it reaches the
fortieth parallel (Aristotelia, Chile, New Zealand). Sloanea with a curious distribution (AM, MA, AU,
NC). Sloanea woollsii, ‘yellow carabeen’, a big tree in Queensland. Vallea is native of the tropical
Andes. Ornamental trees (Elaeocarpus), several non-wood products: Elaeocarpus serratus, Ceylan
olive for preparing curry. Various fruit trees.
! Placed in Oxalidales near Cunoniaceae [2], this family probably originated in the Gondwananan
land mass [3].
! Withering leaves that turn red is not a feature of tropical trees, but is quite common in forest trees
of Northern Asia and America.
13.2 Elaeocarpaceae: 15-16-18-27-34 233
Fig. 13.8 Sloanea sp., buttressed trunk, Belize
Fig. 13.9 Elaeocarpus angustifolius. Corollas with fringed petals, Bali. Rhythmic growth, ramified system in between
orthotropy and plagiotropy by apposition. Young leaf-folding involute
Fig. 13.10 Sloanea sp., petiole pulvinate at both ends, pinnate venation, Belize. Sloanea massonii, sapling, rhythmic
growth, Guadeloupe. Sloanea sp., prickly capsules, French Guiana
13.3 Ericaceae: 12-13-20-29-30
113 genera and 4250 species. Order Ericales. Plants without latex and not aromatic. Rhytidome (i.e.
dead bark) sloughing off in scales (Fig. 13.12). Inner bark without chlorophyllous layer (Fig. 13.12).
Leaves usually spiral (Fig. 13.11), Agarista, Cavendishia, Rhododendron, pseudo two-ranked
(Fig. 13.15) or two-ranked at the extremities of the branches (Vaccinium). Leaves not stipulate.
Leaves simple, entire or toothed, coriaceous. Young leaves are often red or pinkish (Fig. 13.13)
before the onset of chlorophyll synthesis. Petioles with distinctive abscission zone (Fig. 13.12).
Venation often indistinct (Fig. 13.14). Young leaf-folding often revolute. Indument absent or rusty
(Fig. 13.15). Inflorescences racemose or cymose, usually lateral (Fig. 13.14), s.t. pseudoterminal
(Fig. 13.16) or truely terminal (Fig. 13.16). Flowers bisexual, corolla urceolate or tubular with 4–5
fused petals (Fig. 13.16). Stamens-10. Ovary inferior. Fruits a capsule (Fig. 13.14 Ericoideae) or
berry (Gaultheria, Vaccinium). The “ericoid” aspect is frequent under Mediterranean climate, as well
as in the Cape (South Africa), a region hosting more than six hundred Erica species.
Small trees (Rhododendron), numerous

shrubs, epiphytes (Fig. 13.17) and, at higher
latitudes, herbs, some saprophytic without
chlorophyll (Monotropeae). Ericaceae are myc-
orrhizal plants adapted to acid soils on sandstone
or granite, the majority of species are native of
mountain ranges and are quite infrequent in
lowland forests (e.g. Satyria).
Ericaceae are pantropical, diversified in
several subfamilies, such as Ericoideae (Bejaria,
Empetrum, Erica, Rhododendron), Vaccinoideae
(Agapetes, Agarista, Cavendishia, Dimorphan-
thera, Gaylussacia, Macleania, Vaccinium),
Monotropoideae (Pyrola, chlorophyll-less
Monotropeae) and Styphelioideae (mainly Aus-
tralian). Apart from some fruit shrubs (Vac-
cinium), Ericaceae are used above all as
ornamentals.
! Architectural analysis of Ericaceae is a
complicated matter: plants show frequent (and
genetically programmed) abortions of apical
meristems and models are not readily
Fig. 13.11 Typical: sympodial architecture, spiral identifiable.
arrangement of leaves
! One might wonder why Ericaceae are characterized by a complete absence of immediate
branching (branching is always delayed to the penultimate, or even older, GUs). This might be a
preadaptative trait that would explain their success in cold climates (tundra, Himalayan-alpine range,
Andes).
13.3 Ericaceae: 12-13-20-29-30 235
LEEUWENBERG: Gaultheria
reticulata (AM-S), Macleania
rupestris (Andes), Rhododendron
Similar families: aurigeranum (AS).
CYRILLACEAE are taxonomically SCARRONE: Rhododendron
very close to Ericaceae. ferrugineum (EU).
MANGENOT: Vaccinium
corymbosum (AM-N), Vaccinium
sprengelii (AS).
TROLL: probably absent .
Fig. 13.12 Agarista mexicana, Belize, typical: abscission zone (a–z) between stem and petiole, scaly rhytidome and
reddish inner bark, rhytidome sloughing off in small scales, rhy: young rhytidome, rho: old rhytidome, Belize
Fig. 13.13 Bejaria sp., a South American “rhododendron”, Gran Sabana, Venezuela. Cavendishia sp., flush of young
red leaves, leaves glabrous, Venezuela. cf. Vaccinium sp., racemose terminal inflorescence, Flores, Indonesia
Fig. 13.14 Bejaria sprucei, dehiscent capsules, Venezuela. Gaylussacia brasiliensis, terminal inflorescences,
coriaceous leaves, Santa Catarina, Brazil. cf. Notopora sp., venation indistinct, inflorescences lateral, Venezuela
Fig. 13.15 Notopora schomburgkii, sympodial architecture, branches plagiotropic with pseudo-distichous arrange-
ment of leaves. Rhododendron cf. zollingeri, underside of leaves covered by rust-coloured trichomes, Flores
Fig. 13.16 Rhododendron cf. zollingeri, terminal flowering, tubular corollas, Flores, Indonesia. Vaccinium sprengelii,
flowering lateral, but looking terminal, Thailand
13.4 Erythropalaceae: 3-12-13 237
Fig. 13.17 Satyria sp., epiphyte, cauliflory, La Escalera, Venezuela. Vaccinium euryanthum, inflorescences lateral,
truncate calyx, Gran Sabana, Venezuela
13.4 Erythropalaceae: 3-12-13
7 genera and 68 species. Order Santalales. Plants not aromatic Coloured sap uncommon, s.t. exudate
reddish (Fig. 13.20 Coula edulis) or not viscous white latex Heisteria). Patchy suberization of the
periderm (e.g. Heisteria). Small prehensile stems inserted in leaf axils (Erythropalum scandens).
Stem periderm remains green for a long period (Fig. 13.21), i.e. on several consecutive GUs. Young
internodes angular (Fig. 13.21). Leaves spiral (Erythropalum) or two-ranked (Heisteria, Strom-
bosia). Stipules absent. Leaves simple, entire, without glands. Petiole slightly pulvinate distally
(Fig. 13.20 Coula, Octoknema) or not (Heisteria). Young leaf-folding conduplicate (Fig. 13.20).
Venation pinnate, scalariform in Coula (Fig. 13.20). Indument usually absent, rarely stellate hairs
(Octoknema). Inflorescences lateral (Fig. 13.18). Sepals or petals 3, 5 or 6. Ovary superior. Fruits
drupaceous (Fig. 13.21) (Fig. 13.19).
Trees (Coula, AF; Octoknema, AF; Scorodocarpus, AS; Strombosia (AF, AS), shrubs (Heisteria,
AM, AF), lianas (Erythropalum scandens, AS), no herbs.
Pantropical, the majority of the species grows in dense paleotropical rainforests. Timber: Coula,
Scorodocarpus (bark and fruits of Scorodocarpus borneensis,‘kulim’, have a strong smell of garlic),
fruit trees (Coula edulis).

OLACACEAE: Ximenia conforms to ROUX. ROUX (Fig. 13.19 ): Octoknema sp., Strombosia sp. (AS).
SANTALACEAE: Okoubaka: MANGENOT. MANGENOT (Fig. 13.21 -b): Heisteria sp. (Guianas).
! The circumscription of the families is still confused. Some authors [4] place Strombosia in its
own family: Strombosiaceae.
Fig. 13.18 Heisteria sp., shrub, MANGENOT’s model, Fig. 13.19 Strombosia javanica, ROUX’s model, Kebun
flowers with persistent calyx, Guianas Raya, Java
Fig. 13.20 Coula edulis, bark with reddish exudate, young leaves pink-coloured and with conduplicate folding,
Cameroon. Coula, Octoknema, petiole slightly pulvinate distally (pu), venation scalariform
Fig. 13.21 a ROUX: e.g. Strombosia. b MANGENOT: e.g. Heisteria. Heisteria parviflora, leaves glabrous, French
Guiana. Heisteria sp., young internodes angular, pinnate venation, drupe, Cameroon
13.5 Erythroxylaceae: 8-32 239
13.5 Erythroxylaceae: 8-32
4 genera and 240 species. Order Malpighiales. Plants without latex and not aromatic. The family owes
its name to the red-coloured external wood (Fig. 13.22). Rhythmic growth and branching
(Fig. 13.24). Leaves two-ranked (opposite in Aneulophus). Intrapetiolar stipules (Fig. 13.24). Leaves
simple, entire, their underside pale green (Fig. 13.25). S.t. young leaves are involute and after
unfolding their lamina show false longitudinal veins (Fig. 13.25), but this feature is not always
present (Fig. 13.25). Venation pinnate, not densely reticulate. Stems and leaves glabrous.
Inflorescences lateral. Flowers small, with 5 free petals. Ovary superior. Fruits 1-stoned drupes
(Fig. 13.25) (Fig. 13.23).
An unmistakable family of trees (Erythroxylum macrophyllum, E. squamatum) or shrubs
(E. coca, E. havanense, etc.). No lianas and no herbs.
Pantropical, but best represented in tropical deciduous forests of the New World. Cocaine is
extracted from the leaves of Erythroxylum coca and E. novogranatense.
An unmistakable family of small

trees and shrubs.
Architectural models poorly

known
ROUX: Aneulophus sp. (AF).
MANGENOT: Erythroxylum
rotundifolium (AM-C).
TROLL, erect form of the model:
(1 Erytroxylum cf. rotundifoliuim,
AM-C), Erythroxylum longipes
(AM-S).
! Originally coca leaves were mixed with lime

or with burned Chenopodium quinoa plants. The
mixture was chewed to relieve hunger pains and
is still used by farmers in the high Andes.
Fig. 13.22 Erythroxylum sp., thick fissured bark and red

external wood, Belize
Fig. 13.23 Erythroxylum cf. rotundifolium, Belize
Fig. 13.24 Erythroxylum squamatum, erect form of a TROLL’s model, Guadeloupe. Erythroxylum sp., rhythmic
growth, scale leaves and intrapetiolar stipules, Venezuela
Fig. 13.25 Erythroxylum sp., false longitudinal veins, Belize. E. havanense, absence of false veins, inflorescences
lateral, Venezuela. E. novogranatense, false longitudinal veins, berries, Kebun Raya, Java
13.6 Euphorbiaceae + Peraceae … 241
13.6 Euphorbiaceae + Peraceae: 2-4-6-8-14-16-17-18-19-20-22-27-32-34-47-

53-55-A
Euphorbiaceae: 214 genera and 5600 species, Peraceae: 5 genera and 140 species. Order Malpighiales.
Latex (Fig. 13.36) present in many species (Excoecaria, Hevea, Sapium, etc.), s.t. a reddish exu-
date (Croton, Macaranga) but often no special sap (Acalypha). Plants rarely aromatic (Croton spp.).
Cactiform Euphorbia in Southern Africa, Madagascar and India (Fig. 13.26). Twigs supple, periderm
or young bark showing thin longitudinal slits (Fig. 13.26). Bark fibrous in some species (Fig. 13.31).
Spiral arrangement of leaves, s.t. two-ranked (Fig. 13.29 Chaetocarpus, Pera, PERACEAE) or
opposite-verticillate leaf arrangement (Fig. 13.35 Euphorbia sect. Chamaesyce, Euphorbia cotinifo-
lia) and, in some cases, associated with anisophylly (Mallotus). Stipules usually present (modified into
spines (Fig. 13.34) in succulent Euphorbia), rarely absent (Pera). Leaves simple, s.t. trilobate (Dale-
champia, Fig. 13.37 Jatropha), rarely trifoliolate (Hevea, Fig. 13.38 Macaranga spp.) or peltate
(Fig. 13.36 Homalanthus spp., Macaranga spp.). Young leaf-folding conduplicate (Fig. 13.33).
Leaves turning orange in Croton spp. (Fig. 13.31). Petiole often distally enlarged into a pulvinus
(Alchornea, Conceveiba, Fig. 13.40 Pausandra, Ptychopyxis, Spondianthus) or basally pulvinate
(Fig. 13.28 Agrostistachys). Frequently, laminas and petioles vary in size on the same GU. Glands
frequent on laminas or petioles (Fig. 13.38, Fig. 13.41) (Alchornea, Croton, Macaranga, Pausandra,
Sapium, etc.) or leaves without glands (the majority of the species, e.g. Acalypha, Euphorbia, etc.).
Venation pinnate, tripliveined or palmate. Main vein always protruding on the upper side of the lamina
(Fig. 13.32). Indument of simple (e.g. Acalypha) or stellate (e.g. Alchornea) hairs. Monoecy (i.e.
flowers unisexual, male and female on different parts of the inflorescence (Fig. 13.30) or on different
inflorescences (Fig. 13.31), plants never dioecious. Inflorescence often cymose, or a cyathium
(Fig. 13.35 Euphorbia). Flowers, if perianth present, with a radial symmetry (Fig. 13.42), rarely a
bilateral symmetry (Fig. 13.35), almost always reduced or medium-sized, with 5-n stamens. Ovary
superior, usually 2-, 3-, or 4-carpellate, numerous carpels in Hura. Fruits drupes: PERACEAE
(Fig. 13.43), capsules: EUPHORBIACEAE, (rarely a spiny capsule: Fig. 13.29 Chaetocarpus), s.t.
berries or samaras. Capsule usually trivalvate, or bivalvate (Fig. 13.28).
Fig. 13.26 Young bark with Fig. 13.27 Aleurites moluccana, ‘Kemiri’. Smooth trunk. Agroforest (detail:
thin longitudinal slits fissured bark), Flores, Indonesia
Trees (Alchornea, Aleurites, Antidesma, Hevea, Givotia, Hura, Ricinodendron. Shrubs (Acalypha,
Breynia, Croton, Euphorbia, Excoecaria, Jatropha, Macaranga, Mallotus). Lianas (Dalechampia,
Omphalea, Fig. 13.40 Plukenetia, Tragia). Herbs (Cnidoscolus, Euphorbia, Manihot, Ricinus).
Euphorbiaceae grow in most environments, except aquatic habitats, only one representative grows
in mangroves (Fig. 13.42 Shirakiopsis indica, AS). With their reduced flowers, plant are difficult to
identify when using keys or consulting herbaria. However, in the field, each genus has recognizable
architectural features. Uses numerous: natural rubber (Fig. 13.36 Hevea brasiliensis); starchy food,
vegetables (Fig. 13.39 Manihot esculenta of American origin is cultivated everywhere in tropics
(Fig. 13.39 ‘cassava’, ‘mandioca’ Brazil, ‘yuca’ Venezuela, ‘gari’ AF-W, ‘ubi kayu’ Malaya);
oleaginous seeds (Figs. 13.27 and 13.29 Aleurites moluccana), Ricinus communis (castor oil); per-
fumes (Croton spp.); beverages (in Mexico, ‘chaya’ (Fig. 13.30) is prepared with very urticant
Cnidoscolus plants!); medicinals (purgative seeds of Croton tiglium). Numerous ornamentals
(Codiaeum, Dalechampia, Euphorbia, Jatropha). Hippomane mancinella (‘mancenillier’), a small
tree of the Caribbean Coast, has a very vesicant and poisonous latex.
! Of all plant families, Euphorbiaceae show the most fantastic diversity of forms and architectures,
however ROUX’s model is absent, but present in Peraceae (e.g. Pera glabrata).

ACHARIACEAE: pulvinate petiole (e.g. CORNER: Agrostistachys gaudichaudii (shrub,
Ryparosa). Thailand), Trigonostemon wetriifolius (shrub,
CACTACEAE: cactiform Euphorbia can AS).
be distinguished from genuine Cactaceae LEEUWENBERG: Caryodendron orinocense
by their spines disposed in pairs, not (shrub, Peru), Euphorbia krameriana (shrub,
fasciculate (stipular origin). AF-S), Mallotus spp. (AS), Manihot esculenta
(herb, orig. Amazonia, now pantropical!).
DICHAPETALACEAE: leaves with
KORIBA: Balakata baccata (Thailand), Hura
glands.
crepitans (AM), Hippomane mancinella
PANDACEAE: Microdesmis, Panda (coastal small tree, WI).
conform to ROUX’s, a model still not PRÉVOST: Euphorbia pulcherrima (Mexico),
reported in Euphorbiaceae Excoecaria bicolor (shrub, mangrove, AS).
PHYLLANTHACEAE: Several ‘two- NOZERAN: Dichostemma glaucescens (AF),
ranked’ Euphorbiaceae (e.g. Mabea occidentalis (AM-C).
Chaetocarpus) look much like Antidesma. PETIT: Tetrorchidium didymostemon (AF).
SALICACAE (Casearia): leaves with SCARRONE: Croton matourensis (Guianas),
glands. Codiaeum variegatum (shrub, Polynesia).
VIOLACEAE (Rinorea): periderm RAUH : Elateriospermum tapos (Malaysia),
fissured. Euphorbia curtisii (MA), E. confinalis (South
Africa), Hevea brasiliensis (Amazonia).
PERACEAE (Pera conforms to ROUX’s
AUBRÉVILLE: Endospermum. malaccense
model): placed according to APG IV in a
(AS).
separate family.
ROUX: Pera glabrata (AM).
CHAMPAGNAT: Alchornea cordifolia (shrub,
AF) ; Homonoia riparia (rheophyte shrub, AS).
TROLL: Chaetocarpus schomburgkianus (AMS),
Croton fantzianus (AM-C), Sapium cornutum
(AF-W, Tannodia perrieri (MA).
Fig. 13.28 Actinostemon sp., shrub, inflorescence of reduced flowers, Venezuela. Agrostistachys gaudichaudii,
pulvinate petiolar bases, Thailand. Alchornea sp., bivalvate capsules, Venezuela
Fig. 13.29 Aleurites moluccana, leaves with palmate venation and sectioned seed above a fallen leaf, Flores,
Indonesia. Chaetocarpus schomburgkianus, two-ranked leaf arrangement, spiny fruits, Venezuela
Fig. 13.30 Cladogynos orientalis, underside of leaves whitish, Thailand. Cnidoscolus cf. multilobus, inflorescence
cymose, male flowers distal, fruit a basal capsule, Belize. Glass of Chaya (Cnidoscolus sp.), snack bar, Mexico
Fig. 13.31 Croton sp., leaves turning orange, Belize. Croton oblongifolius, bark fibrous, withered leaves orange-
coloured, Thailand. Croton sp., inflorescence cymose, with male and female flowers, Belize
Fig. 13.32 Croton sp., flowers with radial symmetry, Madagascar. Crotonogyne cf. mannii, main vein (V I)
protruding on upper side, Cameroon. Dalechampia sp., inflorescence with two foliaceous bracts, Belize
Fig. 13.33 Dalechampia sp., liana with lobate leaves. Dichostemma glaucescens, trunk apex parenchymatized, tier of
five plagiotropic branches, Cameroon. Elateriospermum tapos, conduplicate leaf-folding, Kebun Raya, Java
Fig. 13.34 Euphorbia bothae, cactus-like, stipules modified into spines. Euphorbia cotinifolia, leaves in whorls of
three. Euphorbia curtisii, chlorophyllous stems. (All three euphorbs: BG LC, France)
Fig. 13.35 Euphorbia (syn. Chamaesyce) articulata, a leaves opposite, Guadeloupe, b cyathium. Euphorbia
tithymaloides, flowers with bilateral symmetry, cult. Gymanthes lucida, base of lamina with two glands, Belize
Fig. 13.36 Hevea brasiliensis, harvesting latex, Kerala, India. Hippomane mancinella, latex white, Guadeloupe.
Homalanthus peltatus, leaves peltate, Flores, Indonesia
Fig. 13.37 Hura crepitans, male inflorescence, Venezuela. Jatropha curcas, red aqueous exudate, lobate leaves and
trilocular capsules, Bali, Indonesia
Fig. 13.38 Mabea sp., young bark with longitudinal slits, pinnate venation, French Guiana. Macaranga sp., foliaceous
stipules (st) and trifoliolate leaves. Macaranga sp., peltate leaf bearing two glands at the base of its main vein, Thailand
Fig. 13.39 Mallotus sp., shrub conforming to LEEUWENBERG’s model, Thailand. Manihot esculenta, in cultivation,
Flores, Indonesia. Omphalea sp., liana, French Guiana
Fig. 13.40 Pausandra sp., petioles distally pulvinate, lateral inflorescence bearing a 3-valvate capsule, Venezuela.
Plukenetia polyadenia, liana, fruits 4-carpellate, Venezuela. Ptychopyxis sp., rhythmic growth, leaves varying in size
Fig. 13.41 Ptychopyxis sp., rust-coloured indument, Sumatra. Sapium caribaeum, periderm with longitudinal slits,
Guadeloupe. Sapium sp., petioles bearing a pair of distal glands, Venezuela
Fig. 13.42 Sapium (syn. Shirakiopsis) indica, mangrove, Bali, Indonesia. Senefelderopsis croizatii, leaves glaucous
beneath, 3-valvate capsules, Sierra de Lema, Venezuela. Trigonostemon sp., floral parts in five, Kebun Raya, Java
Fig. 13.43 Pera glabrata, drupes, Southern Brazil. A typical Euphorbiaceae (e.g. Elateriospermum tapos) with latex,
rhythmic growth, leaves spiral, pulvinate petioles, and glands (gl)
References
1. Roodt, V. 1998. Trees and Shrubs of the Okavango Delta, 224 pp. Botswana: Shell Oil.
2. Savolainen, V., M.F. Fay, D.C. Albach, A. Backlund, M. van der Bank, K.M. Cameron, S.A. Johnson, M.D. Llédo,
J.C. Pintaud, M. Powell, M.C. Sheahan, D.E. Soltis, P.S. Soltis, P. Weston, W.M. Whitten, K.J. Wurdack, and M.W.
Chase. 2000. Phylogeny of the eudicots: A nearly complete familial analysis based on rbcL gene sequences. Kew
Bulletin 55: 257–309.
3. Raven, P.H., and D.L. Axelrod. 1974. Angiosperm biogeography and past continental movements. Annals of the
Missouri Botanical Garden 61: 539–673.
4. Nickrent, D.L., V. Malécot, R. Vidal-Russell, and J.P. Der. 2010. A revised classification of Santalales. Taxon 59:
538–558.
Fagaceae
14
14.1 Fagaceae: 8-9-18
7 genera and 970 species. Order Fagales. Plants without latex and not aromatic. When bark is smooth,
trunks exhibit ‘eye-marks’ (Fig. 14.4). External wood furrowed (Fig. 14.3) (bark of twigs at least
2 cm in diameter has to be peeled off). Spiral leaf arrangement (Lithocarpus, Quercus) or two-ranked
(Castanea, Fagus), very rarely opposite or whorled (Trigonobalanus). Stipules small (Fig. 14.3).
Leaves simple, entire, toothed or crenulate (Fig. 14.5), rarely lobate. Leaves without glands. Petiole
not pulvinate. Underside of lamina pubescent (s.t. whitish beneath Fig. 14.5), s.t. with small groups of
fasciculate hairs. Venation pinnate (Fig. 14.3), densely reticulate, the secondary veins often some-
Fig. 14.1 Quercus kingiana, dry Dipterocarp forest, Thailand
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250 14 Fagaceae
what parallel and secant on lamina margin (Fig. 14.3). Inflorescences of reduced flowers grouped in
catkins. Fruits drupes partially enclosed by a cupule (Fig. 14.4) (excrescence of the receptacle).
Trees (Castanea, Castanopsis, Lithocarpus, Quercus, e.g. Q. oleoides (Fig. 14.2), Q. kingiana
(Fig. 14.1) or shrubs (Quercus spp.). Pantropical distribution, no lianas and no herbs. Trunks of
Fagaceae in dense rainforests have a smooth bark whereas species in dry deciduous forests (Burma,
Laos, Northern Thailand, Yucatan, etc.) show a deeply fissured bark, this feature probably an
adaptation to periodic fires.

DIPTEROCARPACEAE (AS): fasciculate hairs.
RAUH: Quercus argentata, Q. kingiana
MALVACEAE: stellate hairs, but no furrowed external wood. (AS), Q. oleoides (AM-C).
TROLL: Fagus sylvatica (EU), Beech,
and Nothofagus (NOTHOFAGACEAE),
(NG, NZ, NC, AU-N, South temp. AM).
Fagaceae are pantropical, often existing in savannas or mountain ranges of Asia (Castanopsis,
Lithocarpus, Quercus) or Central America (Quercus). Trigonobalanus is a mysterious and archaic
genus of two little-known species, one in Colombia (T. excelsa), the other in Malaysia (T. verticil-
lata). Timber: Quercus spp. Fruit trees: the chestnut tree (Castanea sativa), as well as several
Quercus spp. produce the sweet acorns of the Mediterranean region.
Fig. 14.2 Quercus oleoides, semideciduous forest, Belize

14.1 Fagaceae: 8-9-18 251
Fig. 14.3 Castanopsis armata, persistent stipules, leaves entire, pinnate venation, Thailand. Fagus sylvatica, external
wood furrowed, Switzerland. Lithocarpus lindleyanus, embossed and secant venation, Thailand
Fig. 14.4 Lithocarpus sundaicus, various cupules from the same tree, bracteoles have fused into a spiral or two annular
scars, Kebun Raya. Lithocarpus sundaicus, group of cupules, some with spiral scar turning left, some turning right and
some in concentric circles, Kebun Raya, Java. Lithocarpus sp., ‘eye-mark’ developing after branch shedding, Sumatra
Fig. 14.5 Quercus kerrii, leaves crenulate, Thailand. Quercus sp., bark fissured, Belize. Quercus cf. oleoides, leaves
entire, their underside pubescent whitish, Belize
! NOTHOFAGACEAE, a small family close to the Fagaceae, does not occur in the lowland tropics
(S-AM-S, New Guinea, New Caledonia, New Zealand), the family probably evolved from breakup of
a world-wide Fagales ancestor [1].
! One might wonder why the lobate leaf form of oaks is very rare in the tropics. In Belize, lat. 16–
18 N., among ten species of native oaks, only Quercus segoviensis has deeply crenulate … or weakly
252 14 Fagaceae
lobate leaves. If one considers the oaks native to NE America (about 30 species), it stands out that
northern or inland species have lobate leaves and entire-leaved species grow in the warm temperate or
subtropical part of the U.S.A. (e.g. Quercus chapmanii, Q. incana, Q. laurifolia) [2].
References
1. Harley, R. 1999. Lamiaceae. In Flora of the Venezuelan Guyana, vol. 5. 834 pp. St. Louis (Missouri): ed.
J.A. Steyermark, P.E. Berry, K. Yatskievych, and B.K. Holst.
2. Little E.L. 1992. Field Guide to North American Trees—Eastern Region, 716 pp. New York: Alfred A. Knopf.
Gentianaceae to Goupiaceae
15
15.1 Gentianaceae: 37-38
95 genera and 1850 species. Order Gentianales. Plants without latex and leaves not aromatic.
Opposite arrangement of leaves. Stipules missing but a thin interpetiolar ridge brings together the
petiolar bases, s.t. forming a short collar (Fig. 15.7). Leaves simple, entire, s.t. very large (Antho-
cleista), adorned with a pair of basal secondary veins running to leaf apex (Fig. 15.8). Leaves without
glands, glabrous. Inflorescences cymose (Fig. 15.6) or solitary flowers (Fig. 15.8), lateral
(Fig. 15.5a–c) or terminal (Fig. 15.5b). Corollas contortate, often showy (Figs. 15.1 and 15.8),
tubular at their base (Fig. 15.1), petals and stamens 5, s.t. 4. Ovary superior, 2-carpellate. Fruits
capsules, rarely berries.
Fig. 15.1 Lisianthus axillaris, corolla red and green, Belize
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254 15 Gentianaceae to Goupiaceae
Fig. 15.2 FAGERLIND’s or Fig. 15.3 ROUX’s model,

AUBREVILLE’s model, e.g. e.g. Tachia guianensis
Fagraea sp.
Trees (Fig. 15.2 Anthocleista, Fagraea). Shrubs (Potalia,

Symbolanthus, Tachia). Lianas (Chorisepalum, Guianas). Herbs
(Coutoubea, Gentiana, Irlbachia, Lisianthus). Voyria (Fig. 15.8)
and Voyriella are small mycotrophic herbs in the understorey of
rainforests of tropical AM.
In spite of the bulk of Gentiana species occurring in Alpine and
Himalayan ranges, the family is a group of Gondwanan origin
(numerous endemic genera in the Guianas, e.g. Chorisepalum,
Irlbachia, Saccifolium being basal), [1]. Besides some valuable
timbers (Fagraea spp., ‘tembusu’), the main species of economic
importance are ornamentals and medicinal plants (bark of
Anthocleista).
! Herbs of tribe Gentianeae synthesize bitter substances which
are of great interest to the pharmaceutical industry. Fig. 15.4 STONE model’s, e.g.
Coutoubea sp.
Similar families: CHAMBERLAIN: Potalia amara (AM).
LOGANIACEAE and LEEUWENBERG: Anthocleista nobilis (AF), Symbolanthus
RUBIACEAE also have stems elisabethae (N AM-S).
with an interpetiolar ridge, but SCARRONE: Anthocleista amplexicaulis (MA).
are stipulate.
FAGERLIND Fig. III.7.2: Fagraea crenulata (AS).
Several genera of the former
AUBRÉVILLE: Fagraea fragrans (AS).
Loganiaceae have been
transferred to Gentianaceae, e.g. STONE Fig. III.7.4: Coutoubea ramosa (AM).
Fagraea, Potalia, Anthocleista. ROUX Fig. III.7.3: Lisianthus axillaris (AM-C), Tachia guianensis
(AM-S).
15.1 Gentianaceae: 37-38 255
! The leaves of alpine gentians have a life span lasting more than 12 months, hence leaves remain
green even under snow cover (Fig. 15.6). This evergreen feature is reminiscent of the tropical
behaviour of the Gentianaceae, a family which does not include any deciduous species.
Fig. 15.5 a Chorisepalum nemorosum, shrub, terminal flowering. b Enicostema verticillatum, herb, lateral flowering,
Venezuela. c Irlbachia nemorosa, herb, terminal flowering, Venezuela
Fig. 15.6 a Stem of Fagraea sp., branch plagiotropic by apposition. b–d Various kinds of petiolar bases, b, c petiolar
spurs (b Fagraea spp.), (c Anthocleista, Fagraea). d Auriculate appendages (Anthocleista, Fagraea). Gentiana acaulis
in spring, leaves of the basal rosette remain green after months spent under snow, Swiss Alps. Irlbachia sp., herb
bearing a cymose inflorescence, Venezuela
Fig. 15.7 Irlbachia nemorosa, herb, terminal flowering, Venezuela. Lisianthus axillaris, annular stipuloid organ,
Belize. Potalia amara, cymose terminal inflorescence, French Guiana
Fig. 15.8 Symbolanthus elisabethae, corolla contorted, Venezuelan Guiana. Tachia schomburgkiana, typical venation
and axillary flowers, Guianas. Voyria rosea, mycotrophic, French Guiana
15.2 Gnetaceae: 36-39-53-56
Mongeneric family of 30 species (Gnetum). Order Gnetales, Gymnosperms. Unlike other gym-
nosperms, they possess vessel elements in the xylem. Trees and lianas. Latex in several species
(Gnetum urens), torn leaves with laticiferous threads (Fig. 15.11). Bark with sclerenchymatous
inclusions (Fig. 15.10). Stems enlarged at nodes (Fig. 15.10). Opposite leaf arrangement. Venation
reticulate without blind veinlets (Fig. 15.9a), i.e. all veinlets form closed loops, unlike typical ‘Dicot’
venation with blind veinlets (Fig. 15.9b). Inflorescences in spikes of reduced ‘flowers’ arranged in
whorls (Fig. 15.11). Small trees and lianas (Fig. 15.10).
Fig. 15.9 Gnetum sp. a venation without blind veinlets. b Blind veinlets in a Dicotyledon
15.2 Gnetaceae: 36-39-53-56 257
Fig. 15.10 Gnetum montanum, sclerenchymatous inclusions, Chiang Mai, Thailand. Gnetum urens, reiterating liana,
swollen nodes, leaves opposite and, unusual: two bound trunks, French Guiana
Fig. 15.11 Gnetum montanum, torn leaf showing laticiferous threads, Chiang Mai, Thailand. Gnetum sp., spike of
whorled flowers, Seram, Indonesia. Crackers of Gnetum gnemon, Sumatra

Frequent confusions with opposite-leaved Dicots (Apocynaceae, etc.). ROUX: Gnetum spp.
Uses: young leaves an edible vegetable and seeds of Gnetum gnemon (‘melinjo’) used in Java to
prepare fried chips (Fig. 15.11).
! The prevalence of entomophily in the Gnetales suggest that unspecialized insect pollination
originated before the divergence between Gnetales and Angiosperms [2].
15.3 Goodeniaceae: 24-29
10 genera and 400 species. Order Asterales. Nearly half of the species belong to the almost entirely
extratropical genus Goodenia (all through Australia). Plants sappy. Spiral arrangement of shortly
petiolate leaves (Fig. 15.12). Stipules absent. Flowers with a bilateral symmetry looking like those of
Lobelia (Campanulaceae). Fruits capsules, drupes or nuts.
Scaevola plumieri is a shrub of Indo-Atlantic tropical coasts, S. sericea is similar, but Indo-Pacific
in distribution. Both species conform to AUBRÉVILLE’s model. In these regions, possible confusion
with Corynocarpus (Corynocarpaceae), which also bear entire, ovale and fleshy leaves, but
Corynocarpus leaves are distinctly petiolate.
! Seeds of Scaveola are viable in seawater for long periods, but need fresh water for their
germination [3].
! Scaevola verticillata (Goodeniaceae) is endemic to the summit of the ultramafic Mount
Tambukon in Sabah [4].
Fig. 15.12 Scaevola sp., shrub in back-shore with fleshy leaves, Seram, Indonesia
15.4 Goupiaceae: 9
Monogeneric family of 3 species. Order Malpighiales. NE AM-S. Sympodial architecture

(Fig. 15.13), i.e., trunk consisting of a stacking of modules. The sapling conforms to ROUX’s model
(Fig. 15.13), then the adult tree looks like a LEEUWENBERG’s model, however, the many
peripherical younger parts conform again to ROUX. Stipules small. Two-ranked leaf arrangement.
Leaves simple, toothed. Inflorescences umbellate, lateral (Fig. 15.14). Flowers small, with a disk.
Fruits drupes.
! Goupia glabra, named ‘goupi’ in French Guiana, has uncertain affinities with other Malpighiales
but is a well-known tree of the sylvatic mosaic in NE South America.
References 259
Similar families:
PHYLLANTHACEAE, but leaves entire.
ROUX: Goupia glabra.
Fig. 15.13 Goupia glabra, architectural metamorphosis
Fig. 15.14 Goupia glabra. continuous branching, ROUX’s model, Venezuela. Umbellate lateral infructescences
References
1. Yuan, Y.-M., S. Wohlhauser, M. Möller, P. Chassot, G. Mansion, J. Grant, P. Küpfer, and J. Klackenberg. 2003.
Monophyly and relationships of the tribe Exaceae (Gentianaceae) inferred from nuclear ribosomal and chloroplast
DNA sequences. Molecular Phylogenetics and Evolution 28: 500–517.
2. Kato, M., T. Inoue, and T. Nagamitsu. 1995. Pollination biology of Gnetum (Gnetaceae) in a lowland mixed
Dipterocarp forest in Sarawak. American Journal of Botany 82: 862–868.
3. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3rd ed., 1022 pp. Cambridge: Cambridge University Press.
4. Galey, M.L., van der Ent, A., Iqbal, M.C.M., and Rajakaruna, N. 2017. Ultramafic geoecology of South and
Southeast Asia. Botanical Studies 58: 18.
Hamamelidaceae to Humiriaceae
16
16.1 Hamamelidaceae: 5-6-8-19-29 (incl. Altingiaceae)
28 genera and 100 species. Order Saxifragales. Strong rhythmic growth and branching in Altingia
(Fig. 16.3) and Liquidambar. Arrangement of leaves spiral (Fig. 16.3 Altingia, Fig. 16.4 Rhodoleia)
or two-ranked (Fig. 16.4 Exbucklandia, Maingaya). Stipules present, partly absent in Altingia and
Rhodoleia. Annular stipular scars after leaf shedding (Fig. 16.1) in Maingaya. Petiole slightly
geniculate distally (Fig. 16.4 Rhodoleia). Venation pinnate (Hamamelis, Maingaya) or tripliveined
(Fig. 16.4 Exbucklandia, Liquidambar, Matudaea, the latter genus looking like Celtis spp.). Flowers
usually arranged in spikes or in heads (Fig. 16.4 Altingia, Liquidambar, Fig. 16.2 Parrotia) mim-
icking solitary flowers, rarely showy (Fig. 16.4 Rhodoleia). Fruits woody capsules, s.t. arranged in
heads (Fig. 16.3).
Fig. 16.1 Maingaya malayana, foliaceous stipule enclos- Fig. 16.2 Parrotia persica, male flowers practically
ing a young leaf, the stipule leaving an annular scar, FRIM, reduced to groups of stamens, cultivated tree
Malaysia
Trees (Altingia, Liquidambar, Rhodoleia), shrubs (Corylopsis, Hamamelis). A rather heteroge-

neous group, centred in extratropical regions of the N-Hemisphere (Corylopsis, Hamamelis, Parrotia,
etc.) and also in tropical rainforests where they are represented by other small endemic genera (AS:
Altingia, Exbucklandia, Maingaya, Rhodoleia; AM-C: Matudaea, MA: Dicoryphe). Timber: Altingia
excelsa reaches a height of up to 60 m. Pharmaceutical plants: gum-resin of Liquidambar styraciflua.
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262 16 Hamamelidaceae to Humiriaceae
Hamamelidaceae probably originated in the Northern Hemisphere and in tropical Asia during the
mid Cretaceous [1, 2]. Placed near Saxifragaceae, paeonies and gooseberry shrubs, Hamamelidaceae
are part of a sundry group which that continues to puzzle molecular biologists. Hamamelidaceae
might be one of the earliest extant families endowed with tension wood and conforming to TROLL’s
model (see also Annonaceae).
Fig. 16.3 Altingia excelsa, from herbarium specimens, CBG H. Crenulate leaves and lateral heads of capsules.
Rhythmic growth. Scaly apical bud
Fig. 16.4 Exbucklandia populnea, ramified stem, lobate leaves, stipules (S) spatulate; A1-A2-A3: axes of order 1, 2
and 3, Sumatra. Rhodoleia championii, Sumatra, petiole slightly geniculate distally and showy flowers
16.2 Hernandiaceae: 20-55-57
4 genera and 60 species. Order Laurales. Scattered oil-cells that may unite as cavities. Discontinuous
pith in Sparattanthelium (Fig. 16.7). Spiral leaf arrangement. Leaves simple, entire or palmatilobate
(Fig. 16.6), or leaves trifoliolate (Illigera). Venation palmate or leaves tripliveined. Inflorescences
cymose. Flowers small, regular. Fruits dry, indehiscent, winged (Fig. 16.6 Gyrocarpus, Illigera) or
enclosed in an involucre (Fig. 16.5 Hernandia).
16.2 Hernandiaceae: 20-55-57 263
Shrubs (Hernandia), lianas with pre-

hensile twigs (Fig. 16.7) (Sparattanthelium)
or prehensile petioles (Fig. 16.6 Illigera).
Hernandia has a circumpacific distribution,
in addition Hernandia sonora (Australasia)
and H. guianensis (coastal tropical America)
are very much alike because of their peltate
leaves.
! Fruits of Hernandia sonora look like
small open lanterns, which «sing» when the
wind blows through them, hence the name
‘sonora’.
Fig. 16.5 Hernandia sonora, fruits enclosed by bracts,

Indonesia
Similar families:
Leaves of Hernandia and Gyrocarpus resembles those of MALVACEAE.
Fig. 16.6 a Gyrocarpus sp., leaves palmately lobate, winged drupes. b Hernandia sp., rhythmic growth, leaves
slightly peltate, tripliveined. c. Illigera sp., prehensile petiole
Fig. 16.7 Sparattanthelium tupiniquinorum, liana, Imataca, Venezuela. Sparattanthelium sp., Venezuela. Short shoots
modified into hooks, scalariform venation, leaves tripliveined and discontinuous pith
16.3 Huaceae: 8
A very small families of 2 genera and 3, all African, species. Order Oxalidales. Trees and shrubs with
smell of garlic. Two-ranked leaf arrangement (Fig. 16.9). Stipules present. Stems and leaves pub-
escent (Fig. 16.8), stellate hairs (Afrostyrax), hairs simple (Hua). Venation pinnate, brochidodromous
(Hua) or camptodromous (Afrostyrax). Flowering lateral, flowers small, 5 petals, many short stamens
and stipitate hairy stigmas (Fig. 16.9 Hua). Fruits indehiscent or dehiscent capsules.
Similar families:
MALVACEAE: stellate hairs as Architectural models:
in Afrostyrax, but no smell of MANGENOT: Afrostyrax
garlic. gabonensis (AF-W).
SALICACEAE: small, lateral
flowers as in Hua, but hyaline
margin of leaves present.
Fig. 16.8 Afrostyrax gabonensis, two-ranked leaf arrangement, stem and young leaves pubescent, Cameroond
16.4 Humiriaceae: 9-13-22-30 265
Fig. 16.9 Hua gabonii. Flower with hairy staminodia and plagiotropic branch, leaves two-ranked and flowers
fasciculate, drawn from CBG H specimens
16.4 Humiriaceae: 9-13-22-30
8 genera and 50 species. Order Malpighiales. Trees and shrubs. Rhythmic growth. Spiral arrange-
ment of leaves (Fig. 16.10 Humiria balsamifera) or two-ranked (Fig. 16.11 Humiriastrum,
Sacoglottis, Vantanea). Stipules small or missing. Leaves simple, coriaceous, entire (Humiria) or
slightly crenulate (Humiriastrum, Fig. 16.11 Sacoglottis). Petiole short, its base enlarged
(Fig. 16.11). Leaves glabrous, rarely their underside pubescent (Humiria spp.). Unconspicuous
laminar glands (Humiria, Sacoglottis, Vantanea). Venation pinnate, distinctly reticulate (Fig. 16.11).
Flowers with 5 sepals and 5 free petals, stamens numerous. Fruits drupes containing a plurilocular
embossed stone enclosing resinous cavities (Fig. 16.11), an adaptation for water dispersal.
Fig. 16.10 Humiria balsamifera, Gran Sabana, Venezuela

Fig. 16.11 a Humiriastrum sp., young tree, MANGENOT’s model, frequent meristem abortions (x), Venezuela.
b Humiria sp., short petiole with an enlarged base, internode angular. c Endocarps or their sections. I: Sacoglottis and
Schizostemon. II: Sacoglottis amazonica. III: Sacoglottis trichosperma. Resinous cavities in II–III (redrawn from [5]).
d Sacoglottis gabonensis, Campo forest, Cameroon

IXONANTHACEAE has leaves distinctly toothed. RAUH or SCARRONE: Humiria (AM).
Bonnetia (CLUSIACEAE s.l.) have an indistinct MANGENOT (Fig. 16.11-a): Humiriastrum spp.
venation. (AM).
l b ( ) ( )
Humiriaceae are essentially American, except for a single African species: Sacoglottis gabonensis
(‘ozouga’ in Gabon and with as many names as there are languages in W-Africa). Timber: Humiria
floribunda (Guianas). Tall tree (up to 65 m high): Vantanea barbourii (WI).
Extrafloral nectaries are unexpected organs in this family, formerly placed in Linales [3]. However
description of glands is well documented by [4].
References
1. Xiang, X., K. Xiang, R. Del Ortiz, F. Jabbour, and W. Wang. 2019. Integrating palaeontological and molecular data
uncovers multiple ancient and recent dispersals in the pantropical Hamamelidaceae. Journal of Biogeography 46:
2622–2631.
2. Xie, L., T.S. Yi, R. Li, D.Z. Li, and J. Wen. 2010. Evolution and biogeographic diversification of the witch-hazel
genus (Hamamelis L., Hamamelidaceae) in the Northern Hemisphere. Molecular Phylogenetics and Evolution 562:
675–689.
3. Cronquist, A. 1981. An Integrated System of Classification of Flowering plants, 1262 pp. Columbia University
Press.
References 267
4. Wurdack, K.J., and C.E. Zartman. 2019. Insights on the systematics and morphology of Humiriaceae
(Malpighiales): Androecial and extrafloral nectary variation, two new combinations, and a new Sacoglottis from
Guyana. Phytokeys 124: 87–121.
5. Herrera, F., S.R. Manchester, C. Jaramillo, B. Macfadden, and S.A. da Silva-Caminha. 2010. Phytogeographic
history and phylogeny of the Humiriaceae. International Journal of plant sciences 171: 392–408.
Icacinaceae s.l. to Irvingiaceae
17
17.1 Icacinaceae s.l. (Icacinaceae s.str. I + Metteniusaceae

M + Stemonuraceae S) + Cardiopteridaceae C: 12-13-25-30-44-54-56
A group of four families encompassing 55 genera and 377 species, but placed in different orders.
Plants without latex and leaves not aromatic. Cross-section of lianas with xylem in blocks
(Fig. 17.1). Sclerenchymatic inclusions in bark (Fig. 17.6). Stems with patchy suberization (e.g.
Leptaulus Fig. 17.6). Usually two-ranked leaf arrangement (I: Dendrobangia, Emmotum, S: Dis-
cophora, Gomphandra, C: Gonocaryum, Citronella?), s.t. spiral (M: Apodytes, I: Desmostachys),
rarely opposite (I: Iodes). Stipules missing. Leaves simple, entire, usually shiny (Fig. 17.2) and
coriaceous, without glands. Venation distinctly reticulate and pinnate (palmate in Phytocrene),
camptodromus (Fig. 17.4 Emmotum, Gonocaryum) or brochidodromus (Fig. 17.5 Lasianthera, S?).
Plants glabrous at first sight but buds and apex often pubescent (Poraqueiba), plants entirely glabrous
in Sarcostigma. Inflorescences cymes (Fig. 17.3) or spikes (Fig. 17.3). Flowers small, their petals
free. Ovary superior. Fruits drupes, rarely samaras.
Fig. 17.2 Gomphandra quadrifida, a typical Icacinaceae

with shiny leaves and angular stems with a green periderm,
Malaysia
Fig. 17.1 Iodes africana, xylem in blocks, Cameroon
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270 17 Icacinaceae s.l. to Irvingiaceae
Trees (Citronella C, Dendrobangia M, Emmotum M, Gomphandra S, Nothapodytes I,

Poraqueiba I), geoxylic shrubs (Casimirella, Fig. 17.5 geoxylic Pyrenacantha, I). Lianas with
twining stems or climbing with prehensile petioles (Cardiopteris C, Desmostachys I, Phytocrene I,
Pyrenacantha I, Sarcostigma I, paléo), hooks (Desmostachys I, Pyrenacantha, I) or leaning on their
support (Leretia I, AM, Rhaphiostylis M). Iodes africana I bears terminal twining tendrils becoming
strongly woody. No herbs are reported for this group of families.
ROUX (Fig. 17.6-d): Dendrobangia M (AM), Discophora S (AM-S),
Medusanthera S (AS-OC), Stemonurus S (AS).
MASSART-NOZERAN: Citronella brassii M (NG).
Similar families: PRÉVOST: Lasianthera S (AF), Nothapodytes? I (AS).
ERYTHROPALACEAE have a venation PETIT (Fig. 17.6-b): Citronella C (AS), Gomphandra S (AS-OC),
not densely reticulate. Leptaulus C (AF-MA).
AQUIFOLIACEAE, but stipulate. ATTIMS: Apodytes I (New Caledonia).
TROLL: Alsodeiopsis I (AF).
MANGENOT (Fig. 17.6-a): Desmostachys I (AF-MA), Rhaphiostylis
M (AF), Rhyticaryum I (AS-OC), Leptaulus C (AF-MA).
Fig. 17.3 Dendrobangia boliviana, ROUX’s model (French Guiana) and splintered wood (Venezuela). Desmostachys
sp., inflorescences in spikes, leaves two-ranked, Madagascar
Fig. 17.4 Emmotum fulvum, two-ranked leaf arrangement, venation camptodromous, Venezuela. Gonocaryum
littorale, two-ranked leaf arrangement, BG Singapore. Iodes africana, leaves opposite and shiny, Madagascar
17.2 Irvingiaceae: 5 271
Fig. 17.5 Lasianthera africana, venation brochidodromous, inflorescence an umbel of cymes, Cameroon.
Pyrenacantha malvifolia, geoxylic shrub, (orig. AF) BG LC, France
Fig. 17.6 Leptaulus citriodora, stem with patchy suberization, Madagascar. Various Icacinaceae s.l.;
a MANGENOT’s model (Rhyticaryum); b PETIT (Leptaulus); c Iodes africana, modular construction; d plagiotropy
with two-ranked leaf arrangement (Dendrobangia, Discophora, Emmotum); e sclerenchymatic inclusions in bark
(Gonocaryum, Iodes); f Emmotum sp., peculiar venation
Icacinaceae and closely related families have puzzled molecular biologists. The group is archi-
tecturally highly diversified and encompasses overlooked species of forests and savannas. Sar-
costigma kleinii in ayurvedic, very large tubers Casimirella ampla (I, AM) are cooked to eliminate
toxic substances, and seeds of Poraqueiba are used for cooking oil.
! In understorey, Icacinaceae are not easy to spot because they are neither gregarious nor pioneers.
Attempts to find and identify plants to the correct family may lead botanists to reconsider the
profession!
17.2 Irvingiaceae: 5
3 genera and 10 species. Order Malpighiales. Rhythmic growth (Fig. 17.8) and branching. Two-
ranked arrangement of leaves (Fig. 17.8). Stipules narrow, hood-like, enclosing the apical bud.
Leaves entire, ovate lanceolate. Indument absent (leaves glabrous). Young leaf-folding involute.
Inflorescences paniculate (Fig. 17.9-b). Flowers with an intrastaminal disk (Fig. 17.9). Fruits drupes
(Fig. 17.8) or samaras.
Similar families:
Moraceae when bearing Architectural models:
long and narrow stipules. TROLL: Irvingia malayana.
Ulmaceae: however with
pubescent leaves.
272 17 Icacinaceae s.l. to Irvingiaceae
A small Old World family. Desbordesia glaucescens (Fig. 17.7) and Irvingia malayana (‘pauh
kijang’) are large buttressed trees of W-AF and, South Eastern Asia, respectively.
Irvingia malayana is a lofty tree with large buttresses. Irvingia gabonensis has scented fruits and
its oily seeds are used to prepare Gabon’s ‘chocolate’.
Fig. 17.7 Desbordesia glaucescens, flushes of young leaves, Cameroon
Fig. 17.8 Irvingia malayana, two-ranked arrangement of leaves, Sakaerat, Thailand. Irvingia smithii, rhythmic
growth, buds of young leaves, Cameroon
17.2 Irvingiaceae: 5 273
Fig. 17.9 Irvingia gabonesis, sectioned drupe (a) and axillary paniculate inflorescence (b), (CBG H specimen).
c. Irvingia cf. smithii, intrastaminal disk, Cameroon
Juglandaceae
18
18.1 Juglandaceae: 48
8 genera and 62 species. Order Fagales. Inner bark fibrous in Engelhardia (Fig. 18.3). Septate pith
(Fig. 18.3) in Juglans (AS-N, AM-N) and Pterocarya (AM-N). Rhythmic growth. Leaves compound
pinnate (Fig. 18.3), Stipules absent. Inflorescences in catkins (Fig. 18.1). Flowers much reduced,
usually unisexual. Fruits drupes (Fig. 18.2) or samaras.
Fig. 18.1 Engelhardia spicata, catkins, Northern Thailand
A group mainly temperate in its distribution, tropics: Engelhardia (AS), Oreomunnea (AM),
Juglans boliviana, (AM-S), Juglans neotropica (AM-C & S).
Juglandaceae originated in Europe and North America [1], then underwent migration to South
America (Andes) and South Eastern continental Asia (Thailand, Malaysia).
https://doi.org/10.1007/978-3-031-05942-1_18
276 18 Juglandaceae
Fig. 18.2 Carya alba, dehiscent drupe, in cultivation, orig. AM-N. Juglans neotropica, terminal infructescence of
drupes (AM-C and S)
Fig. 18.3 Engelhardia spicata, Chiang Mai, Thailand. Leaves pinnate and fibrous bark. Juglans regia, septate pith,
Switzerland
! Fertilization is delayed after pollination, as in all of the order Fagales [2]. This characteristic may
be linked to the fact that this order contains no herbaceous plants which need to mature and multiply
rapidly after pollination in order to be competitive.
References
1. Song, Y.-G., Y. Fragnière, H.-H. Meng, Y. Li, S. Bétrisey, A. Corrales, S. Manchester, M. Deng, A.K. Jasińska, H.
V. Sâm, and G. Kozlowski. 2020. Global biogeographic synthesis and priority conservation regions of the relict tree
family Juglandaceae. Journal of biogeography 47: 653–657.
2. Sogo, A., and H. Tobe. 2006. Delayed fertilization and pollen-tube growth in pistils of Fagus japonica (Fagaceae).
American Journal of Botany 93: 1748–1756.
Lamiaceae to Lythraceae
19
19.1 Lamiaceae: 35-38-39-43-44-45-51-52-56
240 genera and 6700 species. Order Lamiales. Plants without latex and, on a tropical scale, leaves not
very often aromatic like those of shrubs and herbs of the Mediterranean region. Bark often thin or
papyraceous or multilayered (Fig. 19.5). Stems usually not lenticellate, quadrangular (Fig. 19.6).
Lianas are twining or lean on their support, or climb by means of petiolar bases modified in hooks
(Fig. 19.4). Internodes quadrangular (Fig. 19.1). Opposite leaf arrangement, exceptionally spiral
(Fig. 19.2 Amasonia, AM). Stipules missing. Leaves simple, entire or toothed, s.t. cordate (Fig. 19.5
Tectona), not coriaceous, compound pinnate in Peronema (AS) or palmate (Fig. 19.6) in Vitex
(Pantrop). Glands s.t. at lamina bases (Aegiphila, Fig. 19.4 Gmelina). Indument of simple hairs,
surprisingly of stellate hairs in Callicarpa and Tectona, these genera formerly placed in Verbenaceae
[1]. Inflorescences in cymes (Fig. 19.3) or in heads (Fig. 19.5), these s.t. surrounded by coloured
bracts (Fig. 19.3). Flowers often bilabiate, more rarely regular, calyx tubular, persistent in Clero-
dendrum (Fig. 19.3) and Karomia. Ovary superior. Fruits drupes (s.t. blue in Clerodendrum
Fig. 19.3) enclosing one stone or consisting of 4 druplets.
Fig. 19.1 Aegiphila integrifolia, stems quadrangular, Venezuela
https://doi.org/10.1007/978-3-031-05942-1_19
278 19 Lamiaceae to Lythraceae
Fig. 19.2 Amasonia campestris, French Guiana
Trees (Gmelina, Premna, Tectona, Vitex), numerous shrubs (Hyptis, Vitex) and even more
extratropical herbs (Hyptis, Ocimum, Plectranthus, Amasonia campestris (Fig. 19.2), the latter
atypical with its alternate leaves and trumpet-like flowers).
Lamiaceae grow in savannas, along trails or in dry deciduous forests. Gmelina arborea (‘gambhar’
or ‘kashmari’) and Tectona grandis (teak), native to Burma and Thailand, are intensively cultivated
for their wood. Medicinal herbs (Ocimum gratissimum, AF). Perfumes (oil of Pogostemon cablin,
‘patchouli’, Malaysia).
! In the system of classification of John Hutchinson [2], Verbenaceae and Lamiaceae have been
placed at the top of two divergent lineages, Lignosae for the former and Herbaceae for the latter. More
recent APG classifications have gone in the opposite direction, re-parenting, even merging Verbe-
naceae and Lamiaceae.
! Possible precautions for deterring ants! Hypenia salzmannii (Fig. 19.4) has waxy stems which
may represent an adaptation to deter ants from reaching the flowers to steal nectar and inadvertently
triggering the delicate explosive pollination mechanism on which cross-pollination depends [3].
Fig. 19.3 Clerodendrum speciosissimum, red persistent calyx, fruits green, then turning blue, naturalized in Seram,
Indonesia. Congea tomentosa, Flowers in small heads surrounded by three bracts, Thailand. Congea tomentosa, leaves
and stem pubescent, Thailand
19.1 Lamiaceae: 35-38-39-43-44-45-51-52-56 279
Fig. 19.4 Clerodendrum sp., liana bearing hooks, Madagascar. Gmelina arborea, glands at base and underside of
leaves, their prints visible on the upper side of the leaf, Thailand. Hypenia salzmannii, a herb with fistulose wax-covered
stems, Venezuela
Fig. 19.5 Hyptis sp., inflorescences in heads, Belize. Tectona grandis, large cordate leaves, Bali, Indonesia. Vitex
gaumeri, outer bark and inner bark multilayered, Belize
Fig. 19.6 Vitex quinata, trunk with thin bark and small thorns, Sakaerat RC, Thailand. Vitex sp., stems quadrangular
with spots looking like stipules, Thailand. Vitex gaumeri, leaves compound palmate, Belize
19.2 Lauraceae: 12-20-21-23-24-29-30-39-44
56 genera and 2700 species. Order Laurales. Plants without latex and leaves or bark s.t. aromatic
(Cinnamomum, Laurus). Trees without buttresses. Trunks or branches often lenticellate (Fig. 19.10).
Rhythmic growth and branching demonstrated by leaf clusters, scale-leaves, etc. (e.g. Actinodaphne,
Aniba, Phoebe) are typical features of the Lauraceae. Branches plagiotropic by apposition for some
species (Fig. 19.10). Trunks have continuous branching in Cryptocarya (Fig. 19.9). Periderm of stems
remaining greenish for several consecutive GUs. Stems somewhat angular at their extremities
(Fig. 19.7). Leaf arrangement most often spiral, rarely opposite (Fig. 19.8) (Beilschmiedia, Cin-
namomum, Nectandra oppositifolia), or s.t. two-ranked (Fig. 19.9) (Cryptocarya, Eusideroxylon).
Leaves grouped distally on GUs: Actinodaphne (AS), Aniba spp. (AM). Stipules absent. Leaves simple,
entire, their underside often blue-green (Fig. 19.9). Young leaf-folding more or less conduplicate
(Fig. 19.12), not involute. Glands absent, s.t. translucent dots (Beilschmiedia spp., AF). Venation
densely reticulate (Fig. 19.7), camptodromous (Fig. 19.9), with two strong basal lateral veins in Cin-
namomum (Fig. 19.8). Inflorescences cymose (Fig. 19.12), expanded (e.g. Nectandra) or contracted
(Litsea). Flowers regular, small with 3–6–9 tepals (Fig. 19.12). Ovary superior (inferior in Hypo-
daphnis). Fruits 1-seeded-berries, often partially enclosed in a cupula (Fig. 19.11), important for birds.
Fig. 19.7 Typical characters: a. internodes angular. b. rhythmic branching. c. venation densely reticulate
19.2 Lauraceae: 12-20-21-23-24-29-30-39-44 281
Trees (Actinodaphne, Aniba, Cinnamomum, Dehaasia, Litsea, Nectandra, Ocotea, Persea), some
shrubs, and an atypical genus of small twining and hemiparasite lianas (Fig. 19.12 Cassytha). Lau-
raceae are mainly forest trees. Before migration to the boreotropics, this family would have originated
in Gondwana [4]. This hypothesis is supported by the bulk of Ocotea species in South America. Uses
numerous. Timber: wood of Chlorocardium rodiei is heavy and very resistant. Fruit trees: avocado
tree (Persea americana); spices: Cinnamon tree (Cinnamomum verum), mountain pepper or ‘kilumu’
(Litsea cubeba); essential oils: camphor tree Cinnamomum camphora).
! Lauraceae are especially difficult to identify in the absence of flowers or fruits. However, in the
course of a short study in the Sierra de Lema (Venezuela), inner bark and rhytidome characteristics
show clear differences between the many species encountered there. “Bark World” is not familiar to
academic and “herbarian” botanists, a lot of field work remains to be done by the generation of
dendrologists to come.
Fig. 19.8 Cinnamomum bejolghota, leaves with two strong lateral veins, Thailand
Fig. 19.9 Cinnamomum camphora, leaf clusters, leaves supratripliveined, BG LC, France. Cryptocarya sp., ROUX’s
model, Belize. Cryptocarya sp., venation camptodromous and underside of leaves blue-green, Moluccas
Fig. 19.10 An unidentified species exhibiting plagiotropy by apposition, Thailand. Aniba bracteata, rhythmic
growth demonstrated by scale-leaves (c), Guadeloupe. Litsea cubeba, lenticellate periderm remaining green for a
long period, Thailand
Fig. 19.11 Litsea sp., inflorescences glomerulate, Thailand. Litsea umbellata, infructescences of grouped berries,
Kebun Raya, Java. Nectandra membranacea, berries inserted in a cupula, Guadeloupe
Fig. 19.12 Ocotea lanceolata, inflorescence cymose, flowers with 6 tepals, Southern Brazil. Conduplicate leaf-folding
of an unidentified Lauraceae, Khao Chong, Thailand. Cassytha cf. filiformis, twining hemiparasite, Seram
19.3 Lecythidaceae: 9-10-13-15-18-25-29
24 genera and 360 species (including Scytopetalaceae). Order Ericales. S.t. trees with impressive
buttresses (Couratari). Leaves and bark not aromatic. Bark fibrous, difficult to tear and detaching in
strips (Fig. 19.14). Branches of Barringtonia exhibit alternate series of long and short internodes,
with leaves somewhat clustered (Fig. 19.15). Leaves spiral (Barringtonia, Chydenanthus, Fig. 19.15
Careya, Gustavia) or two-ranked (Fig. 19.15 Bertholletia, Eschweilera, Lecythis). Stipules missing
19.3 Lecythidaceae: 9-10-13-15-18-25-29 283
or minute (Barringtonia). Leaves simple, entire or toothed (Cariniana, teeth s.t. vestigial (Fig. 19.16)
in Eschweilera-Leythis), lamina large (Grias, Gustavia), shortly petiolate or cuneate at its base
(Careya, Gustavia). Foliar glands in Napoleonaea (Fig. 19.18). Indument always absent? Young
Fig. 19.13 Couroupita guianensis, flowering shoots Fig. 19.14 Eschweilera subglandulosa, fibrous bark,
inserted on trunk, cult Venezuela
Fig. 19.15 Barringtonia sp., rhythmic growth, short (EN c) and long internodes (EN l), Indonesia. Bertholletia
excelsa, two-ranked leaf arrangement, cult., Malaysia. Careya sphaerica, scaly bark, leaves spiral, Thailand
leaf-folding involute (Fig. 19.16). Cauliflory (Grias) or flowering shoots inserted basally on trunk
(Fig. 19.13 Couroupita). Flowers large and showy with 4–6-8 petals and numerous stamens
(Fig. 19.14 Couroupita guianensis, Gustavia). Ovary superior. Fruits woody operculate capsules
(Fig. 19.17), (Bertholletia, Cariniana, Eschweilera, Lecythis), indehiscent capsules (Fig. 19.18
Gustavia), Barringtonia, huge coriaceous berries (Fig. 19.16 Couroupita guianensis, Gustavia) or
drupes (Napoleonaea, Scytopetalum). Winged seeds in Couratari.
Numerous American trees, all evergreen (Bertholletia, Couratari, Eschweilera), small trees in
Asia (Barringtonia, Careya) or in Africa (Napoleonaea, Brazzeia, Scytopetalum), several shrubs
(Gustavia, Napoleonaea), no liana and no herbs.
Fig. 19.16 Couratari guianensis, buttressed tree, Venezuela. Couroupita guianensis, fruiting twigs basally inserted on
trunk, cult., FRIM, Malaysia. Eschweilera sp., involute folding, vestigial teeth
Fig. 19.17 Eschweilera cf. tenuifolia, operculate capsules, Venezuela. Eschweilera sp., two-ranked leaf arrangement,
French Guiana. Gustavia poeppigiana, leaves clustered, terminal bud with scale leaves, Venezuela
Fig. 19.18 Gustavia cf. augusta, Venezuela. Flower with numerous stamens inserted on a disk and coriaceous
indehiscent capsules. Napoleonaea sp., leaf base glandular, Cameroon
19.4 Leguminosae—Introduction 285
Although modest in species number, Lecythidaceae are an important element in Guianas and Ama-
zonia, but they grow also in paleotropical-Pacific mangroves (Barringtonia, Foetidia). Timber: Peter-
sianthus, Lecythis, Eschweilera; fruit trees: Bertholletia excelsa (Brazil nut, ‘Castanheira’), Lecythis
zabucajo (‘coco de mono’). Bark also used to make ties, or even hammock for an overnight camp! Fish
poisons (Chydenanthus, AS). Ornamental shrubs: Gustavia augusta has large showy flowers.
! Eschweilera, the genus with the most species in Lecythidaceae, is represented by more trees than any
other genus of plants in Amazonia. African genera (Brazzeia, Scytopetalum) are basal in the family [5].
19.4 Leguminosae—Introduction
780 genera, 19 500 species. Here divided into three groups, recent studies [6] six subfamilies
(Caesalpinioideae, Cercidoideae, Detarioideae, Dialioideae, MIM, PAP). Leaf types below
(Fig. 19.19), CSL = Caesalpinioid paraclade, MIM = Mimosoid clade, PAP = Papilionoideae.
Fig. 19.19 Leaf types

a. paripinnate (CSL-Detarieae, e.g. Gilbertiodendron, Macrolobium, etc.).

b. bifoliolate (CSL-Cercideae: Bauhinia, CSL-Detarieae: Cynometra, Gilbertiodendron, Peltogyne).
c. leaflets alternate, one leaflet terminal (Detarieae, e.g. Crudia, Kingiodendron).
d. pulvinate base of pinnae, A. cupuliformous glands (MIM: Inga, Pithecellobium), B-C, convex
glands (CSL: Chamaecrista, Senna).
e. paripinnate (CSL-Cassieae: Cassia, CSL-Detarieae: Leonardoxa, Macrolobium).
f. bipinnate, pinnae opposite, leaflets opposite and asymmetrical (MIM-Parkieae: Pentaclethra,
MIM-Ingeae: Pithecellobium).
g. bipinnate, pinnae alternate, leaflets opposite (Dimorphandreae: Burkea).
h. imparipinnate leaflets opposite and (st) stipellate (various PAP).
i. imparipinnate, rachis winged (PAP: Swartzia).
j. trifoliolate, leaflets stipellate (PAP: Erythrina).
k. pinnate, leaflets alternate, one leaflet terminal (PAP-Dalbergieae; CSL-Dialioideae [6]: Dialium).
l. signature of the LEGUMINOSAE: stipule, pulvinus and abscission zone.
19.5 Leguminosae–Cercideae–Cassieae–Detarieae–Caesalpinieae:
47-48-55-D
A group encompassing around 170 genera and 2250 species. Caesalpinioideae do not form a natural
group (but a paraclade). Plants without latex. Leaves and bark not aromatic. Some large buttressed
trees (Fig. 19.29 Mora spp.), s.t. trees with fluted trunk (Fig. 19.30 Vouacapoua americana). Branches
s.t. spiny (Caesalpinia, Mezoneuron). Serial buds in rows in Caesalpinieae (Fig. 19.31). Arrangement
of leaves spiral (Cassia, Delonix, Gleditsia, Haematoxylum) or two-ranked (Cassieae: Cassia, Senna;
all the Detarieae: Brownea, Crudia, Detarium, Gilbertiodendron, Hymenaea, Macrolobium, etc.).
Stipules present. Leaves usually simply compound and paripinnate or bifoliolate (Cynometra),
imparipinnate in Dicorynia. Leaves twice compound in many Caesalpinieae (Caesalpinia, Delonix,
Fig. 19.20 Street trees conforming to CHAMPAGNAT’s model, Delonix regia, Delta Amacuro, Venezuela
19.5 Leguminosae–Cercideae–Cassieae–Detarieae … 287
Fig. 19.21 The emblematic ‘Saman’ of Northern South America, TROLL’s model, Samanea saman, Upata,
Venezuela
Gymnocladus, Haematoxylon). Leaves can also be bipinnate (Dimorphandra, Pentaclethra), these s.t.
with a terminal pinnae (Fig. 19.25 Caesalpinia spp.). Leaves rarely simple and bilobate (Bauhinia
spp.). Leaflets usually opposite (Fig. 19.27), rarely alternate (Crudia spp., Gymnocladus). Petiole
enlarged at its base into a pulvinus. Abscission joint between leaflet (or pinnae) and rachis (see
introduction to the family). Leonardoxa africana has hollow foliar rachis, inhabited by ants
(Fig. 19.28). Inflorescences in racemes or spikes of numerous small flowers, terminal (Caesalpinia,
Fig. 19.27 Dimorphandra, Senna) or lateral (Fig. 19.26 Cassia, Senna), s.t. inserted on thick branches
or even on trunks (Fig. 19.25 Brownea, Saraca, Tamarindus). Flowers or inflorescences usually
showy (Fig. 19.25) with a more or less bilateral symmetry (Fig. 19.24) and 5–10 stamens (Fig. 19.26).
Stamens free. Ovary superior. Fruits dehiscent pods or ‘legumes’ (Fig. 19.29), indehiscent pods:
Fig. 19.30 Tamarindus. Pods usually multi-seeded, 1-seeded in Macrolobium spp. (Fig. 19.28).
Trees (Afzelia, Bauhinia, Brachystegia, Copaifera, Detarium, Daniellia, Delonix, Guibourtia,
Intsia, Peltogyne, Tachigali, Tamarindus and many other genera). Trees are irregularly deciduous, i.e.
they frequently keep their leaves at the end of the dry season. Shrubs (Cassia, Caesalpinia), lianas
(Fig. 19.24 Bauhinia, Mezoneuron), very rarely subshrubs or herbs (Fig. 19.26 Chamaecrista
fasciculata).
‘Caesalpinioideae’ are abundant in Africa and America, with many beautiful trees, often exploited
for cabinetwork, e.g. Peltogyne spp. (‘Bois violet’, ‘Zapatero’). Asia has few big trees (Koompassia
excelsa in Sarawak). Several species are gregarious, Brachystegia spp. (South African Miombo
woodlands), Gilbertiodendron spp. (Equatorial Africa), Colophospermum mopane (Fig. 19.22) in
Southern Africa, Copaifera, Mora spp.in the Guianas. Some “precious woods” such Peltogyne grow
very slowly, but Schizolobium parahyba (Fig. 19.23 Amazonian ‘guapuruvú’) one of the fastest
growing trees in the world. In Guianas, Mora spp. (Fig. 19.29) have the biggest seeds among
‘Dicotyledons’. Fruit trees: Tamarindus indica (Fig. 19.30). Lacquers, copals (oleoresins): AM:
Copaifera (Fig. 19.27), AF: Daniellia and Gossweilerodendron (‘agba’), AS: Kingiodendron (AS).
Dyes: ‘bois de Campeche’ or ‘pau Brazil’: Haematoxylum campechiana (Fig. 19.27), colouring agent
for histology, Biancaea (syn. Caesalpinia) sappan (‘sappannam’ of India) provides a red dye for wool
and cotton. Numerous ornamental trees, some of the best known are ‘flamboyants’ because of their
orange or red flowers: Amhertsia nobilis (Fig. 19.24 Burma), Delonix regia (Fig. 19.20) threatened in
its native environment (MA), is grown everywhere as a street tree; Peltophorum pterocarpum (yellow
flamboyant of Indomalesia). Ornamental shrubs and lianas Caesalpinia spp., Bauhinia spp.
! Monodominance in tropical forests is observed for several Caesalpinioid species, in Africa,
Gilbertiodendron dewevrei and in the Guianas, Mora excelsa [7].
! If one were to attribute a personality to this group, one might say that the Caesalps are big,
beautiful and somehow joyful!
Fig. 19.22 Dry forest with Colophospermum mopane, Fig. 19.23 A fast growing post-pioneer tree: Schizolo-
Chobe, Botswana bium parahyba, Cockscomb, Belize
19.5 Leguminosae–Cercideae–Cassieae–Detarieae … 289
Fig. 19.24 Amherstia nobilis, drooping inflorescence, Kebun Raya, Java. Bauhinia sp., liana ‘Echelle tortue’, French
Guiana. Bauhinia variegata, Chiang Mai, Thailand
Fig. 19.25 Brownea sp., inflorescence in a dense head, Venezuela. Cenostigma gaumeri, leaf bipinnate with terminal
pinnae, Shipstern, Belize. Erythrostemon (syn. Caesalpinia) yucatanensis, flowers with bilateral symmetry, Belize
Fig. 19.26 Campsiandra sp., Orinoco embankments, Venezuela. Cassia moschata, lateral flowering, Upata,
Venezuela. Chamaecrista sp., subshrub, Venezuela
Fig. 19.27 Copaifera sp., leaves paripinnate, Venezuela. Dimorphandra macrostachya, inflorescence terminal,
Aponwao, Venezuela. Haematoxylum campechianum, leaves paripinnate, Guadeloupe
Fig. 19.28 Leonardoxa africana, sliced internode shows hollow stems inhabited by ants, Campo forest, Cameroon.
Macrolobium acaciifolium, flat pod enclosing a single seed, Caura, Venezuela
Fig. 19.29 Mora gongrijpii, Venezuelan Guayana. Buttressed tree and tree reiterating after a stress (many dead
branches). Saraca thaipingensis, terminal inflorescences and fruits, BG Singapore
19.6 Leguminosae–Mimoseae–Acacieae … 291
Fig. 19.30 Senna surattensis, Chiang Mai, Thailand. Tamarindus indica, indehiscent fruits, grown in Belmopan,
Belize. Vouacapoua americana, fluted trunk, French Guiana
Fig. 19.31 Serial buds in Caesalpinieae. a: side view (p: petiole). b: ramified upper bud
19.6 Leguminosae–Mimoseae–Acacieae–Ingeae: 20-30-46-57-D
Including here the Pentaclethra group, the “MIMO” group encompasses around 85 genera and
3300 species. Plants without latex, s.t. producing gums (Senegalia, AF). Leaves and bark not
aromatic. Internodes angular with petiolar base decurrent on the stem. Inner bark s.t. with a
network of fibres (Inga). Trunk or branches with lenticels transversely elongate (Fig. 19.44), these
usually disappearing on old trunks (Fig. 19.45). Branches s.t. spiny, the spines of stipular origin
(Acacia s.l. = Acacia + Senegalia + Vachellia, Mimosa, Piptadenia spp.). Spines in Vachellia
cornigera are hollow and inhabited by ants Pseudomyrmex ferruginea. Spines of Dichrostachys are
modified short shoots, Colateral buds (i.e. two or three buds are grouped in leaf axils Fig. 19.39).
Leaves spiral (Anadenanthera, Enterolobium, Mimosa, Piptadenia, Pentaclethra, Pithecellobium,
all AM) or two-ranked: Acacia s.l. Inga (AM), Dichrostachys (AF-MA). Stipules present. ‘Simple
leaves’ of Australian Acacia are in fact phyllodes (i.e. petioles modified into a lamina Fig. 19.35).
Leaves bipinnate (Fig. 19.42), without a terminal pinnae, or leaves simply pinnate (Fig. 19.39
Inga). Leaflets asymmetric (Fig. 19.42), s.t. very numerous (Parkia) or few (Fig. 19.38
Calpocalyx, Zygia), sensitive to touch in most, but not all, Mimosa species. Abscission joint
between leaflet and rachis (see introduction). Rachis or base of petiole glandular (Cedrelinga, Inga,
Fig. 19.32 Calliandra calothyrsus, SCARRONE’s Fig. 19.33 Falcataria falcata, TROLL, Flores,
model, invasive, Flores, Indonesia
Fig. 19.34 Enterolobium cyclocarpum, TROLL’s model, Belize
Figs. 19.33 and 19.34
Fig. 19.33
Fig. 19.41 Leucaena, Pseudosamanea (AM), Xylia, AS, Acacia s.l.) or without glands (Adenan-
thera (AS-OC), Calliandra, Mimosa, AM). Flowering usually lateral (Inga, Mimosa, Piptadenia,
Stryphnodendron). Inflorescences in spikes (Fig. 19.38 Dichrostachys, Entada, Fig. 19.42 Mimosa,
Fig. 19.43 Piptadenia) or glomerulate (Acacia, Fig. 19.41 Mimosa). Flowers small, with a radial
symmetry, stamens free, not numerous (5–10) or numerous (Fig. 19.40 Inga), s.t. showy, red-
coloured. Ovary superior. Fruits pods, dehiscent or not dehiscent, s.t. flat, coiled (Fig. 19.43) or
splitting in 1-seeded parts (Entada Fig. 19.39). Seeds brown or black, s.t. red (Adenanthera
Fig. 19.36), covered by a mucilaginous sarcotesta in Inga spp.
Trees (AS: Acacia s.l., Albizia, Archidendron, Xylia; AM: Dinizia excelsa, Enterolobium, Inga,
Leucaena, Piptadenia), shrubs (Fig. 19.32 Calliandra), lianas (Fig. 19.36 Acacia, Entada) or herbs
(Fig. 19.41 Mimosa), Neptunia being the single aquatic genus.
Fig. 19.35 Vachellia (syn. Acacia) cornigera, hollow spines inhabited by ants, Belize. Vachellia (syn. Acacia)
hebeclada, deeply fissured bark, Botswana. Acacia podalyriifolia, phyllodized petioles and flat pod, BG LC, France
Fig. 19.36 Acacia sp., liana, Madagascar. Acacia sp., cross-section of a liana showing reddish phloem poles, Campo
forest, Cameroon. Adenanthera pavoniana, dehiscent coiled pods with red seeds, INRA, Guadeloupe
Fig. 19.37 Albizia lebbeck, indehiscent pods (orig. South East Asia), Yucatan. Calliandra calothyrsus, showy stamens
with long pink filaments, (orig. AM), Flores, Indonesia. Calliandra sp., showy stamens with red filaments, Southern Brazil
Leguminosae-’Mimosoids’ are tropical and subtropical in their distribution, many species can
support aridity and very few of them grow at high elevations. The presence of a fissured bark
(Fig. 19.35) or a chlorophyllous layer (Fig. 19.44) is fequent in savanna trees. Calliandra and Acacia
are typical of disturbed vegetation in dry tropics (e.g. the Calliandra-savannas in tropical America).
Leucaena leucocephala (Fig. 19.40), native to AM-C?, became invasive in all tropics. Acacia
auriculiformis trees are grown for restoration of secondary forests, their shade enhance native
seedlings and saplings. Samanea saman (‘saman’) is a shade tree grown in all townships of South
America. Street trees: Albizia lebbeck (AS).
Uses: reverse phenology of Faidherbia albida makes it the most compatible tree in African
agroforestry; firewood (mainly in AF); arabic gum (Senegalia senegal); dye extracted from heart-
wood (Senegalia catechu); fruit trees with sweet sarcotesta covering the seeds (e.g. Inga edulis, ice
cream bean, in Latin America: ‘guamo’, ‘ingá’, ‘shimbillo’). Several Senegalia species produce toxic
amine-derivates. Hallucinogenic: seeds of Anadenanthera peregrina (AM-S) for preparing ‘niopo’.
Acacias are emblematic trees of African savannas, a biome which has greatly extended in the last
centuries and reaches the 10th parallel or even the equator in Tanzania. Large scale planting of acacia
trees in sub-Saharan Africa is an attempt to reverse this process [8] (Fig. 19.37).
Fig. 19.38 Calpocalyx sp., leaves bipinnate each pinnae with 4 pairs of leaflets, Campo forest, Cameroon.
Dichrostachys cinerea, inflorescence in spikes with anthesis acropetal (orig. MA), BG LC, France. Entada polystachya,
spikes of flowers in large compound inflorescences, French Guiana
Fig. 19.39 Entada polystachya, pods breaking in indehiscent segments, Venezuela. Inga sapindoides, winged and
glandular rachis, Belize. Collateral buds, small bud (b) near to a large bud (B); a Inga sp., b Samanea saman
Fig. 19.40 Inga sp., flower with numerous stamens, Southern Brazil. 20. Inga cf. vera., striate pod, Cockscomb,
Belize. Leucaena leucocephala, fascicles of pods, Flores, Indonesia
Fig. 19.41 Leucaena leucocephala, glands disposed at rachis nodes. Lysiloma latisiliquum, flat pods, Belize. Mimosa
pudica, ‘sensitive plant’, Belize
Fig. 19.42 Mimosa myriadenia, inflorescences are racemes of spikes, leaves not sensitive to touch! Bochinche,
Venezuela. Parkia sp., leaf bipinnate, rachis glandular and very numerous leaflets, French Guiana. Zygia cf. collina,
leaf bipinnate with asymmetric leaflets, Venezuela
Fig. 19.43 Piptadenia leucoxylon, inflorescence in spikes, Guri, Venezuela. Pithecellobium s.l., bipinnate leaves,
reddish when young, Venezuela. Pithecellobium unguis-catis, coiled pods, Mexico
Fig. 19.44 Stryphnodendron guianense, inflorescences in lateral spikes, Bochinche, Venezuela. 32–33. Vachellia
(syn. Acacia) tortilis. Lenticels, chlorophyllous layer. Coiled pods, Botswana
Fig. 19.45 Zygia stevensonii, leaves bipinnate, each pinnae bearing, 3 (not 4!) leaflets (fm: missing leaflet), Belize.
Xylia xylocarpa, curious wrinkled rhytidome in an adult tree (lenticels are no more visible), Kerala, India
19.7 Leguminosae–Papilionoideae: 2-6-8-9-46-47-50-53-54-57-D 297
19.7 Leguminosae–Papilionoideae: 2-6-8-9-46-47-50-53-54-57-D
Around 500 genera and 12,000 species. Plants without latex, s.t. their bark producing a red exudate
(Inocarpus, Macropsychanthus, Platysepalum, Fig. 19.57 Pterocarpus). Leaves and bark not aro-
matic. Trunk or branches with lenticels transversely elongate. Lianas are twining (Derris, Phaseolus)
or bear prehensile twigs (Machaerium). Branches or trunks s.t. spiny (Fig. 19.54 Erythrina,
Machaerium lianas). Axillary buds usually solitary, rarely collateral (Sesbania spp.) or in vertical row
(Fig. 19.58 Swartzia). Leaf arrangement usually spiral (Erythrina, Mucuna, Sophora, etc.), s.t. two-
ranked (Dalbergia, Machaerium), rarely opposite (Platymiscium, Fig. 19.59 Tipuana). Stipules
present, rarely absent (Fig. 19.48 Alexa). Leaves compound 1 pinnate, usually imparipinnate
(Fig. 19.58), less frequently paripinnate (Fig. 19.57), rarely ‘unifoliolate’ (petiole with abscission
joint and distal pulvinus), (Baphia nitida, Fig. 19.51 Dalbergia spp., Uraria spp., Flemingia stro-
bilifera). Leaves very rarely truly simple (Fig. 19.59 Zollernia, AM). Leaflets usually opposite
(s.t. more or less alternate, Dalbergia (Fig. 19.51), Diplotropis spp., Machaerium, AM). Leaflets
s.t. stipellate (Erythrina (Fig. 19.19k), Platysepalum, Swartzia). Abscission joint between leaflet and
rachis (see introduction). Leaves or leaflets entire, very rarely toothed (Zollernia). Venation pinnate,
rarely tripliveined (Flemingia), usually camptodromous, rarely with a fimbrial vein (Dussia,
Fig. 19.54 Machaerium, Platysepalum). Foliar glands almost always missing, e.g. Erythryna spp.),
s.t. translucent dots (Myrocarpus, Myrospermum, Myroxylon, AM). Flowering terminal (Machaer-
ium, Ormosia, Sophora) or lateral (Clitoria, Dalbergia). Inflorescences very variable: panicles,
racemes, spikes or 1-flowered. Flowers with a bilateral symmetry (Figs. 19.49 and 19.50) and 5–10
stamens. Flowers resupinate in many Phaseoleae (e.g. Macroptilum Fig. 19.55, Vigna). Stamens free
or fused in a tube. Fruits usually pods (Fig. 19.53), samaras Centrolobium (Fig. 19.49), Machaerium
(Fig. 19.55) or something in between, rarely drupes (Coumarouna, Inocarpus). Seeds s.t. orange and
black or red and black (Fig. 19.48 Abrus, Sophora spp.).
Trees (Pantrop: Dalbergia, Erythrina, Pterocarpus (Fig. 19.46) Sophora; AM: Alexa, Andira,
Centrolobium, Dipteryx, Myroxylon, Lonchocarpus, Ormosia, Swartzia; AF: Cadia, Millettia,
Pericopsis; AS-AU-OC: Butea, Fordia, Inocarpus, Intsia, Millettia, etc.); evergreen (e.g. Alexa) or
deciduous (e.g. Lonchocarpus Fig. 19.54). In savannas, bark becomes deeply fissured and a
chlorophyllous layer appears between dead bark and inner bark (Fig. 19.56). Shrubs: Cajanus,
Dalbergia, Indigofera, Sesbania, etc. Lianas: Abrus, Dalbergia, Dioclea, Machaerium, Mucuna
(Fig. 19.56), Phaseolus. Woody herbs: Arachis, Canavalia, Crotalaria, Desmodium, Indigofera, etc.
Papilionoideae form the most diversified group of the LEGUMINOSAE, they grow in all kinds of
environments, extending from equatorial forests to tundra, steppes of Central Asia and Himalayan-
Alpine mountain ranges. Uses are very numerous, a multipurpose tree: Inocarpus fagifer (Haitian
chestnut) for roasted seeds, charcoal, antidote against fish venom and evidently symbolic (red exudate
in bark!). Cabinetwood: Andira inermis, Dalbergia spp., Centrolobium paraense (‘pau rainha’,
‘cartán’). Essential oils: Aniba rosodora ‘pau rosa’ (from distilled wood). Oil seeds: Arachis hypogea
(groundnut). Shade trees for cacao- or coffee-agroforestry (Erythrina spp., Sesbania grandiflora).
Numerous beans native to America: Cajanus (Fig. 19.49 ‘pois cajun’, pigeon pea), Lablab purpureus
(lablab bean), Phaseolus (‘frijoles’, ‘feijoas’), Psophocarpus tetragonolobus (Goa bean). Sources of
perfumes: Dipteryx odorata (Fig. 19.53 ‘Sarrapia’). Butea monosperma, the sacred red flowering tree
of brahmans provides an astringent gum (Bengal gum). Dyes (indigo, Indigofera tinctoria), gums
(Pterocarpus marsupium). Caution! Seeds of Abrus precatorius (Fig. 19.48), when swallowed and
the outer coating is damaged, are very toxic (toxalbumin).
Fig. 19.46 Buttressed trees (Pterocarpus officinale) in Fig. 19.47 A herb conforming to ACOSTA’s model,
the sanctuary of Crique Toussaint, French Guiana Indigofera sp., Flores, Indonesia
Fig. 19.47
Fig. 19.48 Abrus precatorius, seeds red and black, Venezuela. Alexa imperatricis, no visible stipules! Imataca,
Venezuela. Bowdichia virgilioides, inflorescence paniculate, terminal, Venezuela
! In the lowland tropics of Africa and America it is almost impossible that a hundred-meter square
plot of vegetation does not contain a PAP shrub or tree.
! One may wonder why light tolerant trees valuable for cabinetwork, like Centrolobium paraense
(orange-coloured wood!), and many other PAP, do not benefit from silvicultural programmes taking
advantage of savannized regions or degraded forests instead of logging old-growth forests [9]
(Fig. 19.52).
Fig. 19.49 Cajanus cajan, cultivated Pigeon pea, Upata, Venezuela. Canavalia sp., prostrate herb on a shore, French
Guiana. Centrolobium paraense, samara with prickly basal part, Venezuela
Fig. 19.50 Chadsia sp., flower with bilateral symmetry, Madacascar. Clitoria ternatea, shrub, flowers axillary and
solitary, Bali, Indonesia. Crotalaria cf. retusa, spike of flowers, Guadeloupe
Fig. 19.51 Dalbergia ecastaphyllum, inflorescences lateral, Southern Brazil. Dalbergia latifolia, indehiscent pods,
Bali, Indonesia. Dalbergia latifolia, leaves with alternate leaflets, Bali, Indonesia
Fig. 19.52 Dendrolobium cf. umbellatum, fruits lomentaceous, Indonesia. Uraria oblonga, subshrub, simple leaves,
raceme of flowers, Thailand. Macropsychanthus (syn. Dioclea) macrocarpa, liana producing a red exudate, Fr. Guiana
Fig. 19.53 Dioclea macrocarpa, pods and knife ‘Opinel’ size-12, French Guiana. Dipteryx odorata, BG Ciudad
Bolivar, Venezuela. Dussia sp., leaflets with fimbrial veins, Guadeloupe. Erythrina folkersii, floral buds, Belize
Fig. 19.54 Erythrina subumbrans, spiny trunk, Thailand. Lonchocarpus cf. tubicalyx, deciduous flowering tree, BG
Ciudad Bolivar, Venezuela. Machaerium guaremalense, leaflets alternate, Venezuela. Machaerium sp., fimbrial veins
(fv), Belize
Fig. 19.55 Machaerium sp., terminal panicle of samaras, Venezuela. Machaerium sp., very rhythmic growth
demonstrated by scale-leaves, Venezuela. Macroptilium atropurpureum, herb with resupinate flowers, Flores,
Indonesia. Millettia pinnata, small tree of back-shores, bearing pods, Flores, Indonesia
Fig. 19.56 Mucuna macrocarpa, trunciflorous liana, flowers with greenish standard and purple wings, Thailand.
Mucuna urens, liana with drooping inflorescences, Venezuela. Philenoptera nelsii, chlorophyllous layer, Botswana
Fig. 19.57 Platymiscium trinitatis, yellow flowers, Venezuela. Pterocarpus officinale, red and astringent exudate in
bark, French Guiana. Sesbania punicea, leaves paripinnate, Venezuela
Fig. 19.58 Strongylodon macrobotrys, turquoise blue is an extremely rare colour in angiosperm flowers, (liana native
to Philippines) BG LC, France. Swartzia sp., serial buds, Venezuela. Swartzia sp., leaves imparipinnate, S-Brazil
Fig. 19.59 Tipuana tipu, stem with leaves opposite pinnate mimicking a bipinnate leaf (b: axillary bud), BG Villa
Thuret, Antibes. Zollernia paraensis, leaves simple with spiny teeth, Caura Basin, Venezuela. Camptodromous
venation of an unidentified Papilionoideae, Sumatra
19.8 Linaceae–Hugonioideae (H) + Ixonanthaceae: 13-18-29-30-54 (I)
Two taxa encompassing 12 genera and 50 species (more 230 species in extratropical Linum). Order
Malpighiales. Trees or shrubs (I: Ixonanthes, AS; H: Hebepetalum, Ochthocosmus, Roucheria,
AM), lianas bearing opposite hooks (H: Fig. 19.60 Hugonia, Paleo) or alternate hooks (H: Indor-
ouchera, Andaman isles, AS-SE). Rhythmic growth and branching. Leaves usually spiral (H:
Hugonia, Paleo; I: Ochthocosmus, AM) or two-ranked (I: Phyllocosmus, AF). Stipules present or
missing. Leaves shortly petiolate or subsessile, entire or crenulate. Young leaf-folding involute
(Fig. 19.60) (b). Venation camptodromous (Fig. 19.60) (c) or brochidodromous (d), main vein raised
at upper side of the lamina. Indument none. Inflorescences terminal, cymose (Fig. 19.61a) or lateral,
racemose (Fig. 19.61b, c). Flowers usually regular and 5-merous. Fruits capsules (Ixonanthaceae) or
drupes (Hugonioideae).
Adding to this group Erythroxylaceae, Irvingiaceae and Humiriaceae forms a set equivalent to the
Cronquist’s Linales [10], in which the members shares at least three traits in common: strong
rhythmic growth, leaves glabrous and, at the beginning of leaf development, leaves involute in bud.
Consequently, confusion among these five taxa is to be expected. In addition, in America, possible
confusion of Roucheria with other ‘rhythmic taxa’ such as Ouratea (Ochnaceae), which also have
numerous parallel secondary veins, but the latter bear intrapetiolar stipules (Fig. 19.62).
Fig. 19.60 Hugonia, a: hooks axillate by opposite prophylls, b: involute young leaves, c: venation camptodromous
and scalariform. d. Roucheria humiriifolia, leaf entire, venation brochidodromous with numerous secondary veins,
detail: canaliculate petiole, French Guiana
19.9 Loganiaceae + Gelsemiaceae: 32-33-35-38-56 303
Fig. 19.61 a Ixonanthes icosandra, cymose infructescence. b Ochthocosmus roraimae, crenulate leaves, lateral
racemes of tiny flowers, Venezuela. c Phyllocosmus africanus, entire leaf, fascicle of spikes of flowers, West Africa. All
drawings according to herbarium specimens, CBG H
Ixonanthes icosandra were used for tanning fishing nets in Malaysia. Hugonioideae might be
considered at the family rank (Hugoniaceae). Extratropical Linum, the only genus in Linoideae being
very exotic in this group.
19.9 Loganiaceae + Gelsemiaceae: 32-33-35-38-56
16 genera and 400 species. Order Gentianales. Plants without latex, not aromatic. The majority of the
species are Strychnos, and most of these are lianas climbing by means of short shoots modified into
hooks (Fig. 19.65). Peculiar scattered disposition of phloem (or other tissue) in xylem (Fig. 19.66),
s.t. sclerenchymatous cells coming along with phloem (Fig. 19.64 Strychnos spp.). Opposite leaf
arrangement, s.t nodes enlarged (Fig. 19.67 Geniostoma). Stipules s.t. present (Labordia, Mostuea,
Neuburgia), interpetiolar ridge (Fig. 19.67 Geniostoma, Strychnos). Leaves simple, entire. Glands
missing. Leaf venation of Strychnos is typical: two strong lateral veins run along the lamina
(Fig. 19.65). Plant glabrous except Antonia ovata. Flowers with 4–5 free or fused petals (Fig. 19.63
Similar families:
GENTIANACEAE and RUBIACEAE
are two taxonomically very close
families. Helicoidal anisoclady (Fig.
19.62) exists also in RUBIACEAE.
Logania imbricata (NC), Spigelia
anthelmia (AM, nat. Paleotropics).
MASSART: Strychnos axillaris (AS).
MANGENOT: Strychnos spp.
SCARRONE (Fig. 19.66):
Neuburgia moluccana (AS).
STONE (Fig. 19.62): Antonia ovata
(Guianas), Bonyunia minor (N AM-S).
CHAMPAGNAT: Strychnos
Fig. 19.62 Antonia ovata, helicoidal madagascariensis, Gelsemium
anisoclady, each lateral branch are sempervirens (AM-N).
inserted in a different quadrant, Fr. TROLL: probably absent.
Guiana
Mostuea), corolla contortate (Fig. 19.67). Ovary superior. Fruits capsules (Antonia, Fig. 19.67
Geniostoma), berries (Fig. 19.64 Strychnos) or drupes (Neuburgia).
Small trees (Neuburgia), shrubs (Antonia, Geniostoma, Labordia, Logania, Strychnos), woody
lianas (Strychnos) and herbs (Mitracasme, Mitreola, Schizacme, Fig. 19.63 Spigelia).
Fig. 19.63 Logania imbricata, LEEUWENBERG’s model. Mostuea sp.(Gelsemiaceae), corolla of fused petals,
stipular ruff, Madagascar. Spigelia anthelmia, inflorescence in spikes, naturalized, Bali
Fig. 19.64 Strychnos congolana, young plant, two cotyledons, first axis becoming plagiotropic then erect trunk with
rhythmic growth and branching. Strychnos fendleri, salmon pink inner bark and sclerenchymatous inclusions of phloem
in wood, Venezuela. Strychnos madagascariensis, shrub bearing coriaceous berries, Madagascar
Fig. 19.65 Strychnos nux-blanda, coriaceous berry and seeds, Thailand. Strychnos peckii, hook inserted in the axil of a
fallen scale-leaf (c), Belize. Strychnos sp., typical venation, Thailand
19.10 Lythraceae: 33-37-38-41 305
Fig. 19.66 Strychnos peckii, Belize. Liana cross-section showing phloem poles arranged in a ring inside wood.
Strychnos sp., phloem scattered in wood (other disposition). Neuburgia moluccana, Indonesia, leaves clustered, stipules
adnate to the petiole, SCARRONE’s model
Fig. 19.67 Geniostoma rupestre, nodes enlarged, arillate seed (a.s.), Philippines. Labordia waiolani, contortate
corollas, Hawaii
Essentially tropical. Strychnos extends from AF to AU and Oceania. Strychnine, a poison used in
the past to kill rats, is extracted from seeds of Strychnos nux-vomica. This alkaloid was used in the
preparation of dart poisons (curare), named ‘ourari’ in Guyana (Strychnos toxifera), ‘belai hitam’ in
Malaya (S. scotechinii), [11]. Another toxic plant: Spigelia anthelmia (‘la Brinvilliers’), in remem-
brance of a criminal case. Ground seeds of Strychnos potatorum are used in India for clarifying
muddy water.
19.10 Lythraceae: 33-37-38-41
28 genera and 500 species. Order Myrtales. Plants without latex, leaves and bark not aromatic. Bark
sloughing off in scales (Fig. 19.68, Fig. 19.72 Lagerstroemia) or fibrous strips. Trunks of several
species bear pointed stumps (which are bases of old branches) (Fig. 19.71). Internodes angular, s.t.
alternatively twisted in horizontal stems (Fig. 19.70 Lagerstroemia, Lafoensia). Opposite leaf
arrangement, the leaves of the same pair s.t. staggered (Fig. 19.69). Stipules s.t. present but minute
(Fig. 19.72 Lagerstroemia), absence of interpetiolar ridge. Leaves simple, entire. Glands missing,
except in Sonneratia (Fig. 19.69) (b). Venation usually campto (Fig. 19.72 Lagerstroemia,
Lafoensia). Plants glabrous or pubescent. Flowers with a radial symmetry, 4–6-8 free petals
(Fig. 19.70) and numerous stamens in Sonneratioideae (Fig. 19.73), but bliabiate-tubular with sta-
mens twice the numbers of petals in Lythroideae (Fig. 19.71 Cuphea). Fruits capsules.
Trees (AS: Duabanga, Lagerstroemia, mangrove trees with pneumatophores (Fig. 19.73): Son-
neratia), shrubs (AM: Lafoensia) and herbs (AM: Cuphea), a few aquatics (Lythrum portula, Rotala
spp., Trapa natans). Trees of this family are typical of tropical dry deciduous forests, mainly in Asia
(Lagerstroemia), less often in America (Lafoensia pacari in Southern Brazil and Paraguay). Timber
(AS: Duabanga, Lagerstroemia); fruit trees (pomegranate: Punica granatum, but extratropical);
source of dye (‘héné’: Lawsonia inermis, Paleo); ornamentals as street trees (Lagerstroemia spp.) or
small herbs (Cuphea spp.). Trapa natans, the European water-chestnut, is cultivated in Vietnam.

COMBRETACEAE and MYRTACEAE, MASSART (Fig. 19.69a): Duabanga grandiflora (AS)
two other families belonging to the order ATTIMS: Sonneratia caseolaris (mangrove, AS-OC).
Myrtales. Combretaceae are usually TROLL: Lagerstroemia macrocarpa (AS).
pubescent and glandular and leaves of
many Myrtaceae have translucent dots.
Fig. 19.68 Lagerstroemia duperreana, patchy sloughing off of the rhytidome, Thailand
Fig. 19.69 Lagerstroemia sp., TROLL’s model, Bali, Indonesia. a: Duabanga grandiflora, MASSART’s model,
branch leaves are larger than trunk leaves; b: Sonneratia sp., glands (gl) distally disposed on the internode;
c: Lagerstroemia sp., subopposite staggered leaf arrangement (F1-F2, F3-F4, F5-F6)
19.10 Lythraceae: 33-37-38-41 307
Fig. 19.70 Duabanga grandiflora. Camptodromous venation, Thailand. Angular-winged internodes, these alternately
twisted, Seram, Indonesia. Coriaceous flower buds, Thailand
Fig. 19.71 Cuphea llavea, tubular calyx and bilabiate corolla, cult. Lagerstroemia sp., thorny trunk, scaly bark,
Thailand. Lagerstroemia duppereana, pubescent stem apex, Thailand
Fig. 19.72 Lagerstroemia sp., bark sloughing off in plates, trunk with pointed stumps, Thailand. Lagerstroemia sp.,
leaves shortly petiolate, vestigial prints of stipules, Bali. Lagerstroemia sp., angular internodes and short petioles,
Thailand
Fig. 19.73 Sonneratia sp., flower with numerous stamens, Flores, Indonesia. Sonneratia sp., flower and its 6 sepals
(petals have been removed), Flores, Indonesia. Sonneratia sp., pneumatophores, mangrove, Bali
! Botanists can be classified into “splitters” and “lumpers”, splitting is nowadays a mainstream
tendency in taxonomy. Lythraceae represents an example of a lumping process in APG classifications
where Lythraceae s.str., Punicaceae, Sonneratiaceae and Trapaceae were united in a single family. As
a consequence, Lythraceae becomes a well-balanced assembly of trees, shrubs and herbs.
References
1. Willis, J.C. 1973. A Dictionary of the Flowering Plants and Ferns, 8th ed., 1246 + 66 pp. Cambridge: Cambridge
University Press.
2. Hutchinson, J. 1973. The Families of Flowering Plants, Arranged According to a New System Based on Their
Probable Phylogeny, 3rd ed., 968 pp. Oxford: Oxford University Press.
3. Harley, R. 1999. Lamiaceae. In Flora of the Venezuelan Guyana, vol. 5., 834 pp. St. Louis (Missouri): ed. J.A.
Steyermark, P.E. Berry, K. Yatskievych, and B.K. Holst.
4. Chanderbali, A.S., H. van der Werff, and S.S. Renner. 2001. Phylogeny and historical biogeography of Lauraceae:
Evidence from the chloroplast and nuclear genomes. Annals of the Missouri Botanical Garden 88: 104–134.
5. Mori, S.A., C.-H. Tsou, C.-C. Wu, B. Cronholm, and A.A. Anderberg. 2007. Evolution of Lecythidaceae with an
emphasis on the circumscription of Neotropical genera: Information from combined NDHF and TRNL-F sequence
data. American Journal of Botany 94: 289–301.
6. Legume Phylogeny Working Group (LPWG). 2017. A new classification of the Leguminosae based on a
taxonomically comprehensive phlyogeny. Taxon 66: 44–77.
7. Torti, S.D., P.D. Coley, and T.A. Kursar. 2001. Causes and consequences of monodominance in tropical lowland
forests. The American Naturalist 157: 141–153.
8. Buchter, J.-E. 2019. Reverdir le Sahara, 200 pp. Lausanne: Editions Favre.
9. Falcão, M.T., Cordovil Benezar, R.M., da Silva, W.R., da Silva, G.P., and Ferreira Barbosa Fernandez, M.A. 2010.
Estudo etnobotânico na comunidade indígena Serra da Moça, Boa Vista - Roraima. VI Seminário Latino-
Americano de Geografia Física, 11. Universidade de Coimbra, May 2010.
10. Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants, 1262 pp. Columbia: Columbia
University Press.
11. Burkill, I.H. 1935. A Dictionary of the Economic Products of the Malay Peninsula, vol. I and II, 2402 pp. Oxford:
University Press.
Magnoliaceae to Myrtaceae
20
20.1 Magnoliaceae: 5-14
2 genera and 221 species (Magnolia, Liriodendron). Order Magnoliales. According to Nooteboom [1] and
Sangtae [2], Magnolia would be better split into more than two genera (Elmerrillia, Magnolia, Michelia,
Talauma, etc.). Plants without latex. Some species aromatic in their bark or leaves (e.g. Elmerrillia ovalis,
Michelia alba, Talauma candollei). Bark smooth or rough, often lenticellate, s.t. with eye-marks (Fig. 20.2).
Inner bark often with a network of fibres (Elmerrillia ovalis, Fig. 20.2 Magnolia spp.). Spiral arrangement
of leaves (Fig. 20.2 Magnolia s.str.) or two-ranked arrangement (Michelia, Elmerrillia). Stipules hood-
like, leaving an annular scar on stems (Fig. 20.2). Leaves simple and entire. Inflorescences 1-flowered,
An unmistakable family!
RAUH: Michelia champaca (AS).
SCARRONE Liriodendron
tulipifera (E AM-N).
FAGERLIND: Magnolia
grandiflora, M. obovata.
ROUX: Elmerrillia papuana (AS).
MANGENOT? Talauma candollei
(syn. Magnolia liliifera), Egg
Magnolia or ‘Bunga telur’, a small
tree of Malaysia, about 3m in
height.
TROLL? : Magnolia cf.
denudata.
Fig. 20.1 Magnolia cf. denudata, erect
trunk with two-ranked arrangement
of twigs, in cultivation
https://doi.org/10.1007/978-3-031-05942-1_20
310 20 Magnoliaceae to Myrtaceae
Fig. 20.2 Magnolia sargentiana, fibrous bark, in cultivation. Magnolia liliflora, stipules hood-like, BG Sirikit,
Thailand. Magnolia x soulangeana, follicular fruit, in cultivation
Fig. 20.3 Magnolia cf. denudata, stamens and carpels spirally arranged, cult., Switzerland. Michelia champaca, trunk
smooth, with eye marks (e-m), Khao Chong, Thailand. Michelia champaca, flowers with tepals spirally arranged, in
cultivation, © Richard Lyons
usually terminal. Flowers with numerous tepals and very numerous spirally arranged stamens (Fig. 20.3).
Fruits syncarps consisting of dehiscent follicles, syncarps become fleshy in various species (Fig. 20.2).
Magnoliaceae are of Laurasian origin and exist in tropical and temperate forests of Asia and
America, but are absent from Africa. Ornamental extratropical trees: Liriodendron tulipifera, Mag-
nolia grandiflora, M. kobus, M. x soulangeana. Distilled flowers of Michelia champaca (an Indian
tree) serve as a source of perfumes.
! Magnolia is currently divided in 15 sections [3] and it would be interesting to see how the six
architectural models observed in the genus correlate with the sections [4].
! TROLL’s models would exhibit a peculiar form in this family with erect trunks and two-ranked
leaf/branch arrangement (Fig. 20.1).
20.2 Malpighiaceae: 4-17-33-40-44-56
74 genera and 1300 species. Order Malpighiales. Plants usually without latex (exudate yellowish
in Spachea spp.). If lianas, then twining. Twigs lenticellate. Leaves and bark not aromatic. Opposite
(Fig. 20.7), rarely spiral arrangement of leaves (Acridocarpus, AF). Stipules often present (Fig. 20.8),
intrapetiolar (Fig. 20.6d Byrsonima) or small and inserted on stem or on petiole (Fig. 20.7 Hiraea spp.), s.
t. absent or vestigial. Leaves simple and entire (exceptionally toothed). Glands on the underside of
20.2 Malpighiaceae: 4-17-33-40-44-56 311
lamina, petiole or lamina margins (Fig. 20.6).

Venation brochidodromous (Fig. 20.7). Indument
of simple, appressed hairs (Fig. 20.6e). Inflores-
cences terminal (Byrsonyma, Spachea) or lateral
(Bunchosia, Hiraea, Stigmaphyllon, Tetrapterys).
Flowers highly stereotyped, showy, yellow
(Fig. 20.4), pink or white with 5 unguiculate petals
(Fig. 20.4) and 2 basal glands on each sepal. Pol-
lination usually carried out by bees. Ovary supe-
rior. Fruits very diverse, often bi- or tri-samaras in
lianas (Fig. 20.8), or drupes (Fig. 20.8). Oily seeds
(Byrsonima) (Fig. 20.5).
Small trees (Byrsonima, AM), shrubs (Bun-
chosia, Malpighia, Thryallis, AM), lianas
Fig. 20.4 Stigmaphyllon ellipticum, unguiculate petals,
Belize
Similar families:
(Banisteriopsis, Heteropterys, Hiptage, Hir-
COMBRETACEAE, when stipules are not
aea, Stigmaphyllon, Tetrapterys, etc.), no
observable (Combretum lianas are often twining
and also have leaves with glands).
herbs.
In AF-MA: Acridocarpus with its glandular leaves Distribution in the tropics and subtropics.
can be confused with EUPHORBIACEAE. Malpighiaceae originated in South America,
with basal genera in Guianan Highlands [5].
Very frequent in Neotropics, much less fre-
quent in Paleotropics (AF-MA: Acridocarpus,
Architectural models badly known:
Philgamia; AS: Hiptage). American savannas
LEEUWENBERG (Fig. 20.5) (a), (a variant of the adapted to periodic fires are typically colonised
model with lateral flowering): Bunchosia lindeniana
by Byrsonima species, small trees with thick,
(AM-C).
FAGERLIND ( Fig. 20.5) (c):Byrsonima spp. (AM).
deeply fissured bark (Fig. 20.6). Hallucino-
TROLL’s model probably absent.
genic plants: ‘ayahusca’ or ‘caapi’: Banisteri-
opsis caapi (Amazonia). Ornamental shrubs
(Malpighia, Thryallis). Fruit trees: Malpighia
emarginata (Fig. 20.8 Barbados cherry).
! Flowers bear oil glands (elaiophores) attracting bees, but these organs are absent or vestigial in
paleotropical species. Understanding the terminology used to describe styles and stigmas is essential
for the identification of species [6].
Fig. 20.5 Architecture. a: LEEUWENBERG’s model (Bunchosia lindeniana), meristem abortion (x); b: MANGENOT?
(Malpighia coccigera); c: branch plagiotropic by apposition, FAGERLIND (Byrsonima spp.)
Fig. 20.6 Leaves, glands and indumenta. a: twining liana, leaves with a pair of glands; b: glandular petioles
Stigmaphyllon sp.; c: leaf margin glandular, Spachea sp.; d: intrapetiolar stipules, Byrsonima spp.; e: hairs appressed,
Malpighia emarginata.; f: Byrsonima crassifolia, fissured bark and pink-coloured inner bark, Venezuela
Fig. 20.7 Byrsonima bucidifolia, pubescent leaves, Belize. Bunchosia lindeniana, venation brochidodromous, not
densely reticulate, Belize. Hiraea sp., leaves opposite, stem and leaves with appressed hairs, stipules inserted on the
petiole, Venezuela
Fig. 20.8 Malpighia emarginata, in cultivation, Thailand, leaves stipulate, apices with appressed hairs and drupe.
Stigmaphyllon ellipticum, di-samaras, Belize
20.3 Malvaceae: 5-6-8-14-15-47-55-B
229 genera and 5100 species (including the former Bombacaceae, Sterculiaceae and Tiliaceae). Order
Malvales. S.t. trees with impressive buttresses (Ceiba, Gyranthera, AM; Heritiera littoralis, AF-E;
Kostermansia, AS; Argyrodendron, AU). Trunks or branches spiny (Bombax, Paleo; Fig. 20.16
Ceiba, Pantrop). Wood often light and soft (many pioneer trees). Plants without latex, leaves and bark
not aromatic. Bark with a network of fibres (Fig. 20.10). Plant with cells containing mucilage
(Fig. 20.9).
20.3 Malvaceae: 5-6-8-14-15-47-55-B 313
Fig. 20.9 Durio zibethinus, ROUX’s model, agroforest, Flores, Indonesia
When monopodial branching in trunk is accompanied by spiral leaf arrangement in branches,

growth is usually strongly rhythmic (Fig. 20.26) (e.g. RAUH’s and AUBRÉVILLE’s models).
Spiral arrangement of leaves: Dombeya, AF; Melochia, AM-AS; Hibiscus, Sterculia (Pantrop) or
two-ranked arrangement (Durio zibethinus, Fig. 20.11 Guazuma, AM), Pterospermum (AS) Theo-
broma (AM). Stipules present, hood-like in a few Hibiscus Fig. 20.20. Leaves simple, oval-shaped
(Fig. 20.15), cordate (Fig. 20.14), palmatilobate (Fig. 20.13), or compound palmate (Adansonia,
Bombax, Fig. 20.16 Ceiba). Heritiera and Sterculia have
simple or compound palmate leaves according to the spe-
cies. Leaves may change in form depending on their
location on the growth unit or the development stage of the
individual (Fig. 20.24 Pterospermum, Fig. 20.25 Stercu-
lia). When branches are plagiotropic, leaf lamina often
asymmetrical at its base (Colona, Grewia, Pterospermum,
Fig. 20.24 Scaphopetalum (Paleo), Luehea, AM). Petiole
distally swollen into a pulvinus. Hairs stellate (peltate in
Durio Fig. 20.17, AS; Mollia, AM). Young leaves
drooping and reddish (Fig. 20.15 Catostemma, Theo-
broma). Venation tripliveined or palmate. Lateral flow-
ering (Apeiba, AM; Dombeya, Grewia, Paleo; Sterculia,
Pantrop) or terminal flowering (Abutilon, Ochroma, AM),
when terminal, leaf-opposite inflorescences or flowers give
the appearance of a lateral flowering (Byttneria, Gossyp-
ium, Pantrop; Abroma, AS-AU; Quararibea, AM). Cau-
liflory frequent in understorey (Cola, AF; Herrania
(Fig. 20.19), Theobroma, AM). Corollas usually contorted
(Fig. 20.20) with radial symmetry, 5 petals (Fig. 20.26)
Fig. 20.10 Inner bark with network of fibres,
and numerous stamens (bilateral symmetry in Helicteres
unidentified, French Guiana
Fig. 20.19). Stamens fused into a tube surrounding the style (Fig. 20.22) (androgynophore in
Bombacoideae, Helicterioideae and Malvoideae, Fig. 20.20), or free (Brownlowioideae,
Grewioideae, Tilioideae). Ovary superior. Fruits capsules (Fig. 20.13 Apeiba, Fig. 20.21 Kleinhovia,
Fig. 20.21 Luehea, Fig. 20.23 Pachira, Sterculia), with a hairy endocarp (Adansonia, Ceiba) a
schizocarp (Malva), a berry (Fig. 20.18 Grewia, Microcos), rarely a samara (Gyranthera). Seeds
covered by very numerous long hairs (kapok) in several Bombacoideae (Adansonia, Ceiba,
Ochroma) or covered by an aril (Fig. 20.15 Catostemma, Fig. 20.17 Durio).
Trees (s.t. very tall or very thick, Adansonia, Ceiba, Fig. 20.19 Gyranthera), medium-sized trees
(Catostemma, Pterospermum, Theobroma), pioneer trees (Grewia, Fig. 20.11 Guazuma), shrubs
(Abutilon, Gossypium, Grewia, Hibiscus), prickly lianas (Fig. 20.15 Byttneria), brambles (Sca-
phopetalum amoenum) or herbs (Malva, Sida, Waltheria).
Architectural models: Similar families:

CORNER: Cola buntingii (AF-W), possibly all LOGANIACEAE and RUBIACEAE also have stems
Herrania species (AM). with an interpetiolar ridge (and stipules in Rubiaceae).
TOMLINSON: Alcea rosea, Hibiscus coccineus.
LEEUWENBERG: Triumfetta cordifolia (Congo),
Sparrmannia africana (AF-S).
Architecture (continued):
KORIBA: Ochroma pyramidale (AM-S), Octolobus CHAMPAGNAT: Scaphopetalum amoenum (AF).
sp. (Gabon). TROLL: Hibiscus grewiifolius (AS), very likely all
FAGERLIND: Quararibea funebris (AM-C). Tilia species, Apeiba burchellii (Guianas), Guazuma
ulmifolia (Fig. 20.11).
PETIT: very likely all species of Gossypium,
Waltheria indica (AF).
SCARRONE: Firmiana fulgens, F. simplex (AS),
Heliocarpus appendiculatus (AM).
RAUH: Adansonia za (MA), Hibiscus tiliaceus (AS-
OC), Cola nitida (AF).
AUBREVILLE: Hildegardia erythrosiphon (AS),
Pentace polyantha (Java), Pseudobombax ellipticum
(AM), Sterculia tragacantha (AF).
ACOSTA: Hibiscus sabdariffa (small shrub, AF),
Urena lobata (pantropical ruderal).
NOZERAN: Theobroma cacao (Fig. 20.27),
T. bicolor, T. speciosum (AM).
MASSART: Heritiera rumphii, Pachira cubensis;
Pterygota alata (AS), Ceiba pentandra, Theobroma
grandiflorum (AM), Desplatsia chrysochlamys (AF).
ROUX: Durio zibethinus (Fig. 20.9 AS),
Trichospermum grewiifolium (Cuba).
Fig. 20.11 Guazuma ulmifolia, pioneer tree of sec-
ondary forest, TROLL’s model, Belize
20.3 Malvaceae: 5-6-8-14-15-47-55-B 315
Fig. 20.12 a: Hibiscus tiliaceus (Seram, Indonesia) and b: Thespesia populnea (Flores, Indonesia), two small trees of
seashores with very similar aspect and morphology
Timber: Heritiera cochinchinensis ‘Mengkulang’, Triplochiton (obeche wood), Guazuma,

Ochroma lagopus (balsa wood) provides a light wood often used by model makers. Fruit trees:
Adansonia digitata (‘baobab’), Durio zibethinus (‘durian’), Cola nitida (kola nut), Theobroma cacao
(cacao tree Fig. 20.27). Pot herbs: Hibiscus esculentus (‘gombo’). Source of fibres: Ceiba pentandra
(‘kapok’), Corchorus olitorius (‘jute’), Gossypium spp. (cultivated cotton plant), Hibiscus spp.,
Thespesia populnea (‘portia tree’), Kydia calycina (‘vendai’). Stimulants: Cola anomala, C. nitida.
! In species belonging to the Malvaceae family, uprighting of stems (TROLL’s model) is not caused
by tension wood but is a consequence of the treillis structure of the fibrous bark, which efficiently
redirects transverse extension into longitudinal tension [7], see also Annonaceae for this phenomenon.
! A remarkable convergence exists in two small seashore trees (Fig. 20.12): Hibiscus tiliaceus
(a) and Thespesia populnea (b), (see also Fig. 20.21, Fig. 20.27).
Fig. 20.13 Abutilon angulatum, shrub with terminal flowering, Botswana. Apeiba sp., leaves with sunken venation,
lateral flowering, Venezuela. Apeiba tibourbou, prickly capsule, Venezuela
Fig. 20.14 Bombax ceiba, MASSART’s model, in cultivation (orig. AS), Botswana. Bombax sp., scalloped bark,
BG LC, France. Byttneria divaricata, liana with spiny petioles and cordate leaves, Venezuela
Fig. 20.15 Byttneria sp., spiny fruits and stems, Southern Brasil. Catostemma commune, fruits with arillate seeds,
Bochinche, Venezuela. Catostemma lemense, drooping young leaves, Sierra de Lema, Venezuela
Fig. 20.16 Ceiba pentandra, spiny trunk, spiny trunk and branches (Belize), leaves palmate (ORSTOM, Cayenne).
Cola marsupium, hairy cordate leaves with domatia, Cameroon
Fig. 20.17 Durio lowianus, bark and underside of leaf with metallic aspect, Khao Chong, Thailand. Durio zibethinus.,
conduplicate leaf and peltate hairs, fruit (a small one! with one arillate seed). Gossypium sp., leaves lobate, in
cultivation, Venezuela
20.3 Malvaceae: 5-6-8-14-15-47-55-B 317
Fig. 20.18 Grewia flava, berries, Botswana. Grewia sp., free stamens forming a tube, Kerala, India. Guazuma
ulmifolia, leaf laminas asymmetrical, Tulum, Mexico
Fig. 20.19 Gyranthera caribensis, buttressed tree, Parque nacional Henri Pittier, Venezuela. Helicteres isora, tubular-
shaped flowers, Seram, Indonesia. Herrania lemniscata, fruit inserted on trunk, Venezuela
Fig. 20.20 Hibiscus macrophyllus, stipule hood-like, FRIM, Malaysia. Hibiscus sp., corolla contorted in bud, Tulum,
Mexico. Hibiscus sp., small stipules, BG Sirikit, Thailand
Fig. 20.21 Hibiscus tiliaceus, leaves cordate, Bali, Indonesia. Kleinhovia hospita, leaves cordate, papyraceous
capsules, Seram, Indonesia. Luehea cf. candida, capsule, Gran Sabana, Venezuela
Fig. 20.22 Malva moschata, androgynophore, Switzerland. Melochia umbellata, young leaves are flat, not folded,
Ruteng, Flores, Indonesia. Melochia sp., glomerulate inflorescences, Belize
Fig. 20.23 Microcos hirsuta, fruits, Kebun Raya, Java. Pachira sp., flowers attracting bats, very numerous stamens
united into a tube, Orinoco Delta, Venezuela. Pachira sp., fallen fruit showing its large seeds, French Guiana
Fig. 20.24 Pterocymbium tinctorium, internal bark with fibrous strips, Kebun Raya, Java. Pterospermum
diversifolium, strong heterophylly, Bali. Scaphopetalum thoneri, asymmetrical leaves, Cameroon
20.3 Malvaceae: 5-6-8-14-15-47-55-B 319
Fig. 20.25 Sterculia cf. pruriens, growth unit (GU) with leaves varying in size and shape, stipules (st) enclosing the
apical bud (b), French Guyana. Sterculia apetala, open capsule, Caruachi, Venezuela. Sterculia coccinea, a red pericarp
bearing black seeds, attracting birds!, Sumatra
Fig. 20.26 Sterculia foetida, leaves compound palmate, Bali, Indonesia. Sterculia lanceolata, leaves simple,
Shipstern, Belize. Sterculia multiovula, flowers with 5 fleshy petals, Las Claritas, Venezuela
Fig. 20.27 Theobroma cacao, Cocoa pods, in cultivation, Bali, Indonesia. Theobroma cacao, tier of five young
branches, Lamanai, Belize. Thespesia populnea looks much alike Hibiscus tiliaceus, but its leaves are not cordate and
the brown spots at the base of the petals are not joined (joined in H. tiliaceus), Flores, Indonesia
20.4 Marcgraviaceae: 13-54
8 genera and 130 species, all American. Order

Ericales. When lianas, these climbing with clamp-
roots. Lenticels s.t. disposed in rows (Fig. 20.29).
Plants dimorphic, the climbing juvenile stage with
two-ranked leaf arrangement (Fig. 20.30), the aer-
ial flowering stage with spiral arrangement of leaves
(Fig. 20.29). Leaves simple, entire or toothed,
obtuse, somewhat clustered on canopy stems.
Stipules absent. Leaves can reflect sonar waves of
bats (Fig. 20.30) [8]. Venation brochidodromous
(Fig. 20.30), somewhat indistinct (Fig. 20.29). Fig. 20.28 Norantea guianensis, red inflorescences
attracting humming birds, detail: flower short-pedicellate
Inflorescences terminal racemes, bearing often with its long coloured bract
bright red bracts (Fig. 20.28 Norantea) or
drooping umbels (Marcgravia Fig. 20.30),
these with fasciculate pitcher-like bracteas. Similar families:
Flowers often pollinated by birds. Fruits ERICACEAE when they are epiphytic..
dehiscent capsules.
Lianas and a few small trees or shrubs (e.g.
Architectures unknown.
Norantea adamantium, Norantea guianensis,
AM-S [9].
Fig. 20.29 Marcgravia sororopania, a. decumbent aerial part, b. lenticels (lent) in rows, c. toothed leaf, Venezuela.
Marcgravia sp., leaves of the aerial stage, leaf-folding convolute, French Guiana. Marcgravia umbellata, umbel of
flowers surrounding stalked pitcher-like nectaries, Guadeloupe
Fig. 20.30 Marcgravia umbellata, fruiting stage, Bains Jaunes, Guadeloupe. Marcgravia umbellata, A: climbing
stage with two-ranked leaf arrangement, B: leaves appressed on support, clamp roots. Marcgravia sp. hollow, ear-
shaped leaf reflecting sonar waves
20.5 Melastomataceae: 35-37-39-41-44 321
20.5 Melastomataceae: 35-37-39-41-44
163 genera and 5600 species. Order Myrtales.

Plants without latex and not aromatic. Bark of
trunks or branches sloughing off in scales
(Fig. 20.33) or strips (Fig. 20.37). Stems not lenti-
cellate. Leaf arrangement opposite, s.t. alternate
because of the reduction of a leaf in each pair
(Fig. 20.38 Sonerila, Miconia dimorphica, AM).
Internodes of horizontal branches can be alternately
twisted (e.g. Mouriri Fig. 20.37). Stipules absent.
Leaves simple and usually entire. Young leaf-
folding revolute in Memecyloideae (Fig. 20.37).
Leaves without translucent dots and no glands. Leaf
bases hollow, inhabited by ants (Fig. 20.35 Maieta,
Tococa). Typical venation (Fig. 20.32) with one or
Fig. 20.31 Medinilla sp., epiphytic shrub with drooping
two pairs of major veins parallel with the midrib and
inflorescences, Flores, Indonesia
Similar groups:
Melastomatoideae are unmistakable as a group, but
Memecyloideae are similar to MYRTACEAE, except LEEUWENBERG (Fig. 20.35 ): Marcetia taxifolia
for their indistinct venation and absence of translucent (Guianas), Memecylon sp. (MA), Medinilla spp. (AS),
dots. Miconia hirta (AM).
KORIBA (Fig. 20.35): Clidemia involucrata,
(Guianas), Memecylon spp. (Fig. 20.31 Paleo).
SCARRONE: Leandra divaricata, Miconia plukenetii,
Tococa guianensis, Meriania sclerophylla (Guianas).
RAUH: Henriettea stellaris, H. caudata, small trees in
Guianas, Marcetia taxifolia (N AM-S).
AUBRÉVILLE: Loreya spp., Topobea (Guianas).
MASSART (Fig. 20.39): Mouriri spp. (AM).
MANGENOT: Memecylon guineense (AF), Mouriri
crassifolia, M. guianensis, small trees, Guianas.
TROLL’s model probably absent.
with parallel tertiary veins forming ladders

(Melastomatoideae), but venation indistinct in
Memecyloideae (Fig. 20.37). Plants pubescent
(Melastomatoideae) or glabrous (Memecyloideae).
Inflorescences in panicles (Fig. 20.36), heads
(Fig. 20.34), terminal (Fig. 20.34) or lateral
(Fig. 20.36). Flowers with radial symmetry
Fig. 20.36, pink, blue, purple or white with 4–5
petals, or 3 petals in Sonerila (AS). Stamens
dimorphic, long and short (Fig. 20.34 Melastoma-
toideae). Pollination carried out by insects. Ovary
inferior. Fruits berries (Fig. 20.36), rarely capsules.
Fig. 20.32 Miconia superba, indument of spread hairs,
Venezuela
Few tall trees (Loreya arborescens, Guianas), small trees (AM: Henriettea, Meriania, Mouriri,
MA: Dichaetanthera, Memecylon (Paleo). ‘Geometrical’ shrubs (Melastoma, Miconia, Rhynchan-
thera, Tibouchina, all AM), often growing in open sites (AM: Dissotis, AS: Sonerila). Various
ericoid species in Guianan Highlands (Fig. 20.35), epiphytes (AM: Blakea, Fig. 20.31 Medinilla,
Fig. 20.34 Phainantha, Topobea parasitica) often associated with other plant species in ant gardens
(AM: Maieta, Tococa), few climbing plants (Maieta spp., AM). Herbs (Aciotis, AM).
The subfamily Melastomatoideae is frequent in South America where several of its members are
aluminium-accumulators. Numerous endemics on the tepuis and the Andes. Memecyloideae are
equally distributed in the three tropical land masses (AM, AF, AS). In spite of more than a thousand
species in Miconia, few are of economic importance, except as ornamentals. Berries are edible, but
fruit trees are few. Memecylon edule provides a yellow dye and Dionycha, a black dye. Miconia
calvescens, originating in Mexico, is a formidable alien which has invaded Tahiti and other Poly-
nesian isles, to the detriment of the native flora.
! Melastomataceae are clearly of Gondwanan origin [10]. TROLL’s model seems to be absent in
this family, which is in favour of a Laurasian origin of this model.
Fig. 20.33 Typical leaves of Melastomatoideae, Belize. Papyraceous bark of a Melastomatoideae, Belize. Adelobotrys
sp., a climbing, not anisophyllous species, French Guiana
Fig. 20.34 Miconia (syn. Clidemia) coriacea, terminal flowering, flowers in heads, Gran Sabana, Venezuela.
Dichaeanthera sp., dimorphic stamens, Madagascar. Henriettea sp., cauliflorous tree, Venezuela
20.5 Melastomataceae: 35-37-39-41-44 323
Fig. 20.35 Maieta sp., leaf bases hollow, inhabited by ants, Venezuela. Marcetia taxifolia, ericoid shrub,
LEEUWENBERG’s model, Gran Sabana, Venezuela. Memecylon sp., young plant conforming to KORIBA’s model
Fig. 20.36 Memecylon ovatum, indistinct venation, Thailand. Miconia rigidiuscula, terminal infructescences, Southern
Brasil. Miconia sp., lateral flowering, Venezuela
Fig. 20.37 Mouriri cf. crassifolia, bark sloughing off in strips, French Guiana. Mouriri cf. exilis, sympodial branch,
leaves with indistinct venation, Belize. Mouriri sp., internodes twisted alternately clockwise and counterclockwise,
young leaves folding revolute, French Guiana
Fig. 20.38 Phainantha myrteoloides, epiphytic shrub, Sierra de Lema, Venezuela. Melastoma malabathricum, flowers
with dimorphic stamens, Belize. Sonerila sp., anisophylly, paleotropics. Tococa sp., leaf bases hollow, inhabited by
ants, Venezuela
Fig. 20.39 Sympodial orthotropic branch of an individual conforming to SCARRONE’s or LEEUWENBERG’s

model. MASSART’s model: rhythmic and plagiotropic branches, e.g. Mouriri spp. Anisophylly associated with
myrmecophylly, base of the larger leaf hollow and housing ants (Tococa)
20.6 Meliaceae: 12-24-48-51-C
48 genera and 700 species. Order Sapindales. Plants usually without latex (whitish exudate in Aglaia
spp.), Leaves and bark not aromatic. Adult trees s.t. with thick fissured bark (Fig. 20.46). Tangential
cuts of trunk or branches often display orange-coloured sclerenchymatous inclusions (Fig. 20.43
Dysoxylum). Spiral arrangement of leaves. Usually (big) trees (Cedreloideae) with stout branch
apices. Growth strongly rhythmic (Cedreloid.: Fig. 20.41 Cedrela, Fig. 20.45 Khaya, Chukrasia,
Toona) or weakly rhythmic (Melioideae: Aglaia, Dysoxylum, Guarea, Fig. 20.46 Trichilia). Stipules
absent. Leaves compound pinnate (Fig. 20.45), often large and leaving shield-like petiolar scars
(Fig. 20.47) (a). Leaves, imparipinnate (Aglaia, Aphanamixis, Carapa, Trichila spp.), paripinnnate
(Cedrela, Swietenia, Toona, Xylocarpus), trifoliolate in Sandoricum, rarely simple (Fig. 20.46
Turraea, Vavaea). Leaflets alternate in Trichilia spp. (Fig. 20.47) (e), s.t. their arrangement variable
in a same individual. Absence of translucent dots and glands, but hydathodes can be observed on the
apices of very young leaflets (Fig. 20.47) (f). Leaflets entire, rarely toothed (Azadirachta). In
Chisocheton (AS, OC) and Guarea (AM), the rachis ends in a bud of leaflets (Fig. 20.44), the bud
aborting or continuing its growth for months or even years. Leaflet venation often camptodromous
(Fig. 20.47) (d). Hairs simple, s.t. stellate or peltate (Aglaia, Lepidotrichilia, Trichilia spp.).
Inflorescences in spikes or cymose. Flowers small with a glandular disk, s.t. with petals forming a
tube. Ovary superior. Fruits are most often capsules. Seeds arillate (Fig. 20.44) in Dysoxylum spp.,
Trichilia spp.), s.t. berries or drupes (Melia azedarach).
20.6 Meliaceae: 12-24-48-51-C 325
Fig. 20.40 Xylocarpus cf. moluccensis, three asynchronous branch complexes: defoliate, chlorophyllous and with
senescent yellow leaves, Flores, Indonesia
RAUH (strongly dominant): Carapa (AM),
Entandophragma, Khaya (AF, MA), Swietenia
(AM).
AUBRÉVILLE: Vavaea (West Pacific).
TROLL: Turraea (Paleo).
Predominantly, trees of dense forests (Carapa,

Entandroprama excelsa, the highest known tree in
Africa, Khaya, Sandoricum, Swietenia, Toona).
Xylocarpus spp. are small trees in paleotropical
mangroves (Fig. 20.40). A few shrubs (Turraea),
a single climber (Turraea laurentii, AF)
Meliaceae are an important element of African
forests, but the impressive trunks of mahoganies
have aroused economic interests (Khaya and
Entandrophragma), they are also abundant in
open heath forest (kerangas) of Borneo and
Malaysia. Other timbers: Cedrela, Chukrasia, Fig. 20.41 Carapa procera, flush of young leaves,
Dysoxylum malabaricum, Melia. Swietenia French Guiana
mahagoni is the American mahogany, but Swi-
etenia macrophylla, easiest in cultivation, is intensely planted in tropical Asia. The most frequent
Meliaceae is probably Azadirachta indica (Fig. 20.42 Neem tree, native to Burma?), introduced in
Africa by Muslim pilgrims and naturalized in many regions of Asia. Neem oil serves for body care
Fig. 20.42 Aglaia argentea, dense indumentum of peltate-stellate hairs on underside of leaflets, Bali, Indonesia.
Azadirachta indica, toothed leaflets and cymose inflorescence, BG Ciudad Bolivar, Venezuela. Chisocheton
cumingianus, young tree, Thailand
Fig. 20.43 Chisocheton siamensis, bud of leaflets, Thailand. Dysoxylum alliaceum, sclerenchymatous inclusions in
bark, Kebun Raya, Java. Dysoxylum sp., capsule, Moluccas, Indonesia
Fig. 20.44 Dysoxylum sp., seeds with chlorophyllous tegument and white aril, Moluccas, Indonesia. Guarea sp., bud
of leaflets, Belize. Lansium domesticum, fruiting tree, Seram, Indonesia
and for medicinal purposes. Fruit trees: ‘sentul’ (Sandoricum koetjape, Malaysia), arils of ‘langsat’
seeds (Fig. 20.44 Lansium domesticum, AS-SE), seed-oil used for candel- and soap-making for Natal
mahogany (Fig. 20.46 Trichilia emetica, AF-C). Gum (Chukrasia tabularis, S-E AS). Anti-
inflammatory and anticancerous substances in Aglaia.
! As far as vegetative characters are concerned, the different families of the Sapindales are difficult
to distinguish. Each feature, appearing at first typical for a family, may exist, as an exception, in other
groups (just say that the supplementary key C covers five pages in order to identify only half a dozen
families!).
20.6 Meliaceae: 12-24-48-51-C 327
Fig. 20.45 Khaya senegalensis, (orig. AF-W), rhythmic growth, fallen leaves in the next to last GU, Sirikit BG,
Thailand. Melia azedarach, raceme of fruits, in cultivation, Maun, Botswana. Swietenia macrophylla, sapling with
paripinnate leaves, Belize
Fig. 20.46 Toona sinensis, thick and deeply fissured bark of an adult tree, Kebun Raya, Java. Trichilia emetica, weak
rhythmic growth, in cultivation, Botswana. Turraea pubescens, entire leaves and corollas with free petals, Bali
Fig. 20.47 Meliaceae. a: Enlarged petiolar bases, large petiolar, shield-like scars (Cedreloideae); b: rhythmic growth,
four consecutive GUs are represented, e.g. Toona sp.; c: trunk apex bearing scale-leaves, in side view and seen from
above; d: leaflet with camptodromous venation; e: leaf bearing alternate leaflets and a terminal leaflet (Trichilia spp.); f:
young leaf of Carapa procera, the leaflet apices with water-secreting glands (hy: hydathodes)
20.7 Menispermaceae: 19-55-57
72 genera and 450 species. Order Ranunculales.

Plants not aromatic. Cross-sections of lianas display
a radial structure with phloem rings alternating with
xylem rings (Fig. 20.48) and a furrowed external
wood. Spiral arrangement of leaves. Stems stiff and
elastic. Leaves simple, rarely palmatilobate
(Fig. 20.51), trifoliolate (Syntriandrium, AF-W), or
peltate (Cyclea), their margin being always entire.
Petioles genicu-
late at both ends
Similar families: None !
(Fig. 20.51).
Venation palmate
or leaves 3-veined
Architectural models little
(Fig. 20.49).
known:
Plants glabrous.
LEEUWENBERG (Fig.
Venation often 20.51): Abuta sp.
Fig. 20.48 Furrowed wood and concentric rays and camptodromous. MANGENOT: Cocculus sp.
rings, e.g. Abuta Inflorescence OLDEMAN: many lianas.
cymose
(Fig. 20.50). Flowers with parts in three. Ovary
with numerous free carpels. Stamens variable in
numbers. Fruits polydrupes (Fig. 20.50). Seeds
with ruminate endosperm.
The majority of the species are woody lianas
(Abuta spp., Chondrodendron, Cocculus, Hyper-
baena) but the family contains also several trees or
shrubs (Abuta spp.). Menispermaceae is a forest-
linked family, reported elsewhere in tropics, but
highly endemic at the genus level: Abuta (AM),
Burasaia (MA), Chondrodendron (AM), Cocculus
(AM-N, AF, AS), Epinetrum (AF), Hyperbaena
(AM), Tiliacora (AF, AS, AU). Menispermaceae
Fig. 20.49 Leaf, drupes and seeds of an unidentified are known for their arrow-poisons (Chondroendron
specimen, Venezuela
tomentosum to prepare curare), medicinal plants
(cycleins of Cyclea peltata).
! Menispermaceae have invented the pulvinate (or geniculate) petiole, a structure not reported in
basal Angiosperms and Magnoliides. Among Monocots, Marantaceae and Dioscoreaceae also have
pulvinate petioles, but anatomically, they differ from those of ‘Dicots’.
20.8 Monimiaceae s.l.: 40-44-45 329
Fig. 20.50 Abuta sp., liana, camptodromous venation, Belize. Epinetrum sp., drupes, Cameroon. cymose inflorescence
of an unidentified species, Venezuela
Fig. 20.51 Menispermaceae. a: trunk cross-section, corky bark; b: twining liana with modular structure (prehensile
extremities; various types of leaves: c: typical leaf; d: palmatilobate leaf (Burasaia, Tinospora); e: shrub conforming to
LEEUWENBERG’s model
20.8 Monimiaceae s.l.: 40-44-45
28 genera and 210 species, including Atheros-

permataceae and Siparunaceae. Order Laurales.
Plants aromatic (Siparuna) or not aromatic
(Kibara, Mollinedia, etc.). Inner bark with
(Kibara, Matthaea) or without (Glossocalyx,
Mollinedia, Siparuna) a network of fibres. Main
stems have immediate branching. Periderm
remains green for several consecutive GUs
(Fig. 20.52 e.g. Ephippiandra madagascarensis,
Madagascar). Extremities of stems somewhat
flattened (Fig. 20.53). Opposite leaf arrange-
ment, looking two-ranked in Glossocalyx
because of the reduction of one leaf in each pair Fig. 20.52 Ephippiandra madagascariensis, stem with
green periderm, Madagascar
(Fig. 20.54). Leaves simple, entire (Matthaea,
Similar families:
LAURACEAE when bearing opposite
leaves with pinnate venation.
RAUH: Hedycarya tapeinospermifolia
(OC).
MASSART: Siparuna crassifolia (AM).
MANGENOT: Ephippiandra (MA),
Matthaea (AS), Mollinedia, Siparuna
guianensis (AM).
ATTIMS: Nemuaron veillardii (NC).
Fig. 20.53 Typical: flattened stem apex
ROUX: Glossocalyx longicuspis (AF).
TROLL’s probably absent.
Tambourissa) or toothed (Fig. 20.55), teeth spaced (Kibara,
Mollinedia, Ephippiandra), s.t. with laticiferous threads
(Tambourissa, Fig. 20.55 Xymalos). Venation brochidodro-
mous with secondary veins forming angular arcs (Fig. 20.55), see also Chloranthaceae. Indument of
simple hairs, s.t. hairs stellate or grouped in fascicles (Palmeria, Siparuna spp.). Inflorescences
lateral (Fig. 20.55). Flowers small, unisexual, regular. Ovary with numerous free carpels. Stamens
variable in numbers (up to 2000 in Tambourissa!). Fruits are fascicles of drupes (Fig. 20.54), s.t.
plunged in the receptacle.
Essentially trees (but these never tall) (Hedycarya, Peumus, Xymalos), shrubs (Glossocalyx,
Kibara, Tambourissa) and some lianas (Palmeria), no herbs.
Monimiaceae are of East-gondwanan origin [11], mostly represented in Australasia and Oceania.
American and African genera are few, AM: Mollinedia, Siparuna, AF: Glossocalyx, Xymalos.
Endemism is high at the genus level Ephippiandra (MA), Glossocalyx (AF-W), Hedycaraya (West
Pacific), Kibara (AS, AU), Laurelia (NZ, Chili, Peru), Matthaea (Malaysia), Monimia (Mauritius and
Réunion), Nemuaron (New Caledonia), Palmeria (NG, AU), Peumus (Southern Chile), Siparuna
(AM), Tambourissa (MA and Mascarene), Xymalos (AF-S), etc. Monimiaceae are strictly associated
with forests and their immediate vicinities where they are somewhat inconspicuous. Some plants are
used for dyes (Kibara moluccana) or medicinal purposes (Peumus boldus (orig. Chile), Siparuna).
Fig. 20.54 Glossocalyx longicuspis, strong anisophylly, Campo forest, Cameroon. Glossocalyx longicuspis, inflated
branch bases and reduced leaf (rl) in each pair. Kibara coriacea, fasciculate carpels, Java
20.10 Moraceae: 2-4 331
Fig. 20.55 Mollinedia sp., leaves toothed, secondary veins forming angular arcs, Belize. Secondary veins forming
angular arcs, laticiferous threads (e.g. Xymalos). Siparuna thecaphora, lateral inflorescences, Belize
20.9 Montiniaceae: 44
Very small family of 3 genera and 5 species.

Order Solanales. Trees and shrubs with a pep-
pery scent. Whorled leaf arrangement. Leaves
simple. Stipules absent. Flowers small, unisex-
ual, 3- or 4-merous. Fruits capsules containing
Similar families:
CORNACEAE also have a
camptodromous venation.
GENTIANACEAE, but almost
absent in Madagascar.
winged seeds. Architecture sympodial and mod- Fig. 20.56 Grevea madagascariensis, leaves clustered at
ular (LEEUWENBERG’s model). AF-S and branch extremities, Madagascar
Madagascar (Fig. 20.56).
20.10 Moraceae: 2-4
40 genera and 1200 species. Order Rosales. Formerly Moraceae included at the subfamily rank
Cecropioideae (i.e. Cecropia-Pourouma-Musanga), a group actually placed in Urticaceae. Plants not
aromatic. Trunks often exhibiting small buttresses (Fig. 20.60) or producing a profusion of
adventitious roots (Fig. 20.59, e.g. Ficus), these often anastomosed. Several Ficus species are
stranglers (Fig. 20.57). Bark often fibrous. Spiny trees uncommon, e.g. Poulsenia armata (AM).
Many species are deciduous, but not necessarily in the course of the dry season. Latex aqueous, in
bark and leaves, usually white, s.t. opalescent (Fig. 20.58), pinkish or beige, usually not toxic (but see
Antiaris further). Monopodial architecture. Spiral arrangement of leaves (Artocarpus, Fig. 20.62
Ficus spp.), or two-ranked arrangement (Brosimum, Ficus spp., Fig. 20.64 Pseudolmedia, Trophis),
very rarely leaves opposite (Bagassa). Stems lenticellate. Stipules always present and s.t. hood-like
(Fig. 20.60), leaving an annular scar on the stem (Brosimum, Ficus, Perebea). Leaves simple, shortly
petiolate, entire (Ficus spp.), toothed (Morus),

s.t. lobate (Fig. 20.59 Artocarpus, Fig. 20.61
Clarisia, Ficus spp.). Petioles not pulvinate dis-
tally. Venation brochidodromous (Fig. 20.65) or
leaves tritripliveined (Fig. 20.62 Ficus spp.).
Venation very densely reticulate (s.t. with
domatia (Fig. 20.65). Dioecious trees (Ficus).
Inflorescences lateral, often on specialized
shoots inserted on the trunk in Artocarpus
(Fig. 20.60) and Ficus (Fig. 20.63). In Artocar-
pus, inflorescences in heads, then the numerous
flowers swell to form a, s.t. huge, compound
fruit. Inflorescences pauciflorous in Prainea
(Fig. 20.64). In Ficus, fruit peduncles swell and
develop into a hollow receptacle enclosing
numerous minute flowers. In Dorstenia flowers
are inserted on a flat receptacle (Fig. 20.61).
Fruits tiny drupes (Ficus), somewhat large dru-
pes (Brosimum, Prainea). Trees (Antiaris, Arto-
carpus, Brosimum, Castilla, Ficus, Maclura,
Milicia excelsa ‘Iroko’, Morus, Sorocea, Stre-
blus, Trophis), shrubs (Ficus), s.t. epiphytes Fig. 20.57 An epiphytic Ficus has overtopped its host
and killed it, Thailand
Similar families:
When latex is observable, no possible
confusion.
CORNER: Dorstenia gigas (treelet,
Socotra).
RAUH (dominant): Artocarpus altilis, A.
integrifolia (AS), Ficus spp.
ROUX: Antiaris, Castilla, Milicia excelsa,
Broussonetia papyrifera (AS, OC).
AUBRÉVILLE: Ficus ischnopoda (AS).
CHAMPAGNAT: Maclura pomifera (AM-
N, Morus alba (China, cult. EU, AM-N).
TROLL (dominant): Artocarpus spp.,
Brosimum, Ficus spp., Milicia excelsa,
Sorocea, Streblus, Trophis.
with roots at first hanging, then reaching the soil

(strangler fig trees). Herbs only in Dorstenia and
Fatoua. Several species are completely decidu-
Fig. 20.58 Aqueous and whitish latex, an unidentified ous (Antiaris, Morus alba, white mulberry).
specimen, French Guiana
20.10 Moraceae: 2-4 333
Moraceae are essentially tropical forest trees present on all land masses, predominantly in Asia.
Uses are numerous. Timber: AF: Milicia excelsa (‘iroko’), AF: Antiaris toxicaria (‘ako’). Rubber:
Castilla, Ficus. Dyes: Maclura pomifera. Fibres: Poulsenia armata (AM). Broussonetia papyrifera
(AS) has been introduced by Austronesian people as far as Polynesia and the Easter Islands to make
clothes and fishing nets. Fruit trees: Artocarpus altilis (breadfruit tree), A. integrifolia (jackfruit tree),
Brosimum, Ficus spp. (fig trees), Prainea limpato (‘kesusu, Malaysia’). Cardiotonic arrow poisons
(Antiaris toxicaria, Paleo). The general purpose tree of the Mayan civilisation: Brosimum alicastrum
(‘ramón’), a much appreciated tree, still provides wood, fibres, edible leaves, latex and seeds.
! Many species of deciduous Moraceae, among them Ficus spp., Broussonetia payrifera have a
chlorophyllous layer between the inner bark and the thin rhytidome. It is very probable that these trees
take advantage of cortical photosynthesis during the dry season [12].
Fig. 20.59 Radicular system of a tree dug up from an embankment (Ficus indica), Bali, Indonesia. Artocarpus
gomezianus, copious milky latex, Thailand. Artocarpus altilis, Breadfruit tree, Bali
Fig. 20.60 Artocarpus heterophyllus, stipule hood-like, Thailand. Artocarpus heterophyllus, fruits inserted on trunk,
and WIFI available over here! Seram, Indonesia. Brosimum alicastrum, buttressed tree, Belize
Fig. 20.61 Brosimum alicastrum, venation brochidodromous, Belize. Clarisia illicifolia, lobate leaf, French Guiana.
Dorstenia sp., herb, its inflorescence inserted on a flat receptacle bearing two linear expansions, Cameroon
Fig. 20.62 Ficus thonningiana, spiral leaf arrangement, Botswana. Ficus sp., leaves tripliveined, Belize. Ficus
auriculata, fruits inserted on trunk, in cult. BG LC, France
Fig. 20.63 Ficus sp., drooping branches bearing fruits, Seram, Indonesia. Ficus sp., lateral flowering (small figs),
Venezuela. Ficus sp., trunciflory, Flores, Indonesia
Fig. 20.64 Morus alba, CHAMPAGNAT’s model, leaves toothed, Thailand. Prainea limpato, lateral infructescence
of drupes. cf. Pseudolmedia sp., two-ranked leaf arrangement, venation brochidodromous, Venezuela
20.11 Moringaceae: 48 335
Fig. 20.65 Trophis mexicana, two-ranked leaf arrangement, venation brochidodromous, Belize. Buttressed tree, Ficus
sp., Flores, Indonesia. Ficus sp., venation densely reticulate, two main lateral veins with domatia, Sumatra
20.11 Moringaceae: 48
A small monogeneric family of 13 species. Order Brassicales. Subarid regions of Africa and Asia.
Small trees with tuberous rootstock. Bark with smell of mustard-oils and a network of fibres.
Leaves 1–3 compound pinnate (Fig. 20.66). Leaflets entire. Conspicuous glands on rachis nodes (1).
Flowers scented, irregular (i.e. without radial symmetry Fig. 20.66). Fruits are woody pods. Seeds
3-winged.

LEGUMINOSAE with nipple-like glands, e.g. OLDEMAN: because branching process is too irregular for
Senna!, but their leaves are not stipulate and considering CHAMPAGNAT’s model [72].
crushed bark has a different smell.
Moringa oleifera (Horseradish tree of Northern India or, in Africa, ‘Nebedies’ for ‘never dies’ tree)
is widely naturalized and cultivated. Uses are many, among them cooking oil, vegetable, pesticide,
medicinal. When crushed, moringa seeds can help to purify dirty water, by flocculation of proteins.
Fig. 20.66 Moringa oleifera. Flowers with bilateral symmetry, 5 sepals and 5 petals, Bali. Part of a 3x-compound leaf
(two opposite pinnae), glands on rachis node, Java
20.12 Muntingiaceae: 6
A tiny family with 3 genera and 3 species. Order Malvales. Branches plagiotropic with asymmetrical
leaf bases (TROLL’s model), Fig. 20.67. Inflorescences lateral, 1-flowered. Muntingia calabura (Jam
tree or ‘Calabura’) is a small American tree cultivated everywhere in the tropics as a shade tree.
‘Calabura’ is a perpetual flowerer but with flowers lasting only one day. Fruits are berries with
numerous seeds.
! Muntigiaceae are very similar to Malvaceae, as “Tiliaceae” or Grewia. Observing their non-
pulvinate petioles, one will not fall into the trap of mistaking them for Malvaceae.
Fig. 20.67 Muntingia

calabura. Solitary flower and
leaves two-ranked,
asymmetrical
20.13 Myricaceae: 30
4 genera, 50 species. Order Fagales. Aromatic trees. Latex absent. Nitrogen-fixing Actinomycetes in
roots (Morella). Stems lenticellate (Fig. 20.68). Branching weakly rhythmic (i.e. without short
internodes and scale-leaves). Spiral leaf arrangement. Stipules absent. Leaves toothed-crenelate,
shortly petiolate (Fig. 20.68). Venation somewhat indistinct. Inflorescences lateral, in spikes.
Flowers small, wind pollinate. Fruits drupes or nutlets. Morella and Canacomyrica are tropical in
their distribution. Some are sources of wax.

BORAGINACEAE and SYMPLOCACEAE have RAUH: Morella (Fig. 20.68) (AM, Macaronesia, AS),
leaves with distinct venation and are not aromatic. Canacomyrica (New Caledonia), Myrica
(N temp.).
Fig. 20.68 Morella cerifera. RAUH’s model, stem lenticellate and toothed leaves, Belize
20.14 Myristicaceae: 3-12 337
20.14 Myristicaceae: 3-12
20 genera and 500 species. Order Magnoliales. Plants not aromatic, sectioned bark producing a red or
reddish exudate (Fig. 20.70). The exudate tasting astringent (polyphenols). Branches distinctly
disposed in tiers (Fig. 20.69). Two-ranked leaf arrangement. Stipules absent. Leaves simple, entire,
often chewed by insects. Leaves often glaucous beneath. Venation pinnate, not densely reticulate.
Apices covered by a rust-coloured indument (Fig. 20.71). Lateral flowering (Fig. 20.72), inflores-
cences cymose or racemose. Flowers small, 3–5 petals. Fruits fleshy or coriaceous berries enclosing
one seed, the seed surrounded by a red aril (Fig. 20.72). Seeds with endosperm presenting complicate
folds. Fruits taken by birds, which eat the arillas but regurgitate the seeds, these being toxic.
Similar families:
EBENACEAE: when a red exudate not observable (Diospyros),
underside of leaves are often glaucous in both families.
MASSART’s model is exclusive in this
family (Fig. 20.69 ).
Trees, never very tall (Horsfieldia, Knema, Iryanthera,

Myristica, Pycnanthus, Virola), a single species of liana
(Pycnanthus dinklagei).
Essentially in tropical rainforests of all land masses.
Several species grow in swampy sites (Iryanthera, Virola).
Cephalosphaera (AF-W), Knema (AS), Iryanthera (AM),
Myristica (AS, AU), Pycnanthus (AF), Virola (AM). Tim-
ber: Cephalosphaera usambarensis (AF-E), Pycnanthus
angolensis (AF-W). Seeds of Pycanthus angolensis
(Ilomba), Horsfieldia spp., Virola sebifera, etc. used by soap
factories and for the fabrication of candles. Seeds of nutmeg
trees (Myristica fragrans) are used for seasoning, although
they are toxic when eaten in large quantities. Several Virola
species (‘epéna’, ‘nyakwana’) are used for hallucinogenic
snuffs by Amazonian forest dwellers.
! Myristicaceae are a relictual family containing very
similar species, they are considered as the sister group of the
remaining Magnoliales, i.e. Magnoliaceae, Annonaceae,
Himantandraceae and Degeneriaceae [13]. These two last
families, encompassing only 2 genera and 4 species, all Fig. 20.69 Virola michelii, reiterating
MASSART’s model, Fr. Guiana
endemic to Australasia and Oceania, are not described in this
book.
Fig. 20.70 Knema furfuracea, red exudate, Thailand. Myristica fragrans, capsule and an arillate seed, Seram. Typical:
buds and young leaves hairy
Fig. 20.71 Horsfieldia irya, Seram, hanging fruits, Indonesia. Iryanthera sp., aqueous red exudate, French Guiana.
Knema sp., leaves chewed by insects, their underside bronze-coloured, Seram
Fig. 20.72 cf. Knema sp., fallen tree with tiers of branches (MASSART’s model), two-ranked leaf arrangement,
secondary veins regularly spaced, Khao Chong, Thailand. Pycnanthus sp., lateral inflorescences, Cameroon
20.15 Myrtaceae: 4-23-39-40-41-42-44 339
20.15 Myrtaceae: 4-23-39-40-41-42-44
134 genera and 5500 species. Order Myrtales. Plants with leaves s.t. aromatic and always without
latex. Bark sloughing off in scales (Fig. 20.77) or in strips (Fig. 20.79). Trunk and branches not
lenticellate in Myrtoideae, lenticellate in several Leptospermoideae (Leptospermum laevigatum).
Inner bark usually not fibrous (fibrous in Leptospermum spp. Figure 20.80). Early suberization of
the periderm of the last formed GU (Fig. 20.73), turning ochre-coloured (Fig. 20.77). Leaf
arrangement opposite (Myrtoideae) or spiral in the adult tree (Leptospermoideae). Internodes can
twist alternately on plagiotropic twigs (Fig. 20.80 Pimenta dioica, Psidium guajava). Stipules absent
(very minute in Psidium). Leaves simple, shortly petiolate, entire, usually with translucent dots
visible against light. Venation brochidodromous, often with one or even two submarginal veins
(Fig. 20.77), (e.g. Eugenia, Syzygium). Inflorescences lateral (Fig. 20.80) (usually racemose or
greatly reduced and appearing glomerulate, fasciculate in Eugenia and Calyptranthes, Myrciaria,
AM), axillary panicles (Myrcwia, AM), terminal cymes or panicles (numerous Syzygium), uniflorous
(Fig. 20.78 Feijoa, Psidium spp., AM). Clusters of flowers mimicking a spike but flowers lateral on
monopodial stems (Fig. 20.78 Callistemon). Flowers with 4–5 petals and numerous stamens
(Fig. 20.81). Ovary with axile placentation (Fig. 20.75). Fruits berries (Fig. 20.79 Eugenia), cap-
sules (Leptospermoideae), drupes or nuts.
Trees (Calyptranthes, Eugenia, Syzygium,
Similar families: Eucalyptus, Myrcia), shrubs (Callistemon, Psid-
ium), several lianas and stranglers
LYTHRACEAE (Lagerstroemia): translucent
dots absent. (Metrosideros), no herbs.
MELASTOMATACEAE - Memecyloideae: Although Myrtaceae are easily recognized,
venation indistinct. identification at the genus level is difficult when
relying only on vegetative characters. Although a
lot of useful vegetative keys help to identify
species at a local level, botanists need more
Architectural models (Fig. 20.76): general identification systems. This situation
KORIBA: Eugenia spp. (E. buxifolia?), concerns especially Brazilian savannahs
Syzygium cumini (India, cult.). (‘restingas’, ‘campos cerrados’) and Andean
RAUH: Syzygium myrtifolium, S. malaccense. mountain ranges where Myrtaceae-Myrtoideae
ATTIMS : Eucalyptus spp. (Moluccas, are highly diversified.
Australia).
MANGENOT? : Eugenia lambertiana (West
Indies).
TROLL: Psidium spp. (AM).
Treelets of the understorey exhibit a modular

architecture (Eugenia or Syzygium, or both?)
where two branches develop at the trunk extrem-
ity, one of the pair will become the leader, the
other will grow obliquely or even horizontally
(Fig. 20.76). This structure is easily visible from a
distance and provides a reliable feature to identify
Myrtaceae; moreover, it also exists in Mouriri, a
genus of the Memecyloideae (Melastomataceae).
Myrtaceae exist in all kinds of shrubby or Fig. 20.73 An unidentified Myrtaceae, flaky rhyti-
wooded environments, especially in the Southern dome, stem with early suberization, Belize
Fig. 20.74 Cloves (Syzygium aromaticum, ‘Cenke’) in drying process, Ambon, Indonesia (yr. 1990)
Hemisphere but is poorly represented in tropical

Africa. Myrtoideae are pantropical but Lep-
tospermoideae represent essentially an Aus-
tralasian group. Timber or paper pulp: Eucalyptus.
Essential oils: Eucalyptus, Melaleuca. Spices:
cloves are dried flower buds of Syzigium aro-
maticum (Fig. 20.74). Pimenta dioica provides
‘allspice’ or Jamaica pepper. Fruit trees: Feijoa
sellowiana (‘pineapple Guava’), Psidium guajava
(guayava tree), Syzygium malaccense (Malay ap-
ple), Syzygium cumini (‘jambolan’).
! Architectures of Myrtoideae are almost
Fig. 20.75 Psidium guajava, axile placentation unknown in light of the two thousand species of
Fig. 20.76 KORIBA’s model, one branch of each tier becomes the leader (e.g. Syzygium cumini). Syzigium
myrtifolium, monopodial architecture, young individual growing in shade with tiers of plagiotropic branches, then
becoming erect in sunny places (RAUH’s model), cult., Bali. Eugenia lambertiana, MANGENOT’s model?,
Guadeloupe. Apex abortions (x)
20.15 Myrtaceae: 4-23-39-40-41-42-44 341
Fig. 20.77 Papyraceous rhytidome, an unidentified Myrtaceae, Belize. Ocre-coloured young bark, an unidentified
Myrtaceae, Thailand. Typical: intramarginal veins
Fig. 20.78 Feijoa sellowiana, flowers solitary, stamens with red filaments, BG INRA-Antibes. Eucalyptus sp., young
tree conforming to ATTIM’s model. Callistemon sp., crowded flowers mimicking a spike, but each of them axillate by a
very reduced leaf (a terminal UC bears normal leaves), in cultivation
Fig. 20.79 cf. Eugenia buxifolia, internal rhytidome ochre-coloured, bark brown-reddish and foliage, dry forest,
Belize. Eugenia uniflora, berries, in cultivation, Southern Brazil. Eucalyptus deglupta, bark sloughing off in strips,
Seram, Indonesia
the Eugenia-Syzygium complex and, in (sub)tropical America, other very similar species of Caly-
colpus, Calyptranthes, Campomanesia, Marlierea, Myrcia, Myrciaria, Myrcianthes, Myrteola,
Siphoneugena, etc. In other words, in order to understand how the different species grow and develop,
a huge amount of field-work remains to be done, a task requiring only pencils and notebooks.
Fig. 20.80 Leptospermum laevigatum, scaly bark, inner bark fibrous, Flores. Pimenta dioica, internodes angular
alternately twisted, Belize. Psidium guineense, lateral flowering, Venezuela
Fig. 20.81 Syzygium cumini, submarginal vein, cultiv., Thailand. Syzigium malaccense, flowers with numerous
stamens, cult., Venezuela. Syzygium? sympodial branch, opposite leaf arrangement, Thailand
References
1. Nooteboom, H.P. 1985. Notes on Magnoliaceae, with a revision of Pachylarnax and Elmerrillia and the Malesian
species of Manglietia and Michelia. Blumea 31: 65–121.
2. Sangtae, K., P. Chong-Wook, K. Young-Dong, and S. Youngbae. 2001. Phylogenetic relationships in family
Magnoliaceae inferred from ndhF sequences. American Journal of Botany 88: 717–728.
3. Wang, Y.-B., B.-B. Liu, Z.-L. Nie, H.-F. Chen, F.-J. Chen, R.B. Figlar, and J. Wen. 2020. Major clades and a
revised classification of Magnolia and Magnoliaceae based on whole plastid genome sequences via genome
skimming. Journal of Systematics and Evolution 58: 673–695.
4. Hallé F. 2004. Architectures de plantes, 162 pp. JPC Edition.
5. Davis, C.C., P.W. Fritsch, C.D. Bell, and S. Matthews. 2004. High-latitude Tertiary migrations of an exclusively
tropical clade: Evidence from Malpighiaceae. International Journal of Plant Sciences 165 (Suppl. 4): 107–121.
6. Anderson, W.R. 2001. Malpighiaceae. In Flora of the Venezuelan Guayana, vol. 6., 804 pp. St. Louis (Missouri):
ed. J.A. Steyermark, P.E. Berry, K. Yatskievych, and B.K. Holst.
7. Clair, B., B. Ghislain, J. Prunier, R. Lehnebach, J. Beauchêne, and T. Alméras. 2019. Mechanical contribution of
secondary phloem to postural control in trees: The bark side of the force. New Phytologist 221: 209–217.
8. Simon, R., M.W. Holderied, C.U. Koch, and O. von Helversen. 2011. Floral acoustics: Conspicuous echoes of a
dish-shaped leaf attract bat pollinators. Science 333: 631–633.
9. Da Silva Júnior, M.C., & B.A. Da Silva Pereira. 2009. 100 Árvores do cerrado - matas de galeria, 288 pp. Brasilia:
Rede do Sementes do Cerrado.
10. Renner, S.S. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic Journal of
Botany 13: 519–540.
References 343
11. Renner, S.S., J.S. Strijk, D. Strasberg, and C. Thébaud. 2010. Biogeography of the Monimiaceae (Laurales): A role
for East Gondwana and long-distance dispersal, but not West Gondwana. Journal of Biogeography 37: 1227–1238.
12. Girma, A., A.K. Skidmore, C.A.J.M. Bie, F. de Bongers, and M. Schlerf. 2013. Photosynthetic bark: Use of
chlorophyll absorption continuum index to estimate Boswellia papyrifera bark chlorophyll content. International
Journal of Applied Earth Observation and Geoinformation 23: 71–80.
13. Sauquet, H., J.A. Doyle, T. Scharaschkin, T. Borsch, K.W. Hilu, L.W. Chatrou, and A. Le Thomas. 2003.
Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: Implications for character
evolution. Botanical Journal of the Linnean Society 142: 125–186.
14. Harden, G., B. McDonald, J. Williams. 2006. Rainforest trees and shrubs: a field guide to their identification in
Victoria, New South Wales and subtropical Queensland using vegetative features, xxxiv + 266 pp. Gwen Harden
Publishing, Nambucca Heads.
Nyctaginaceae
21
21.1 Nyctaginaceae: 36-41-44-56
27 genera and 390 species. Order Caryophyllales. Plants not aromatic and
without latex. External wood often furrowed (Fig. 21.3b). Modular archi-
tecture with clustered leaves (Fig. 21.4). Several species bear short shoots
modified into spines (Bougainvillea, Pisonia, Fig. 21.3d). Leaf arrangement
spiral (Bougainvillea), pseudowhorled (Guapira fragrans) or opposite
(Neea, Fig. 21.5 Pisonia). Stems of herbaceous species are often swollen at
nodes (Mirabilis, Boerhavia). Stipules absent. When leaf arrangement is
opposite, the two leaves of the same pair often unequal in size, Fig. 21.3c,
(anisophylly). Leaves simple, shortly petiolate, entire. Underside of leaves
often glaucous. Venation pinnate, not densely reticulate, or even indistinct.
Apices and buds covered by a rust-coloured indument of short hairs
(Fig. 21.4). Inflorescences lateral, cymose (Guapira, Neea, Pisonia). Flow-
ers small, forming a 3–5–8 lobate short tube, s.t. rimmed by showy bracts
mimicking sepals (Fig. 21.2 Bougainvillea). Ovary superior, unilocular.
Fig. 21.1 KORIBA’s
Fruits achenes or small nuts, often hidden by the perigon (Fig. 21.1). model
Fig. 21.2 Bougainvillea spectabilis, two different cultivars
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346 21 Nyctaginaceae
Fig. 21.3 a CHAMPAGNAT’s model; b external wood furrowed; c opposite leaves and anisophylly; d short twigs
modified into spines
Trees (Guapira, Fig. 21.4 Neea, Pisonia spp.), shrubs (Guapira cuspidata), climbing or spiny
plants (Pisonia aculeate), herbs (Fig. 21.6 Boerhavia, Mirabilis).
Nyctaginaceae are centered on the Neotropics with Pisonia extending to Asia. They are more
frequent in semi-deciduous forests and there is a dramatic radiation of genera in the arid regions of
North America. A few species are adapted to saline environments and grow in upper shores of Pacific
islands (Pisonia grandis). Pisonia grandis ‘Alba’ (Moluccan cabbage, ‘kol banda’, lettuce tree).
Bougainvillea and Mirabilis jalapa (marvel of Peru) are ornamentals grown everywhere in warm
climates (Fig. 21.6).
! Several members of the Nyctaginaceae host ectomycorrhiza, like Pisonia grandis growing on
seashores rich in guano. The complex nutritional requirements and the unusual distribution of the
latter species may be related to breeding populations of birds living in these islands [1].
Fig. 21.4 Guapira fragrans, lenticellate twig, pseudowhorled leaf arrangement, Guadeloupe. Neea sp., sympodial
branches, cymose inflorescences, venation indistinct, rust-coloured indument, Venezuela. Neea sp., anisophylly, Tulum,
Mexico
Fig. 21.5 Neea sp., modular architecture, Tulum, Mexico. Pisonia sp., anisophylly, Kebun Raya, Java. External wood
furrowed (Pisonia or Neea), inner bark with a chlorophyllous layer, Venezuela
Reference 347
Fig. 21.6 Boerhavia sp., herb with sympodial, ramified inflorescences (overtopping a papilionoid), Indonesia
LEEUWENBERG: Guapira cuspidata
Similar families: (Guianas); Mirabilis jalapa (AM), Boerhavia
spp. (AM, subcosmopolitan).
OLEACEAE but these do not exhibit anisophylly.
KORIBA (Fig. 21.1): Guapira fragrans, Neea
(AM), Pisonia (AM, AS).
CHAMPAGNAT: Bougainvillea (AM, cult.).
Reference
1. Ashford, A.E., and W.G. Allaway. 1982. A sheathing mycorrhiza on Pisonia grandis R.Br. (Nyctaginaceae) with
development of transfer cells rather than a Hartig net. New Phytologist 90: 511–519.
Ochnaceae to Oxalidaceae
22
22.1 Ochnaceae: 8-9-18-34-50
33 genera and 585 species. Order Malpighiales. Plants not aromatic and without latex. Bark s.t. thick
(Fig. 22.3). Branching distinctly rhythmic (Fig. 22.2). Leaf arrangement spiral (Fig. 22.1 Ces-
pedesia, Elvasia, Lophira), two-ranked (Ochna, Ouratea) or opposite/whorled (Lacunaria, Quiina),
(Fig. 22.1). Stipules intrapetiolar (Fig. 22.1). Leaves simple, shortly petiolate (Fig. 22.2), entire or
toothed, rarely compound leaves (sapling of several Quiinoideae). A very peculiar venation is
observed: raised on upper side of the leaf, and tertiary or even secondary veinlets more or less parallel
(scalariform venation: Cespedesia spathulata Fig. 22.1b). Quiina leaves exhibit a ‘finger-print’
venation (Fig. 22.1c). Plants glabrous with leaves shiny and coriaceous. Inflorescences usually ter-
minal panicles (Fig. 22.4 Poecilandra), Flowers showy, yellow or white, with 5 petals (Fig. 22.4) and
numerous stamens. Fruits often a group of 4–5 drupeoles on a fleshy receptacle (Fig. 22.3 Ochna,
Ouratea) or dehiscent berries in Quiinoideae (Froesia, Lacunaria, Fig. 22.4 Quiina, all AM).
CORNER: Campylospermum spp.(AF).
CHAMBERLAIN (Fig. 22.1c):
Euthemis leucocarpa (Malaysia), Lacunaria
sp. (AM).
LEEUWENBERG: Cespedesia spathulata,
Similar families: (N AM-S), Luxemburgia gaudichaudii
(Brazil), Quiina sp. (Guianas),
As a rule, no possible confusion with other families . Schuurmansia henningsii (New Guinea).
KORIBA: Lophira alata (N AM-S).
SCARRONE: Poecilandra sp. (AM-S).
MASSART (Fig. 22.1a):Ochna kirkii
and Ouratea affinis (AF), Ouratea spp.
(AM).
https://doi.org/10.1007/978-3-031-05942-1_22
350 22 Ochnaceae to Oxalidaceae
Trees, never very tall (Lophira), shrubs (Ochna, Ouratea, Quiina, Sauvagesia), woody herbs
(Sauvagesia, Indovethia)
Ochnaceae are highly diversified in tropical America (Cespedesia, Luxemburgia, Ouratea, etc.). In
savannas, trunks of trees develop deeply fissured bark (Fig. 22.3). Numerous species of Poecilandra,
Sauvagesia and Tyleria are endemics of Guianan highlands. Lophira (only two species) are African
trees bearing large leaves. In the Guianas, Cespedesia spathulata, although smaller, exhibits the same
features. A single pantropical genus: Ouratea. Sauvagesia erecta has become a pantropical ruderal.
! A number of dykes in the Netherlands are still lying on rot-proof trunks of ‘azobé’, or ‘ekki’
(Lophira alata), wood of this species was also used for harbour facilities and sluice doors [1].
! The monotypic genus Medusagyne was recently included in Ochnaceae (leaves opposite, not
stipulate and curious septicidal capsules), M. oppositifolia is an endangered species endemic to
Seychelles.
Fig. 22.1 a MASSART’s model; bud with scale-leaves (sc-l), stipule (st), toothed leaf margin and venation raised on
upper side (Ochna, Ouratea). b KORIBA (Lophira), leaf with scalariform venation (Cespedesia). c CHAMBER-
LAIN’s model (Lacunaria, Quiina), leaves in whorls of four, finger-print venation (Quiina)
Fig. 22.2 Cespedesia spathulata, long cuneate leaves, Venezuela. Indovethia calophylla, a woody herb bearing a
curiously bent inflorescence axis, Pasir Mayang, Sumatra. Ochna integerrima, scaly buds, Thailand
22.2 Olacaceae (incl. Schoepfiaceae): 12-13-21 351
Fig. 22.3 Ochna integerrima, bark thick and fissured, adapted to periodic fires, Northern Thailand. Ochna kirkii,
‘Mickey Mouse’ fruit, BG Singapore. Ouratea sp., yellow flowers, Venezuela
Fig. 22.4 Poecilandra retusa, terminal inflorescence, Venezuela. Quiina sp., baccate fruits, Venezuela. Quiina sp.,
stipules, French Guiana
22.2 Olacaceae (incl. Schoepfiaceae): 12-13-21
23 genera and 160 species. Order Santalales. Plants not aromatic and without latex. Trunk s.t. fluted
(Fig. 22.5). Stems thorny in Ximenia americana (Fig. 22.7). Leaves spiral (Fig. 22.7 Ximenia) or
two-ranked (Aptandra, Coula, Schoepfia). Stipules absent. Leaves simple, entire, shortly petiolate,
not coriaceous. Venation usually not densely reticulate (Fig. 22.7), camptodromous in Minquartia
(Fig. 22.6). Plants glabrous. Inflorescences lateral. Flowers small with a cupuliformous perianth in
3–6 parts. Ovary superior with 2-3-5 fused carpels. Fruits drupes (Fig. 22.7) or capsules.
Similar families: ROUX: Coula edulis (AF), Ximenia
americana (AM, naturalized in paleotropics).
ERYTHROPALACEAE, SANTALACEAE, two
NOZERAN: Minquartia guianensis (N AM-S).
other families of the order Sapindales.
TROLL: Aptandra (AM, AF).
ICACINACEAE, however with coriaceous leaves.
MANGENOT: Okoubaka aubrevillei (AF).
CHAMPAGNAT: Ximenia caffra (AF-S).
Trees (Ochanostachys, Ongokea), shrubs (Olax, Ximenia), lianas (Anacolosa), root parasitic
(Olax), no herb
Mainly African (Anacolosa, Ximenia caffra) and American (Cathedra, Dulacia, Ximenia ameri-
cana) but also Asiatic (Ochanostachys), with about ten monospecific genera (e.g. Coula, AF,
Maburea, AM, Minquartia, AM, Ongokea, AF). Olacaceae grow in dense forests, rarely in savannas
(Ximenia). Timber: Minquartia guianensis, Ochanostachys amentacea, Ongokea gore. Fruit trees
(Anacolosa spp., Coula edulis, Ximenia caffra, all AF). Oil seeds: Ongokea gore (AF), for manu-
facturing lacquers, varnishes and linoleum.
! At first sight, this family looks somewhat undistinguished because species do not show any
striking features (i.e. leaves simple, entire, not stipulate, almost no latex, and so forth). However their
stems undergo suberization which is patchy at early stages (see also Heisteria, Erythropalaceae and
Lepionurus, Opiliaceae).
Fig. 22.5 Fluted trunk, Minquartia guianensis Fig. 22.6 Minquartia guianensis, camptodromous venation
Fig. 22.7 Ongokea gore, capsules with yellow arils, Cameroon. Ximenia americana, ROUX’s model, thorny stems.
Ximenia americana, venation indistinct, observe the praying mantis, Flores, Indonesia
22.3 Oleaceae: 35-40-44-45-51-52-56-57 353
22.3 Oleaceae: 35-40-44-45-51-52-56-57
25 genera and 565 species. Order Lamiales. Plants not aromatic and without latex. Bark multilayered
(Fig. 22.13). Sclerenchymatic inclusions in bark (e.g. Fig. 22.13 Forsythia). Suberized stems
showing protruding, almost circular, lenticels (Fig. 22.8). Internodes not angular, except in
(Fig. 22.10) Myxopyrum and Nyctanthes. As a rule, interpetiolar ridge missing (Fig. 22.11), but
present in Myxopyrum, Nyctanthes and Jasminum spp. Leaves opposite, rarely spiral (Jasminum
sect. Alternifolia). Stipules absent.
Leaves simple, shortly petiolate, entire or toothed, compound pinnate (Fraxinus, Jasminum) or
trifoliolate (Jasminum). Venation brochidodromous (Fig. 22.9), tripliveined (Fig. 22.11) in Myx-
opyrum. The underside of L. covered by very numerous minute trichomes (hand lens 20x !).
Domatia in vein axils (Jasminum spp.). Inflorescences cymose. Flowers regular, usually with 4
petals and 2 stamens, or with 6 petals (Fig. 22.9 Jasminum, Nyctanthes), corolla tubular Jasminum).
Ovary superior, bicarpellate. Fruits drupes (Chionanthus, Ligustrum, Fig. 22.11 Noronhia, Olea),
berries (Fig. 22.10 Myxopyrum), capsules (Schrebera) or samaras (Fraxinus).
Fig. 22.8 Chionanthus microstigma, protruding lenticels, Thailand
Fig. 22.9 Chionanthus microstigma, multilayered bark, and leaves with brochidodromous venation, Thailand.
Jasminum elongatum, Thailand. Interpetiolar ridge, liana and corollas with 6 petals
Fig. 22.10 Jasminum fluminense, a tubular corollas, b scandent shrub, Maun, Botswana. Jasmimum sp., liana,
trifoliolate leaves, thin and linear sepals (corollas shed), Madagascar. Myxopyrum smilacifolium, stems angular, not
lenticellate, berry, Thailand
Fig. 22.11 Myxopyrum smilacifolium, leaves tripliveined, Thailand. Noronhia sp., drupes, Madagascar. Olea
madagascariensis, interpetiolar ridges absent, Madagascar

LAMIACEAE-VERBENACEAE but internodes quadrangular. LEEUWENBERG: Syringa vulgaris.
MELASTOMATACEAE: Memecylon spp. have a similar aspect. PREVOST: Myxopyrum smilacifolium.
Trees (Chionanthus, Noronhia, Olea), never very tall, shrubs (Syringa), lianas, twining (Jas-
minum) or scandent (Fig. 22.10 Jasminum, Myxopyrum), no herbs.
Mainly African and Asian in their distribution (Noronhia, Olea), where they grow in warm
temperate or tropical climates. Chionanthus (incl. Linociera) is pantropical. The mainly temperate
genus Fraxinus reaches equatorial latitudes (Fig. 22.12 F. griffithii). Olea europaea (Olive tree) is
native to the Middle East. Timber (Fraxinus, Olea), wood of Olea capensis holds the record for
highest density (1,4 kg/dm3). Numerous ornamental shrubs (Jasminum, Osmanthus, Syringa). Plants
used for perfumes (Jasminum, Nyctanthes).
! Oleaceae are very scattered in rainforests and for this reason, difficult to spot. Could this rarity be
due to a slow growth related to high density wood?
22.4 Opiliaceae: 13-30 355
Fig. 22.12 Fraxinus griffithii, bark smooth, Flores, Indonesia. Linociera (syn. Chionanthus) sp., sympodial
architecture, doubtfully TROLL, Sumatra. Olea salicifolia, sympodial architecture, doubtfully a TROLL’s model,
Thailand. Olea europaea, absence of interpetiolar ridge, stems lenticellate
Fig. 22.13 Syringa vulgaris, 4-petals flowers, young trunk lenticellate, multilayered dead bark and internal bark.
Forsythia intermedia, orange sclerenchymatous inclusions in bark, in cultivation
22.4 Opiliaceae: 13-30
A small family of 10 genera and 32 species. Order Santalales. Plants not aromatic and without latex.
Bark scaly or, in old trees, fissured (Fig. 22.14a, b). Patchy suberization of the periderm (Fig. 22.14).
Leaves spiral (Agonandra, Fig. 22.15d Meliantha) or two-ranked (Fig. 22.14 Cansjera, Lepionu-
rus). Stipules absent. Leaves simple, soft, entire, shortly petiolate. Venation somewhat indistinct, not
densely reticulate (Fig. 22.15c). Inflorescences in spikes (Fig. 22.14), panicles or umbels. Flowers
small, perianths cupuliform with parts in 4 or 5. Ovary superior. Fruits drupes. Oily seeds.
Similar families:
ERYTHROPALACEAE and Architecture almost unknown:
OLACACEAE, two other families of the CHAMPAGNAT: Cansjera rheedei (AS).
order Santalales.
Trees (Agonandra, Melientha) and shrubs (Lepionurus), these s.t. climbing (decumbent and spiny
in Cansjera Fig. 22.14), no herbs.
Opiliaceae are centered in the paleotropics (Cansjera, Lepionurus, Melientha, Opilia), only one
genus is American (Agonandra), they grow in deciduous forests or savannas. Timber: Agonandra
brasiliensis, oil seeds and medicinals.
Santalales present few distinctive vegetative features. Try this trick: when a patchy suberization of
the periderm is observed, one probably has a member of the Olacaceae-Opiliaceae-Santalaceae.
Fig. 22.14 Cansjera rheedei, Thailand. Short shoots modified into spines, patchy suberization of the periderm,
Thailand. Leaves two-ranked. Lepionurus sylvestris, inflorescences in spikes, Thailand
Fig. 22.15 a Agonandra brasiliensis, cracked bark, Venezuela. b Melientha suavis, fissured bark, Thailand.
c Melientha suavis, venation indistinct, not densely reticulate. d Branches decumbent with leaves spirally arranged,
Thailand
22.5 Oxalidaceae: 27-48-49-54 357
22.5 Oxalidaceae: 27-48-49-54
5 genera and 565 species, of which some 500 Oxalis. Order Oxalidales. Plants not aromatic and
without latex. Leaf arrangement spiral (Fig. 22.16) Averrhoa bilimbi) or two-ranked (Fig. 22.17)
Averrhoa carambola). Stipules absent. Leaves compound pinnate (Averrhoa, Biophytum), trifoliolate
(Oxalis, Fig. 22.18 Sarcotheca) with entire leaflets. ‘Simple’ leaves of Sarcotheca monophylla and
Dapania spp. have an articulation between the petiole and the lamina and can be considered as
unifoliolate. Leaflets moving in response to stimuli (i.e. pulvinate petiolules, Biophytum, Sar-
cotheca). Venation often with a pair of ascending secondary veins (Fig. 22.18). Main vein sunken on
upper side of the leaf. Trunciflory in Averrhoa bilimbi and A. carambola (Fig. 22.17, Fig. 22.18).
Flowers with 5 free petals and 10 stamens. Ovary superior. Fruits capsules or berries (Averrhoa,
Sarcotheca).

CONNARACEAE much CORNER (Fig. 22.18):
like Sarcotheca, Biophytum.
however, the former
TROLL: Averrhoa carambola.
usually lianescent.
LEGUMINOSAE, but OLDEMAN?: Averrhoa bilimbi.
stipulate.
Small trees (Averrhoa), shrubs (Sarcotheca, Malaysia and Sulawesi), lianas (Dapania, MA,
Malaysia) or herbs (Biophytum). Many Oxalis species in South America and Cape province.
Pantropical distribution.
Fruit trees: Averrhoa bilimbi (‘bilimbi’, cucumber tree) and A. carambola (‘carambola’, star fruit),
two species of uncertain origin, Malaysian or Brazilian? Potherb: Oxalis tuberosa (‘oqa’), an edible
tuber in the Andean region. Condiments (Sarcotheca spp.).
Fig. 22.16 Averrhoa bilimbi, flush of young leaves, Fig. 22.17 Averrhoa carambola, TROLL’s model and
Bali trunciflory
Fig. 22.18 Biophytum sp., unbranched herb, Madagascar. Averrhoa bilimbi, trunciflory, flowers and unripe fruits,
Thailand. Sarcotheca sp., leaves trifoliolate, with basal ascending veins
Reference
1. Van de Kluien, J.W.G., and H.J. Blass. 2015. Mechanical properties of azobé (Lophira alata). Holz als Roh- und
Werkstoff 63: 1–10.
Pandaceae to Putranjivaceae
23
23.1 Pandaceae: 8-A
A small family with 3 genera and 16 species, all paleotropical. Order Malpighiales. Panda oleosa
(AF-W) is a tree conforming to ROUX’s model. Leaves two-ranked (Fig. 23.1), simple, entire,
apically mucronate. Inflorescences small cymes (Microdesmis) or cauliflorous thyrses (Fig. 23.2
Panda oleosa). Fruits drupes (Fig. 23.2) enclosing one stone. Seeds of Panda oleosa provide
cooking oil.
Fig. 23.1 Microdesmis sp., leaves two-ranked Fig. 23.2 Panda oleosa, redrawn from ‘Die natürliche
Pflanzenfamilien’ (Engler & Prantl). a Drupe; b cauliflorous
thyrse; c seed
https://doi.org/10.1007/978-3-031-05942-1_23
360 23 Pandaceae to Putranjivaceae
23.2 Pandanaceae: 1-54 (Monocotyledons)
4 genera (5 if Benstonea is segregated from Pandanus [1, 2] and 575 species. Order Pandanales,
Monocotyledons. Impressive adventicious roots (Fig. 23.3). Trunk with branching often continuous
(Fig. 23.4a). Spirotristichous leaf arrangement (Fig. 23.4b). Leaves simple, entire, coriaceous,
bearing marginal spines, up to 5 m in length (Pandanus). Buds present at leaf axils (absent in
Arecaceae!). Venation parallel. Inflorescences are terminal panicles (Sararanga) or 1-n spadices
subtended by coloured spaths. Flowers small with very numerous stamens. Ovary superior con-
sisting of 10–80 uniovulate carpels (multiovulate in Freycinetia). Fruits berries (Freycinetia, Sar-
aranga) or drupes, these often grouped in cylindric to globose ‘phalanges’ (Fig. 23.4c) Pandanus,
Fig. 23.4d Freycinetia). Pachycaulous trees (Pandanus), shrubs (Pandanus) or lianas (Freycinetia).
Pandanaceae grow in paleotropics and Oceania. They are known for their spectacular, very ‘ge-
ometric’, habit (screw pines). Fibres for thatching, flavourings (Pandanus amaryllifolius), fruit trees
(Karuka nut, New Guinea).
! Pandanaceae look like Palms, but are more complex in structure. As Monocotyledons, Pan-
danaceae are architecturally very diversified [3], perhaps due to the existence of axillary buds and a
peculiar stem anatomy where xylem and phloem vessels are arranged in 2 or 3 discreet vascular
strands [4].
Fig. 23.3 Pandanus sp., adventicious roots, back-shore, Flores, Indonesia

23.3 Passifloraceae: 6-7-17-22-55-57 361
Fig. 23.4 a Pandanus sp., trunk with continuous branching, Seram, Indonesia. b Pandanus sp., compound fruit,
BG FIBV, Venezuela. c Pandanus sp., spirotristichous leaves, Madagascar. d Freycinetia sp., fascicle of female
inflorescences, Papua, herbarium specimen, CBG H
23.3 Passifloraceae: 6-7-17-22-55-57
29 genera and 920 species. Order Malpighiales. Plants not aromatic and without latex. Geoxylic
(Fig. 23.6 Adenia) or climbing plants. Passiflora bear tendrils at leaf axils (Fig. 23.7).
Ancistrothyrsus bear tendrils on their inflorescences (Fig. 23.6) Hollow stems inhabited by ants in
Barteria (Fig. 23.7). Leaves spiral (Passiflora) or two-ranked (Barteria, Fig. 23.6 Paropsia, Pas-
siflora). Stipules present. Leaves very variable in shape, simple, lobate (Fig. 23.6), trifoliolate
(Fig. 23.7), their margins entire or toothed.
Fig. 23.5 Passiflora foetida, 3 fringed sepals, Bali

Venation pinnate (Barteria, Fig. 23.9 Turnera) or palmate, sometimes leaves tripliveined (Pas-
siflora). Glands in stipular position (Barteria) or inserted on petioles or on laminas (Passiflora,
Paropsia, Turnera, Fig. 23.7). Passiflora glands mimic egg laying, which discourage insects to lay.
Inflorescences lateral (Fig. 23.6). Flowers large, showy (Fig. 23.8), with 3 sepals (Fig. 23.5), 5 or
10 free petals and often 5 stamens. Carpels topping gynophore, style with 3 stigma (Fig. 23.5
Passiflora). Ovary superior rimmed by a crown of staminodes (Fig. 23.8). Fruits berries (Fig. 23.9
Passiflora) or capsules.
Fig. 23.6 Adenia glauca, bulbous plant, leaves trifidous, BG Monaco. Ancistrothyrsus tessmanii, lateral inflorescences
ending in a hook, a b, minute prophylls with axillary peduncles, Brazil, from a herbarium specimen, CBG H.
Paropsia edulis, a: plagiotropic branch with lateral inflorescences, b: fascicle of flowers, from herbarium specimens
(Madagascar), CBG H
Fig. 23.7 Barteria sp., ROUX’s model, branches inhabited by ants (see the small apertures). Passiflora sp., axillary
tendril, Belize. Passiflora sp., trifoliolate leaves with glands on petioles, Belize
Fig. 23.8 Passiflora caerulea, leaves trilobate, staminodes blue, Belize. P. glandulosa, leaves entire, French Guiana.
P. vespertillo, leaf delta-shaped, flower with five short and five long petals, Venezuela
23.4 Pentaphylacaceae: 12-23-29-30 363
Fig. 23.9 Passiflora cf. boenderi, leaves bilobate, Venezuela. Passiflora sp., berries with coriaceous pericarp,
Venezuela. Turnera scabra, five free petals, toothed leaves, Venezuela
Similar families:
SALICACEAE (compared with Barteria).
ROUX (Fig. 23.7): Barteria fistulosa (AF).
VITACEAE have also tendrils, but these are
oppositifoliate. ATTIMS: Turnera ulmifolia (AM).
Small trees (Barteria, AF, Passiflora magnoliifolia, Colombia), shrubs (Turnera, AM) s.t.
tuberous (Adenia, Paleo), lianas (Passiflora, Ancistrothyrsus, Adenia) or herbs (Passiflora, Turnera).
Almost all Passiflora are native to tropical America, but several species are naturalized or even
invasive (Pacific islands). Fruit lianas (Passiflora edulis). Numerous ornamentals (Adenia, Passiflora,
Turnera).
Passiflora (Passion flower) was given its name by Carl Linnaeus because the form of the floral
organs had been thought to symbolize the crucifixion of Jesus Christ [5].
23.4 Pentaphylacaceae: 12-23-29-30
12 genera and 340 species. Order Ericales. Plants not aromatic and without latex. Leaf arrangement
spiral (Fig. 23.10 Ternstroemia) or two-ranked (Fig. 23.11 Adinandra, Fig. 23.12 Eurya, Freziera).
Stipules absent. Leaves simple, entire (Adinandra, Ternstroemia) or toothed (Fig. 23.12 Eurya).
Young leaf-folding involute (Fig. 23.12) (Adinandra, Freziera). Venation pinnnate, often indistinct
(Fig. 23.13 Ternstroemia). Underside of leaf with (glandular?) trichomes (Anneslea, Fig. 23.13
Ternstroemia). Plants glabrous (Ternstroemia) or apices with appressed hairs (Fig. 23.11 Freziera).
Inflorescences lateral (Fig. 23.10). Flowers large, with free petals (Fig. 23.11). Ovary superior 5-
carpellate. Fruits berries or woody capsules.
Small trees (Adinandra, Eurya, Freziera), shrubs (Ternstroemia), no lianas and no herbs.
Pentaphylacaceae are poorly represented in Africa (Balthasaria), the larger genus, Ternstroemia,
being native to Asia and America. Uses are few (timber, ornamentals).
Fig. 23.10 Ternstroemia gymnanthera, leaves clustered, lateral flowering (red line), Thailand
Fig. 23.11 Adinandra dumosa, two-ranked leaf arrangement, Kebun Raya, Java. Adinandra, Freziera, involute
folding and petiole with appressed hairs. Anneslea fragrans, fleshy flowers, Thailand
Fig. 23.12 Eurya acuminata, erect form of a TROLL’s model, toothed leaves, Flores, Indonesia
23.5 Phyllanthaceae: 2-3-7-8-27-47 365
Fig. 23.13 Ternstroemia tapezapote, spiral leaf arrangement, venation indistinct and minute trichomes, Belize

PENTAPHYLACAEAE are very similar to RAUH: Anneslea fragrans (AS), Ternstroemia
THEACEAE and both families were formerly united. tapezapote (AM-C), T. gymnanthera (AS).
ROUX: Adinandra dumosa (AS).
TROLL (Fig. 23.12): Eurya acuminata (AS).
23.5 Phyllanthaceae: 2-3-7-8-27-47
57 genera and 2000 species. Order Malpighiales. Plants not aromatic and without latex (possibly brownish
exudate in Uapaca bark). Young bark or periderm exhibiting thin longitudinal slits (Fig. 23.14).
Sclerenchymatous inclusions in bark in some species (Fig. 23.16). Twigs supple. Leaf arrangement
usually two-ranked, but also spiral (Fig. 23.21 Bischofia, Fig. 23.14e Hieronyma, Fig. 23.19 Richeria,
Uapaca). Numerous Phyllanthus species have plagiotropic branches mimicking compound leaves
(phyllomorphic branches Figs. 23.15 and 23.21b–d). Leaves almost always simple and entire (trifo-
liolate in Bischofia Fig. 23.21a). Stipules present (s.t. foliaceous Fig. 23.20) Venation pinnate, usually
camptodromous (Fig. 23.16), secondary veins ending in a fimbrial vein in Bridelia (Fig. 23.17d). Foliar
glands almost always absent. Inflorescences usually in lateral position (4–13). Cauliflory in Baccaurea
(Fig. 23.17a), Maesobotrya (Fig. 23.18), Sauropus (Fig. 23.20). Flowers reduced, with free petals.
Ovary superior 3-carpellate, each carpel enclosing 2 ovules (Fig. 23.18). Fruits capsules (winged in
Hymenocardia), s.t. with explosive dehiscence, berries (Fig. 23.18) or schizocarps.
Trees (Antidesma, Aporosa, Baccaurea (AS), Bischofia (AS), Uapaca (stilt roots (Fig. 23.20),
shrubs (Phyllanthus, Pantrop) or herbs (Phyllanthus).
Phyllanthaceae have a pantropical distribution, they grow in forests, savannas and along trails
(many are ruderal). Uses not many considering the large number of species, but some fruit trees:
Phyllanthus acidus (West Indian gooseberry), Baccaurea angulata (‘belimbing darah’). A supposed
aphrodisiac: Richeria grandis (‘bois bander’ of the West Indies). Phyllanthus emblica is a sacred tree
of India and Sri Lanka, its fruits are edible (Indian gooseberry, ‘amalaki’, ‘buah amla’), for healing
oil, herbal juice. Medicinal: Antidesma ghaesembilla (black currant tree, AS), and the very toxic
Cleistanthus collinus (AS, AU). Bischofia javanica spread by Polynesians (dye for tapa cloth).
! Unlike for many Euphorbiaceae, the sister family, leaves of Phyllanthaceae are almost never
glandular, except Phyllanthus vaccinifolius (Fig. 23.19) (AM-S) and Hymenocardia spp. (AF-C).
! Immediate-basal branching would be exceptional (see Introduction), but it exists in Phyllanthus
juglandifolius (Fig. 23.21). However, branches of this species are functionally like leaves, so the
plant can escape the rule.
Fig. 23.14 Fissured periderm, Sauropus sp., Thailand Fig. 23.15 Phyllanthus niruri, herb with phyllomor-
(diam. 15 mm) phic twigs
Fig. 23.16 Antidesma cf. montanum, TROLL’s model?, Bali. Aporosa villosa, venation camptodromous, axillary
spikes of minute flowers, Thailand. Aporosa cf. villosa, sclerenchymatous inclusions in bark, Thailand
Fig. 23.17 a Baccaurea javanica, trunciflory, Sumatra. b Breynia cf. coronata, ROUX’s model, Bali. c Bridelia
stipularis, leaves with a fimbrial vein, Thailand. d Glochidion perakense, bark and inner bark, leaves with
camptodromous venation, Thailand. e Hieronyma sp., foliaceous stipules, Venezuela
23.5 Phyllanthaceae: 2-3-7-8-27-47 367
Fig. 23.18 Maesobotrya sp., trunciflory, Cameroon. a–b–c, Phyllanthus emblica. a «hammered» trunk, Thailand.
b Infructescences inserted laterally on a branch, Belize. c Berry enclosing six seeds
Fig. 23.19 Phyllanthus vaccinifolius, reiterated ROUX’s model, leaf with a single gland on its underside, Venezuela.
Richeria grandis, RAUH or AUBRÉVILLE’s model, Guadeloupe
Fig. 23.20 Sauropus thorelii, cauliflory, Thailand. Uapaca sp. Stilt-roots and foliaceous stipules, Campo forest,
Cameroon

EUPHORBIACEAE, these also have supple twigs with thin ROUX (dominant): Glochidion (AS, OC), 16
longitudinal slits (but carpels are uniovulate). Phyllanthus (Pantrop), Sauropus (AS, AU).
SALICACEAE when these have entire leaves. RAUH: Bischofia javanica (Indomalesia), 17 Richeria
grandis (or AUBRÉVILLE’s?, AM-S, WI), Uapaca
(AF, MA).
AUBRÉVILLE: Baccaurea javanica (AM).
TROLL: Bridelia stipularis (AS), Cleistanthus
polyphyllus (AS), Antidesma thwaitesianum (AS).
Fig. 23.21 a Bischofia javanica, leaves spirally arranged, trifoliolate, FRIM, Malaysia. b Phyllanthus mimosoides,
scars of fallen phyllomorphic twigs, Guadeloupe. c–d Phyllanthus juglandifolius, Venezuela. c: phyllomorphic
branches, in situ; d branching immediate leading to plagiotropic branches, these, however not arranged in tiers, but
evenly disposed on trunk GUs
23.6 Physenaceae: 30
A monogeneric family of 2 species, Madagascar. Order Caryophyllales. Spiral leaf arrangement.

Leaves not stipulate, glabrous. Venation pinnate, not densely reticulate. Flowers in axillary racemes.
Fruits are inflated and 1-seeded (Fig. 23.22). A very bitter plant used in household medicine (an-
timicrobial flavanones).
Fig. 23.22 Physena madagascariensis, inflated fruits, Montagne d’Ambre, Madagascar

23.7 Phytolaccaceae (incl. Petiveriaceae): 30-54 369
23.7 Phytolaccaceae (incl. Petiveriaceae): 30-54
Two families totalling 13 genera and 50 species, all native to America, but several species have
become cosmopolitan. Order Caryophyllales. A curious pair of families encompassing trees, shrubs,
small lianas (Seguieria and Trichostigma spp.) and herbs (Phytolacca americana). Trees with soft
wood, brittle branches, simple and entire leaves. Succession of concentrical cambia in Phytolacca
dioica (“anomalous” secondary thickening). Prophylls? modified into spines in Segieria (Fig. 23.24).
Petiveria, Rivina and Seguieria have been excluded from PHYTOLACCACEAE to be placed in
PETIVERIACEAE.
Ombú trees or ‘Umbu’ (Phytolacca dioica) grow in dry deciduous forests of Southern AM-S, the
thick trunk reiterates profusely (Fig. 23.23). Phytolacca americana is a large herb (Fig. 23.24)
conforming to LEEUWENBERG’s model, ruderal in regions with mild winters. Rivina humilis
(Fig. 23.24) is very frequent in the Caribbean region. Petiveria alliacea is a cosmopolitan herb of
open ground which has a smell of garlic. Fruits of Phytolacca, Rivina (inkweeds) are sources of dyes.
Ombú (Fig. 23.23) is thought to have evolved from a herbaceous ancestor, this «inverted» evo-
lution occurred due to the formation of successive cambia (see Sect. 30.3).
Fig. 23.23 Phytolacca dioica, ‘Umbu’, with its sprouting massive trunk, in cultivation, France
Fig. 23.24 Phytolacca americana, ruderal herb, London, UK. Rivina humilis, herb, terminal flowering, berries, Belize.
Seguieria paraguayensis, spines in stipular position, Kebun Raya, Java
23.8 Picrodendraceae: 31-33-44-49-52-A
25 genera and 105 species. Order Malpighiales. Trees and shrubs without latex. Bitter bark and
strong rhythmic growth in Picrodendron (WI). Leaf arrangement usually spiral, s.t. opposite
(Oldfieldia), s.t. whorled (Mischodon). Stipules absent or minute. Leaves simple (Austrobuxus,
Malaysia, OC, Mischodon, India, Sri Lanka), trifoliolate (Picrodendron, AM) or compound palmate-
trifoliolate (Fig. 23.25 Oldfieldia africana, AF). Ovary with axile placentation. Fruits capsules with 2
ovules per locule.
Many endemic genera, but the family is cosmopolitan. In Australasia Austrobuxus is easily
confused with Chionanthus (Oleaceae) and in tropical America Piranhea longepedunculata
(Fig. 23.25) has a buttressed trunk and branches resembling those of Tabebuia (Bignoniaceae).
Fig. 23.25 Piranhea longepedunculata, buttressed trunk. Venezuela. Oldfieldia africana, opposite trifoliolate leaves
and capsule (cp) enclosing seeds (s)
23.10 Piperaceae: 13-20 371
23.9 Pinaceae: 1-37 (Gymnosperms)
Family encompassing 12 genera and 225 species. Gymnosperms. Pinus species native to tropics are
few, e.g. P. caribaea (Fig. 23.26a–c, a two-leafed pine) in West Indies and Central America,
P. merkusii (Fig. 23.26d a three-leafed pine) in Indomalesia. All Pinus conform to RAUH’s model.
! Never confuse the angiosperms Casuarina (Casuarinaceae) with pines!
Fig. 23.26 Pinus caribaea (a–c) a Landscape with Pinus caribaea, Cockscomb, Belize. b twig and leaves. c Fallen
cone. d Pinus merkusii, Thailand
23.10 Piperaceae: 13-20
5 genera and 3600 species. Order Piperales. Plants aromatic tasting spicy. Stem nodes or base of
internodes swollen in Piper (Fig. 23.29). Leaves spiral (Peperomia, Pothomorphe) or two-ranked
(Piper). Branches sympodial (Fig. 23.30c). Leaves simple, entire. Petioles canaliculate, bearing s.t. a
pair of appendages looking like stipules (Fig. 23.30d). Venation pinnate or leaves tripliveined.
Flowering terminal (Fig. 23.27 Piper) or lateral (Fig. 23.30e Pothomorphe). Flowers small, much
reduced, in terminal (looking lateral) spikes. Fruits one-seeded berries or drupes.
Fig. 23.27 Piper aduncum, terminal flowering, French Guiana (native to Asia)
TOMLINSON: Pothomorphe peltata (N AM-S).
PETIT (Fig. 23.30a), mainly Piper species.
MANGENOT: Lepianthes umbellatum (Pantrop).
Shrubs and very numerous herbs (Lepianthes, Peperomia, Piper, Pothomorphe), rarely small
trees. Climbing Piper are exceptional (Fig. 23.29). Piperaceae grow in all tropical rainforests,
Peperomia being much more frequent in highlands and cloud forests. Spices or condiments:
P. nigrum (black pepper), P. longum (Bengal pepper), Piper cubeba (Java pepper), P. lolot (syn.
P. sarmentosum (‘lalot’ of vietnamese recipes), P. retrofractum (Fig. 23.28 Javanese chilli). Stim-
ulants: Piper betle (Fig. 23.28), Piper methysticum (‘kawa’, Vanuatu Islands).
Fig. 23.28 Piper betle (leaves), Areca catechu (red fruits) and Uncaria gambir (Rubiaceae, fruits yellow) serve to
prepare betel, Sumatra. Piper sp., spikes of flowers, Belize. Piper retrofractum, Bali
Fig. 23.29 Piper sp., swollen base of an internode, Thailand. Piper sp., swollen nodes of a trunk of a climbing species,
Seram, Indonesia. Piper sp., petiole canaliculate, Belize
Fig. 23.30 Piper spp. a PETIT’s model; b terminal spike of reduced flowers; c false stipules (fs) and canaliculated
petiole; d modular and sympodial construction of a branch (each module with the first internode very short and a long
second internode (long IN) ending in an inflorescence; e Pothomorphe peltata, a herb with lateral flowering, Venezuela
23.12 Poaceae (Bambuseae): 1 (Monocotyledons) 373
! TROLL’s models possibly absent in Piperales and all other paleoherbs [6] given the absence of
an uprighting mechanism which is specific to this model.
23.11 Pittosporaceae: 29
9 genera and 210 species. Order Apiales. Plants not aromatic, without latex, but producing s.t. gums
or mucilage. Spiral leaf arrangement with leaves clustered in module extremities (Fig. 23.31b).
Stipules absent. Leaves simple, entire. Venation pinnate, brochidodromous. Flower bases tubular
with 5 petals and 5 stamens (Fig. 23.31c). Ovary superior. Fruits berries (Fig. 23.31a, orange-
coloured in Auranticarpa, blue in Billardiera) or capsules. Seeds surrounded by an aril or a viscid
pulp, but not in Auranticarpa [7]. Winged seeds in Hymenosporum.
Fig. 23.31 a Auranticarpa sp., (syn. Pittosporum sect. Auranticarpa), orange berry, Flores, Indonesia.
b–c Pittosporum tobira, leaves clustered, flowers yellow. d Pittosporum sp., KORIBA’s model, in cultivation

PENTAPHYLACACEAE (Ternstroemia), but their KORIBA (Fig. 23.31d): Auranticarpa, Pittosporum.
leaves have an indistinct venation. FAGERLIND: Hymenosporum flavum (AU-E).
Trees (Hymenosporum, Pittosporum), shrubs (Pittosporum) or lianas Billardiera (AU

extratropical).
Pittosporaceae grow in Madagascar and Asia, with a majority of species in Australia.
! Robert Brown, Scottish botanist and discoverer of “brownian motion”, named the family because
of the pitchy (i.e. ‘pittos’ in Greek) viscous pulp of Pittosporum fruits.
23.12 Poaceae (Bambuseae): 1 (Monocotyledons)
750 genera and 11,000 species. Order Poales. Bambuseae are the only tribe in the family with
arborescent species (Fig. 23.32, i.e. bamboos), the remaining species named «grasses» in the
common sense. Bamboos are about 20 cm up to 40 m in height, with stem tissues containing silica
Fig. 23.32 A widely cultivated bamboo, cf. Gigantochloa sp., Flores, Indonesia. Large foliar sheaths inserted in each
node, Sakaerat, Thailand
Fig. 23.33 Dendrocalamus sp., two collateral stems with basal scaly leaves, Thailand. Phyllostachys nigra, two
collateral stems, cult. Chusquea sp., clusters of ramified short twigs, Santa Catarina, Brazil
bodies which make them very hard. Bamboos are rhizomatous plants conforming to TOMLINSON’s
(sympodial rhiozomes) and BELL’s models (monopodial rhizomes). Stems, or trunks, are hollow
except at nodes but Chusquea stems are full of pith. Young axis ramify readily, then undergo a fast
intercalary growth. Two-ranked leaf arrangement. Basal leaves reduced into scales (Fig. 23.33).
Leaves linear, s.t. scabrous and sharp, with parallel veins. Inflorescences spikes or panicles of
spikelets of reduced flowers. Flowering sporadic or staggered in long unpredictable cycles of several
decades, s.t. simultaneous for all individuals, over a large geographical area.
Bamboos are important components of tropical forests of Asia, where they can form thick groves.
Neotropical lowland forests provide a habitat for medium-sized bamboos (e.g. Guadua). Slopes of
Andean mountain ranges are often covered by impenetrable Chusquea thickets. Bamboos provide,
especially in Asia, multi-purpose material, for construction, thatching, etc. Bamboo trunks do not lend
themselves to sawing, they can be split but are most often used for poles and posts. When used for
construction, they are attached with ties, rather than with nails or screws.
! Are Bamboos doing maths? Mast flowering intervals (some as long as 120 years), can be
factorized into small prime numbers which may represent the initial synchronization intervals [8].
23.13 Podocarpaceae: 1 375
23.13 Podocarpaceae: 1
A gymnospermous family encompassing 17 genera and 172 species, centred in the Southern
Hemisphere. Trees often resinous. Multilayered inner bark (Fig. 23.36). Buds distinctly developed,
with several scales. Leaves spiral, opposite with twisted petioles in Retrophyllum (NC, OC). Leaves
flat, narrow and elongate (Podocarpus) or narrow, bent and acute (Dacrydium, South eastern AS and
Oceania). Petioles short, with their bases decurrent along the stem (Fig. 23.35). Male cones
resemble catkins and have numerous scales. Female cones uniovulate, often two seeds per ‘cone’,
e.g. Dacrydium beccarii. The ovules s.t. with their peduncles swelling and becoming fleshy
(Fig. 23.36), a feature for attracting birds (Podocarpus, Prumnopitys). In Dacrydium, the same
function is performed by scales basal to the ovule. Dominant architectural model: RAUH
(Fig. 23.34).
! The majority of Podocarpaceae can be found in montane rainforests, however in Central
America, Podocarpus guatemalensis can grow at elevations as low as 200 m (Fig. 23.35) and in
Malaysia, Podocarpus polystachyus grows on seashore and sandy ridges in mangrove. Uses: timber
and ornamentals.
Fig. 23.34 Podocarpus coriaceus, RAUH’s model, Guadeloupe

Fig. 23.35 Podocarpus sp., strong rhythmic growth, decurrent petiolar bases, Seram, Indonesia. Podocarpus
guatemalensis, sapling, Cockscomb, Belize
Fig. 23.36 Podocarpus tepuiensis, multilayered inner bark (bark pink, wood yellow), Venezuela. Podocarpus
drouynianus, blueish and fleshy peduncles bearing orange coloured ovules, in cultivation (orig. Western Australia),
with the courtesy of Richard Clark
23.14 Polygalaceae: 11-54
24 genera and 700 species. Order Fabales. Plants not aromatic and without latex. Lianas with pre-
hensile twigs (Fig. 23.37d) (Bredemeyera, Securidaca) or twining (Moutabea). Leaf arrangement
spiral (Fig. 23.37h) (Bredemeyera, Diclidanthera) or two-ranked (Fig. 23.37e Securidaca,
Fig. 23.37b Xanthophyllum). Stipules absent, s.t. glands in stipular position (Fig. 23.37d Securidaca).
Leaves simple, entire. Young leaf-folding conduplicate (Fig. 23.38). Venation pinnate. Glands
scattered on underside of lamina (Fig. 23.37f Moutabea, Fig. 23.37c Xanthophyllum). Plants glabrous
(Xanthophyllum) or pubescent (Bredemeyera, Securidaca). Inflorescences racemes or panicles
(Fig. 23.37j), spikes. Flowers with a bilateral symmetry looking like those of the Papilionoideae
(Fig. 23.38). Ovary superior rimmed by a crown of staminodes. Fruits capsules, samaras or drupes.
23.14 Polygalaceae: 11-54 377
Fig. 23.37 Xanthophyllum sp. a sapling, MANGENOT; b stem extremity showing apex abortion (+); c: lamina with
glands; d prehensile twig, cauline glands (gl), (Securidaca sp.); e two-ranked leaf arrangement (Securidaca sp.); f foliar
glands (gl) disposed on veinlets (Moutabea); g imbricate aestivation of young leaves (Bredemeyera, Securidaca);
h spiral leaf arrangement (Bredemeyera); j continuous branching and spiral leaf arrangement in Diclidanthera sp
Fig. 23.38 Bredemeyera sp., young leaf-folding conduplicate, Venezuela. Diclidanthera sp., flowers with weak
bilateral symmetry, Venezuela. Securidaca sp., “papilionoid” flowers, Venezuela
Similar families: Architectural models badly known:

EBENACEAE (Diospyros): leaves with LEEUWENBERG: Polygala spp. (subshrubs and
scattered glands such as Xanthophyllum. herbs, Pantrop).
MANGENOT (Fig. 23.37a): Xanthophyllum (AS).
CHAMPAGNAT (Fig. 23.37h): probable in
Diclidanthera (AM).
Trees (Xanthophyllum, AS, AU), shrubs (Monnina, AM; Polygala, Xanthophyllum), lianas
(Bredemeyera, Securidaca, AM; Moutabea, AF) or herbs (Monnina, AM; Polygala, Pantrop).
Polygalaceae are mainly native to America, Asia and Australia. Uses: some dyes and oil seeds
(Xanthophyllum spp., ‘minyak berok’, Malaysia).
Xanthophyllum, or ‘yellow leaf’ in Greek, have leaves turning yellow soon after a drying process
(for making herbarium parts), because of the accumulation of aluminium ions in tissues. Several other
families, for instance the Melastomataceae, are also aluminium accumulators [9].
23.15 Polygonaceae: 5-14-54
52 genera and 1200 species. Order Caryophyllales. Plants not aromatic and without latex. Lianas s.t.
with phloem poles (Fig. 23.39b). Pigments are anthocyanic (Fig. 23.40) and not betalains which are
common in Caryophyllales. Short shoots develop especially in lianas (Fig. 23.39a). Hollow stems
inhabited by ants in Triplaris. Leaf arrangement spiral (Coccoloba) or two-ranked (Coccoloba,
Triplaris). Stipules annular or cylindrical (Fig. 23.39c) leaving an annular scar on the stem
(Fig. 23.41). Leaves simple, entire (Fig. 23.40), rarely peltate (Fig. 23.42). Young leaf-folding
revolute (Fig. 23.39a). Venation pinnate. Inflorescences usually in spikes (Fig. 23.42) or panicles
(Fig. 23.40). Flowers small, trimerous, with 3 + 3 tepals (Fig. 23.43). Ovary superior. Fruits nuts or
three-sided achenes.
Trees (Ruprechtia, Fig. 23.43-e Triplaris, AM), shrubs (Fig. 23.42 Gymnopodium, AM), lianas
(Fig. 23.40 Antigonon, Coccoloba, AM) or herbs (Fig. 23.43 Oxygonum, Persicaria).
Fig. 23.39 a Afrobrunnichia erecta and Coccoloba sp. a Short shoot, revolute leaf-folding; b–c Coccoloba spp.,
flatened stem of a liana with two phloem poles, ochrea
Fig. 23.40 Antigonon leptopus, liana, cordate leaves, inflorescences paniculate, naturalized Seram. Coccoloba uvifera,
seashore, Belize
23.15 Polygonaceae: 5-14-54 379
Fig. 23.41 Coccoloba sp., sapling (precursor of a liana) with arched branches, one of these bearing an erect flagellum,
Belize. Coccoloba sp., short shoot bearing red-coloured young leaves, Venezuela. Coccoloba sp., annular scar of an
ochrea (main stem), ochrea visible on the lateral stem, Venezuela
Fig. 23.42 Coccoloba sp., peltate leaves, Belize. Coccoloba cf. orinocense, inflorescence in spike, Venezuela.
Gymnopodium floribundum, ROUX’s model, Belize
Polygonaceae are mostly American. Few uses are known in the tropical rain belt, although at
higher latitudes, Sorrel (Rumex patientia), Buckweed (Fagopyrum esculentum) and Rhubarb (Rheum
rhabarbarum) are well known food plants in Europe. Seagrape (Fig. 23.40 Coccoloba uvifera) are
small trees growing on almost all seashores of the Caribbean region. Fallopia baldschuanica is an
ornamental liana native to Central Asia.
! Quoting Burkill [10], p. 2185: «Triplaris surinamensis was introduced into the Botanic Gardens,
Singapore, in 1887. … It is exceedingly interesting that ants live in the stems in Singapore just as they
do in Suriname, though the Singapore ants must be a species formerly quite unfamiliar with the tree».
Fig. 23.43 a Gymnopodium floribundum, flowers trimerous, Belize. b Persicaria limbata, herb, inflorescences in
spikes, Thailand. c Muehlenbeckia platyclada, flat chlorophyllous stems bearing reduced leaves, TROLL’s model.
d Oxygonum sinuatum, annual herb, prostrate ACOSTA’s model, detail: node and leaf (inflorescence in red), Botswana.
e Triplaris surinamensis. flowering tree, Scarrone’s model
ROUX (Fig. 23.42): Coccoloba uvifera,
Gymnopodium floribundum (AM).
Similar families: SCARRONE (Fig. 23.43e): Triplaris
surinamensis (AM).
An unmistakable family when the peculiar shape of
stipules (ochrea) is observed. CHAMPAGNAT: Coccoloba spp. (AM).
TROLL (Fig. 23.43c): Coccolobaspp. (AM),
Muehlenbeckia platyclada, Melanesia.
ACOSTA (Fig. 23.43d): Oxygonum sinuatum
(AF-S).
23.16 Primulaceae: 11–23-29
57 genera and 2645 species. Order Ericales. Plants not aromatic and without latex. Inner bark
displaying s.t. a network of fibres (Fig. 23.47). When branches are shed, they leave a large oval-
shaped scar on the trunk (Fig. 23.46c, Fig. 23.51). Branches often enlarged at their base
(Fig. 23.45). Leaf arrangement spiral, less often two-ranked (Ardisia spp.). Stipules absent. Leaves
simple, entire or toothed, mucronate in Jacquinia (Fig. 23.48). Venation pinnate. Underside of leaves
with minute glandular trichomes (Fig. 23.46e) or scales (Fig. 23.46d). Resiniferous ducts in leaves
(Fig. 23.50 Maesa). Inflorescences racemose (Fig. 23.48), umbellate (Fig. 23.44), terminal
(Deherainia, Fig. 23.49 Jacquinia), but most often lateral (Ardisia, Fig. 23.48 Clavija, Fig. 23.50
Maesa, Myrsine, Oncostemum). Flowers usually with 5 sepals, 5 petals, 5 stamens and often 5
23.16 Primulaceae: 11-23-29 381
Fig. 23.44 Umbella of 5-merous flowers, an unidentified Fig. 23.45 Swollen branch base, an unidentified
specimen, Thailand specimen, Thailand
staminodes (Fig. 23.49). Elaiophores attracting bees. Ovary superior (semi-inferior in Theophrasta),
uniloculate and with central placentation. Fruits capsules (Primula), berries (Fig. 23.51) or drupes
(Fig. 23.47).
Fig. 23.46 a CORNER’s model (Clavija, Theophrasta). b RAUH’s model (Cybianthus). c: Scar left by a fallen twig
(bs), leaf scar (ls). d Serrulate leaves with scaly (sc), or e glandular trichomes (trich)
Fig. 23.47 Inner bark with a network of fibres, an unidentified specimen, Thailand. Spiral, pseudo two-ranked leaf
arrangement, an unidentified specimen, Thailand. Lateral small drupes, an unidentified specimen, Venezuela
Fig. 23.48 Clavija sp., contracted infructescences of berries, one of these peeled, Venezuela. Clavija lancifolia,
CORNER’s model, spikes of orange-coloured flowers, Venezuela. Jacquinia macrocarpa, leaves mucronate, Belize
Fig. 23.49 Jacquinia macrocarpa, inflorescence terminal, Belize. Jacquinia barbasco, corollas with 5 petals and 5 flat
staminodes, Guadeloupe. Jacquinia macrocarpa, section of a berry, Belize
Fig. 23.50 Maesa ramentacea, inflorescences lateral, Thailand. Maesa sp., leaf with resiniferous ducts and glandular
trichomes. Myrsine ferruginea, inflorescences lateral, Southern Brazil
Primulaceae-Myrsinoideae (Ardisia, Cybianthus, Myrsine) are dificult to identify at the species

level, even in the presence of flowers.
Trees (never very tall), shrubs (Myrsine, Maesa), s.t. pachycaulous (Clavija, Oncostemum,
Tapeinosperma, Theophrasta), climbers (Embelia spp.) or herbs (Labisia, Androsace, Primula). The
“old” Myrsinaceae, dating from the pre-APGan classifications, have been merged into Primulaceae
(i.e. primroses Fig. 23.51, androsaces and cyclamens). The Asian genus Labisia represent the
23.16 Primulaceae: 11-23-29 383
Fig. 23.51 Oncostemum sp., berries, Madagascar. Oncostemum umbellatum, shield-shaped scar left by a fallen branch,
Madagascar. Primula vulgaris, Switzerland
Fig. 23.52 a CORNER, e.g. Myrsine grandifolia, New Caledonia; b LEEUWENBERG, e.g. Deherainia smaragdina,
AM-C; c ATTIMS, e.g. Myrsine asymmetrica, New Caledonia; d MASSART, e.g. Ardisia crenata, AS;
e CHAMPAGNAT, e.g. Maesa lanceolata, AF-S, MA
“not-missing” link between arborescent and herbaceous species. Apart from numerous ornamental
uses, fish poisons (Jacquinia spp., ‘barbasco’) and medicinals (Embelia ribes, Labisia, Maesa).
With a distribution extending from the equator to circumboreal mountain ranges, few features are
of general occurrence in Primulaceae. However, the family avoids aridity (except Maesa, placed in
the segregate family Maesaceae!), we do not know of any species growing in African (among Acacia
trees) or in American savannas (with Calliandra and Curatella), or in the steppes of Central Asia.
Primulaceae grow in the understorey of tropical rainforests or on high altitude grassy steppes and
rocks in a humid climate (Alps, Himalaya, etc.).
Similar families: Architectural models (RAUH’s being dominant):

PITTOSPORACEAE (in CORNER (Fig. 23.52a): Clavija lancifolia (AM), Oncostemum
Paleotropics): if glandular trichomes cauliflorum (MA), Tapeinosperma pachycaulum (Solomon Islands).
not properly observed.
LEEUWENBERG (Fig. 23.52b): Deherainia smaragdina (AM-C),
Tapeinosperma psaladense (New Caledonia).
KORIBA: Jacquinia barbasco (AM-C). SCARRONE: Aegiceras
corniculatum, shrub of paleotropical and Pacific mangroves.
RAUH (Fig. 23.46b): Ardisia elliptica (Thailand), Cybianthus spp.
(Guianas), Myrsine africana (Paleo), Myrsine erythroxyloides (MA),
Stylogyne standleyi (AM-C), Tapeinosperma vieillardii (New
Caledonia).
ATTIMS (Fig. 23.52c): Myrsine asymmetrica (New Caledonia).
MASSART (Fig. 23.52d): Ardisia crenata (AS), Ardisia
polycephala (Papua New Guinea), Oncostemum palmiforme (MA).
ROUX: Embelia (Paleotrop).
CHAMPAGNAT (Fig. 23.52e): Maesa lanceolata (MA), this genus
often placed in a family of its own: the Maesaceae [11].
23.17 Proteaceae: 25-44-45-48
75 genera and 1710 species. Order Proteales. Plants not aromatic and without latex. Inner bark with a
network of fibres and external wood furrowed (Figs. 23.54e, 23.57). Stems lenticellate (Fig. 23.54f).
S.t. serial buds at leaf axils (e.g. Panopsis Fig. 23.54-f). Leaf arrangement spiral, rarely subopposite
(Fig. 23.56 Panopsis spp.) or whorled (Fig. 23.55). Stipules absent. Leaves coriaceous, very variable
in shape: simple, pinnatifid (Fig. 23.54c) or compound (Fig. 23.54d Roupala spp.), entire or toothed.
Several species are heterophyllous: leaves of the adult plant have a different shape than those of the
juvenile plant. Venation pinnate, brochidodromous (Fig. 23.55) in broad-leafed plants. Indument
usually absent (plants glabrous). Inflorescences often in
showy spikes (Fig. 23.56). Flowers with tubular corolla
(Fig. 23.53) with 4 petals. Fruits follicles (Roupala), ach-
enes, nuts or drupes (Helicia, Macadamia, Panopsis).
Trees (AM: Roupala, AF: Faurea; AS: Helicia, Australasia:
Grevillea). Shrubs (AU: Banksia, AF: Protea) or small shrubs
(AU: Hakea), geoxylic in S-African Protea.
Typically a Gondwanan group, almost confined to Aus-
tralia and Southern Africa. In savannas or shrubby environ-
ments: Faurea saligna (AF-S), Roupala spp. (AM), but
Helicia spp. (AS) grow also in rainforests. Numerous orna-
mentals (Banksia, Persoonia, Protea, etc.). Australian hazel-
nut: Macadamia integrifolia, M. tetraphylla. Grevillea
robusta is grown to make shfade to coffee plantations.
! At the end of the 1990s, at the time of the first APG
classifications, the botanic world was shocked by results
showing that Platanaceae were much closer to Proteaceae
than conventionally admitted. This result is consistent with
anatomical observations: both families exhibit a furrowed
external wood. Fig. 23.53 Oreocallis grandiflora, Cuenca,
Ecuador
23.17 Proteaceae: 25-44-45-48 385
Fig. 23.54 Leaves of Proteaceae. a Juvenile leaf of Roupala sp.; b simple and toothed, Helicia sp.; c pinnatifid
(Heliciopsis, Stenocarpus); d compound pinnate (adult leaves of Roupala sp.); e typical: bark fibrous and external wood
furrowed; f typical: stem lenticellate, Panopsis sp. has serial buds
Fig. 23.55 Helicia sp., leaves toothed, venation brochidodromous, Seram, Indonesia. Macadamia tetraphylla, leaves
in whorls of four, Flores, Indonesia
Fig. 23.56 Panopsis rubescens, subopposite leaf arrangement, Caura, Venezuela. Roupala montana, inflorescence in
spike, Venezuela
Fig. 23.57 Roupala montana, external wood furrowed and bark fibrous, Belize. Roupala montana, leaf compound
pinnate, Venezuela. Stenocarpus sinuatus, leaves pinnatifid, BG Geneva

Several! when leaves are simple and if LEEUWENBERG: Beauprea pancheri
observation of typical bark has failed (e.g. (NC), Aulax umbellata, Leucadendron
Pentaphyllacaceae, Polygalaceae, ...) eucalyptifolium, Protea cynaroides.
RAUH: Macadamia ternifolia (AU),
Leucospermum reflexum (AF-S),
Embothrium coccineum (AM-S, Andes).
SCARRONE: Faurea galpinii (AF-S),
Grevillea robusta (AU), Leucadendron
argenteum and L. rubrum (AF-S),
Oreocallis grandiflora (Andes of Ecuador).
TROLL’s model: probably absent.
23.18 Putranjivaceae: 8-9-13-A
A small family of 2 genera and 205 species. Order Malpighiales. Plants not aromatic and without
latex. Inner bark not fibrous. Glucosinolates or mustard oils. Trunk with continuous branching
(Fig. 23.59). Trunk or branches lenticellate (Fig. 23.59). Branches with two-ranked leaf arrangement
(Fig. 23.58). Very rarely, hollow stems inhabited by ants (Drypetes longifolia). Stipules present
(Fig. 23.59), s.t. vestigial, never leaving an annular scar on the stem. Leaves simple, entire (e.g.
Drypetes confertifolia (India), D. longifolia, Thailand) or toothed (e.g. D. arguta (AF-S), D. fan-
shawei (Venezuela). s.t. spiny (Fig. 23.58). In a few cases, leaves are toothed in the juvenile tree and
entire in the adult tree. Venation pinnate. Leaves without glands. Flowering lateral (Fig. 23.59).
Flowers reduced, with stamens variable in numbers. Ovary superior, 1-3-(6) loculate with axil
placentation. Fruits drupes (Fig. 23.59).
23.18 Putranjivaceae: 8-9-13-A 387
Fig. 23.58 Drypetes sp., leaves spiny, Cameroon
Fig. 23.59 a Drypetes sp., fruits axillary, Madagascar. b Drypetes laterifolia, bark lenticellate, Belize. c Drypetes sp.,
trunk with continuous branching, foliaceous stipules (st) Cameroon
Similar families:
PHYLLANTHACEAE, the sister family
of Putranjivaceae.
SALICACEAE: also plagiotropic ROUX (Fig. 23.59): Drypetes spp
branches and stipulate, toothed leaves.
Trees (Drypetes is pantropical, but numerous African species, Putranjiva in Asia), no lianas and
no herbs. Drypetes and Putranjiva are evergreen trees of rainforests. In India, Putranjiva roxburghii
is used for treating female infertility and the oil of the seeds can be used for burning.
! Drypetes variabilis is a tree named in Venezuela ‘kerosen’, because of the smell of its bark!
References
1. Callmander, M.W., T.J. Booth, and S. Buerki. 2013. Update on the systematics of Benstonea (Pandanaceae): When
a visionary taxonomist foresees phylogenetic relationships. Phytotaxa 112: 57–60.
2. Stone, B.C. 1978. Revisio Pandanacearum, Part I. Pandanus subgenera Coronata and Acrostigma. Flora Malesiana
precursores. Federation Museums Journal, New Series 23: 1–74.
3. Hallé, F., and R. Keller. 2019. Mais d’où viennent les plantes?, 180 pp. Arles: Actes Sud.
4. Zimmermann, M.H., P.B. Tomlinson, and J. LeClaire. 1974. Vascular construction and development in the stem of
certain Pandanaceae. Botanical Journal of the Linnean Society 68: 21–41.
5. D’Orbigny, C. 1847. Dictionnaire universel d’histoire naturelle, Tome neuvième, 776 pp. Renard, Martinet et Cie, Paris.
6. 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.
7. Cayzer, L.W., M.D. Crisp, and R.H. Telford. 2000. Auranticarpa, a new genus of Pittosporaceae from northern
Australia. Australian Systematic Botany 13: 903–917.
8. Veller, V.C., M.A. Nowak, and C.C. Davis. 2015. Extended flowering intervals of bamboos evolved by discrete
multiplication. Ecology Letters 18: 653–659.
9. Jansen, S., T. Wanatabe, and E. Smets. 2002. Aluminium accumulation in leaves of 127 species in
Melastomataceae, with comments on the order Myrtales. Annals of Botany 90: 53–64.
10. Burkill, I.H. 1935. A dictionary of the economic products of the Malay peninsula, vols. I and II, 2402 pp. Oxford:
University Press.
11. Anderberg, A.A., C. Rydin, and M. Källersjö. 2002. Phylogenetic relationships in the order Ericales s.l.: Analyses
of molecular data from five genes from the plastid and mitochondrial genome. American Journal of Botany 89:
677–687.
Rhabdodendraceae to Rutaceae
24
24.1 Rhabdodendraceae: 23
A monogeneric family with only 3 species, endemic to the Guianas and Amazonia. Order
Caryophyllales. Monocaulous plants with leaves rather large and cuneate at their base (Fig. 24.1).
Young leaf-folding revolute (Fig. 24.1). Leaves with peltate hairs and glandular dots. Inflorescences
racemose (Fig. 24.1). Fruits drupes.
The family is basal in the order Caryophyllales [1].

RUTACEAE: In Amazonia and in Guianas Erythrochiton spp. are also CORNER (Fig. 24.1):
pachycaul, non-ramified treelets. Rhabdodendron spp.
Fig. 24.1 Rhabodendron sp., CORNER’s model, Imataca, Venezuela. Rhabdodendron amazonicum, young leaf-
folding revolute (rev), St Elie, French Guiana. Rhabodendron sp., inflorescence lateral and racemose, Imataca,
Venezuela
https://doi.org/10.1007/978-3-031-05942-1_24
390 24 Rhabdodendraceae to Rutaceae
24.2 Rhamnaceae: 6-8-9-17-18-31-32-33-54-55-56
63 genera and 1050 species. Order Rosales. Plants without latex, not aromatic. Inner bark without
network of fibres, very bitter in Maesopsis eminii and smelling of methyl salicylate in Lasiodiscus
fasciculifloris. Lianas with prehensile twigs (Ventilago) or tendrils (Fig. 24.4 Gouania). Spiny stems
(Fig. 24.6 Ziziphus). Branches usually with two-ranked leaf arrangement (Fig. 24.6) or leaves op-
posite (Fig. 24.5), Maesopsis, Scutia. Stipules present. Leaves simple, entire or toothed. Venation
pinnate, often scalariform (Fig. 24.6) (Maesopsis, Ziziphus). Leaves usually without glands (glands
present in Colubrina spp. Fig. 24.3). Flowers small wih 5 petals and 5 petal-opposed stamens. Ovary
superior rimmed by a nectariferous disk. Fruits drupes (Fig. 24.6), s.t. dehiscent, or samaras
(Fig. 24.5). Trees (Alphitonia, Berchemia, Maesopsis, Ziziphus). Shrubs (Colletia, Colubrina),
lianas (Gouania, Ventilago), almost all arborescent or shrubby, only Crumenaria is herbaceous [2].
Fig. 24.2 Ziziphus mucronata growing on a termite mound, Botswana
Fig. 24.3 Colubrina glandulosa, treelet, MASSART’s model, Venezuela. Alphitonia sp. ROUX’s model
24.2 Rhamnaceae: 6-8-9-17-18-31-32-33-54-55-56 391
Fig. 24.4 Berchemia discolor, treelet, MASSART’s model, Botswana. Gouania polygama, infructescence of samaras,
Belize. Gouania sp., liana bearing tendrils, Belize
Fig. 24.5 Maesopsis eminii, series of unramified nodes and ramified nodes, these exhibiting anisoclady. Scutia
myrtina, a climber with spines, MASSART’s model. Ventilago sp., samaras of a unidentified species, Thailand
Fig. 24.6 Ziziphus sp., a zigzagging branch bearing stipules modified into spines. Ziziphus mauritiana, fruit and
discoloured leaves, Belize. Typical: camptodromous and scalariform venation
The family is cosmopolitan in range, especially in warm temperate and tropical regions. Timber
(Maesopsis eminii, ‘muzizi’), fruit trees (Berchemia, Hovenia, Ziziphus), dyes (Alphitonia, Rhamnus),
numerous medicinals.
RAUH: Karwinskia humboldtiana (Mexico), Krugiodendron
ferreum (AM-C), Reynosia latifolia (Bahamas).
Similar families: MASSART (Fig. 24.5): Berchemia discolor (AF-S),
PHYLLANTHACEAE, but with periderm Colubrina glandulosa (Guianas), Scutia myrtina (MA).
with thin longitudinal slits.
SALICACEAE: but leaves with a hyaline ROUX (Fig. 24.3): Alphitonia spp. (AS, AU, OC), Colubrina
margin. arborescens (WI), Maesopsis eminii (AF); Ventilago cristata
(AS), Ziziphus papuana (liana).
TROLL: Ziziphus mauritiana, an Indomalesian tree, often
introduced, very common in the tropics.
! Nesiota elliptica, an endemic species of a monotypic genus of St Helena Island (Southern

Atlantic), has disappeared in the wild since 1994. In 2004, the single existing cultivated individual
died, so the species is now extinct [3].
24.3 Rhizophoraceae: 32-33-34
15 genera and 125 species. Order Malpighiales. Plants not aromatic and without latex. Trunks of
forest trees exhibiting ‘eye-marks’ (i.e. scars of fallen branches Fig. 24.8). Mangrove trees with stilt
roots (Fig. 24.7 Rhizophora). Trunk or branches lenticellate (Fig. 24.8). Inner bark without network
of fibres. External wood often furrowed (Fig. 24.8 Carallia, Sterigmapetalum). Leaf arrangement
Fig. 24.7 Rhizophora mucronata, mangrove, Bali

24.3 Rhizophoraceae: 32-33-34 393
opposite (3–4-whorled in Anopyxis, AF). Stipules interpetiolar (Fig. 24.9) leaving an annular scar,
elongate and enclosing buds in Rhizophora (Fig. 24.9). Leaves simple, entire or with minute teeth
(Blepharistemma). Venation pinnate, brochidodromous. Glands absent, but s.t. underside of leaves
with minute trichomes (Carallia, Fig. 24.10-c Rhizophora). Flowers with 4–5 petals (Fig. 24.9)
(petals fringed in Cassipourea), stamens variable in numbers. Fruits berries or capsules.
Fig. 24.8 Blepharistemma sp., trunk lenticellate, with eye-marks, Kerala, India. Cassipourea sp., eye-marks, lenticels,
Venezuela. Carallia sp., external wood furrowed, Seram, Indonesia
Fig. 24.9 Cassipourea guianensis, stipules interpetiolar, Guadeloupe. Rhizophora mangle, Belize. Stipules hood-like.
Fleshy persistent flowers, open capsule
Fig. 24.10 a Apposition growth in AUBRÉVILLE’s model (e.g. Ceriops, Rhizophora). b Rhizophora mangle, stilt-
roots, Belize. c Rhizophora mangle, underside of leaves covered by trichomes, Belize. d Cassipourea guianensis,
young tree conforming to ATTIMS’s model, Guadeloupe (author’s field book)
Trees: Cassipourea, Sterigmapetalum (AM), Gynotroches (AS, OC), Carallia (MA, AS, AU).
Shrubs of mangroves: Bruguiera, Ceriops (Paleo), Rhizophora (Pantrop). No lianas and no herbs.
Several Rhizophoraceae are very common in mangroves (Bruguiera, Ceriops, Rhizophora),
however some members of the family grow in forests (Carallia, Blepharistemma). Not a sustainable
source of charcoal: Bruguiera and Rhizophora in mangroves! Tanins.
! Mangrove species (Rhizophora, Ceriops, Bruguiera, Kandelia) form a monophyletic group
which is also the most highly evolved in the classification published by [4]. This group is clearly
separate from the inland representatives of the family.
24.4 Rosaceae: 7-11-15-46 395
AUBRÉVILLE (Fig. 24.10a):
Bruguiera (AF, AS, OC), Ceriops (AS,
Similar families: OC).
ATTIMS: Anopyxis (AF), Crossostylis
RUBIACEAE have also interpetiolar
(OC), Rhizophora (Pantrop).
stipules, however they are rarely
lenticellate and have smooth external MASSART: Carallia, MA, AS, AU.
wood. ROUX: Cassipourea (AM, AF), Pellacalyx
(AS).
ATTIMS (Fig. 24.10d ): Cassipourea
guianensis (Guadeloupe).
24.4 Rosaceae: 7-11-15-46
83 genera and more than 3500 species, but poorly represented in tropics. Wounded bark often
producing gums. Trunk and branches typically lenticellate, these transversely elongate (Fig. 24.11).
Leaf arrangement usually spiral (Prunus spp., etc.), but s.t. two-ranked (Prunus spp.). Leaves
stipulate, usually simple and often toothed (compound in Hagenia, Fig. 24.14 Rubus). Flowers with
5 petals and numerous stamens. Fruits drupes (Fig. 24.14 Prunus), groups of small drupes
(Fig. 24.14 Rubus) or achenes (Geum, Hagenia, Polylepis), these embedded in a fleshy receptacle to
form an “apple” or a “pear” (Malus, Pyrus).
Fig. 24.12 Hagenia abyssinica, RAUH’s model
Fig. 24.11 Prunus cerasoides, lenticels transversely

elongate, Thailand
Trees, several Prunus species grow in South East Asia (P. arborea, P. cerasoides), East Africa
(P. africana) and Mexico (P. capuli). Some trees and shrubs in the Andean mountain range
(Kageneckia, Hesperomeles, Polylepis), (Fig. 24.13), Hagenia abyssinica in Central Africa highlands
(Fig. 24.12). Eryobotrya japonica (Nispero) is grown in the subtropics, or even naturalized
(e.g. Madagascar). Shrubs, brambles (Rubus) and herbs. Uses are many: fruit trees, phytosterols
(Prunus africana). Black dye extracted from leaves (Kageneckia lanceolata).
! Sanguisorbeae (Sanguisorba, Acaena, Hagenia, Polylepis) may represent an example of evo-
lution from herbs (like Sanguisorba) to trees (like Hagenia or Polylepis) [5].
Polylepis australis (AM-S).
FAGERLIND: Eryobotrya
japonica (AS, cult.).
RAUH (Fig. 24.12): Hagenia
abyssinica (AF-E), Prunus spp.
CHAMPAGNAT: Rubus spp.
Fig. 24.13 Polylepis australis, LEEUWENBERG’s

model (with the courtesy of Francis Hallé)
Fig. 24.14 Rubus fraxinifolius, flowers and fruits, a native bramble in Flores, Indonesia. Prunus mahaleb, leaves
toothed, drupes, Switzerland
24.5 Rubiaceae: 32-56-E 397
24.5 Rubiaceae: 32-56-E
576 genera and 12,000 species. Order Gentianales. Plants not aromatic and without latex. Peeling
bark in Calycophyllum (Fig. 24.17b). Rarely buttressed trees (Fig. 24.17c). Trunk or branches usually
not lenticellate. Inner bark without network of fibres. Some species with short shoots modified into
thorns (Fig. 24.26b) or hooks (lianas, Fig. 24.15f). Leaf arrangement opposite (Fig. 24.19,
Fig. 24.15), infrequently whorled (e.g. Hamelia spp. Fig. 24.21). Leaf arrangement can also be
opposite but with the successive pairs spirally arranged (bigugate phyllotaxy Fig. 24.26d). Stipules
present and almost always interpetiolar (Fig. 24.16), leaving an annular scar, or intrapetiolar
(Fig. 24.16k). Leaves simple, entire, very rarely lobate. Venation pinnate, often camptodromous
(Fig. 24.18, Fig. 24.25). Leaves without glands but s.t. with domatia on the underside of laminas
(Coprosma). Inflorescences cymose (Fig. 24.20), rarely 1-flowered, terminal or lateral. Showy
bracts in inflorescences are relatively frequent (Fig. 24.24, Fig. 24.25). Flowers with 4–5 petals and
Fig. 24.15 Axis morphology. a Modular construction, branches erect (e.g. Psychotria). b Modular construction,
branch plagiotropic (e.g. Morinda citrifolia). c Branch plagiotropic by apposition and terminal flowering (e.g. Genipa).
d Branch plagiotropic and monopodial. e Branch decumbent. f Liana bearing short shoots modified into hooks (e.g.
Mussaenda, Uncaria). g Monocaulous pachycaul (e.g. Duroia)
Fig. 24.16 Various kinds of stipules. a Curvilinear, Mussaenda; b bilobate, Isertia, Psychotria; c notched, Palicourea;
d forming a short collar, Pagamea; e triangular, Coffea, Gaertnera, Randia, Rondeletia, Uncaria; f forming a short
collar with a linear appendage, Argocoffeopsis; g fringed, Borreria; h spatulate, Cinchona; j acuminate, Catunaregam,
Faramea; k intrapetiolar, Capirona
Fig. 24.17 a Atractogyne sp., liana, Cameroon. b Calycophyllum spruceanum sp., peeling bark, BG FIBV.
c Chimarrhis turbinata, buttressed trunk, Fr. Guiana
Fig. 24.18 Chioccoca sp., white berry, Belize. Cinchona pubescens, camptodromous venation, BG Geneva
greenhouses. Coccocypselum guianense, prostrate herb bearing blue fruits, French Guiana
24.5 Rubiaceae: 32-56-E 399
Fig. 24.19 Coprosma lucida, domatia, cult. BG Menton, France. Coussarea cf. racemosa, internodes angular,
camptodromous venation, Venezuela. Diocrescis erythroclada, small deciduous tree, Thailand
Fig. 24.20 Ernodea littoralis, decumbent shrub bearing short shoots, Guadeloupe. Faramea crassifolia, terminal
cymose infructescence, Venezuela. Gaertnera sp., terminal cyme of fruits, Madagascar
Fig. 24.21 Guettarda speciosa, stipules oval-shaped, Bali, Indonesia. Hamelia patens, leaves in whorls of three,
flowers tubular, Belize. Ixora sp., flowers tubular with 4 petals, Thailand
Fig. 24.22 Ladenbergia sp., infructescences of capsules, Venezuela. Lasianthus sp., blue fruits, Flores, Indonesia.
Morinda citrifolia, cymose inflorescence, Bali, Indonesia
as many stamens (Fig. 24.21). Fruits berries (Fig. 24.26e), frequently blue (Fig. 24.18) or capsules
(Cinchona, Ladenbergia Fig. 24.22).
Similar families:
RHIZOPHORACEAE also have
interpetiolar stipules, but in contrast to
almost all Rubiaceae, their twigs are
lenticellate.
Trees, rarely tall (Chimarrhis, Calycophyllum), often medium-sized (Cinchona, Coffea, Gardenia,
Mitragyna, Wendlandia). Very numerous shrubs (Coffea, Hamelia, Ixora, Psychotria, Palicourea,
Schumanniophyton, Tarenna). Lianas (Atractogyne, Fig. 24.17a, Canthium, Manettia, Uncaria),
epiphytes (Hydnophytum, Myrmecodia), these s.t. with thick stems enclosing cavities inhabited by
ants, or herbs (Argostemma, Rubia, Spermacoce). Herbs in the understorey often have bold striped
patterns on their leaves (Fig. 24.26c) (a physiological adaptation for maximizing light capture).
Rubiaceae are very frequent in forest understoreys, savannas and along trails. Considering that
there are 11,000 species, with the exception of coffee (Coffea arabica, C. liberica) and quinine
(Cinchona pubescens), there are very few products of economic importance. Numerous species
became widely cultivated as ornamental plants (Gardenia, Ixora, Mussaenda, Pavetta). Morinda
citrifolia (Fig. 24.23 cheese fruit, ‘bois douleur’, ‘noni’, ‘luoling’ and many other names!) is grown
everywhere in the tropics and is used in traditional medicine.
! Most tropical representatives of the Rubiaceae reiterate sparsely and therefore lend themselves as
a good material for architectural analysis.
Fig. 24.23 Morinda citrifolia, garden tree, Bali. Mussaenda kerrii, white bracts, Thailand. Nauclea subdita, globular
lateral inflorescences Kebun Raya, Java
24.5 Rubiaceae: 32-56-E 401
Fig. 24.24 Palicourea rigida, shrub adapted to periodic fires, Gran Sabana, Venezuela. Posoqueria latifolia, twig not
lenticellate, Belize. Psychotria coccinea, blue fruits and red bracts, French Guiana
Fig. 24.25 Psychotria poeppigiana, yellow flowers and orange-coloured bracts, Belize. Psychotria glabrata, pale blue
fruits, Guadeloupe. Psychotria sp., cymose infructescence, Venezuela
Fig. 24.26 a Randia cf. armata, short shoots, Venezuela. b Randia sp., short shoots modified into spines, Southern
Brazil. c Unidentified specimen, small plant in the understorey, “leaf veins” violaceous, Venezuela. d Unidentified
specimen, opposite-bijuguate leaf arrangement, Campo forest, Cameroon. e Sabicea sp., pubescent plant with lateral
inflorescences and berries, Venezuela
CORNER (Fig. 24.27-b): Bertiera sp. (AF-W), Duroia sp. (AM-S), Coffea macrocarpa (Mauritius).
TOMLINSON: Spermacoce octodon (herb, AF-W).
LEEUWENBERG (Fig. 24.27-f): Margaritopsis sciadephora (shrubs, AF-E), Psychotria poeppigiana (shrub,
Guianas), Strumpfia maritima (shrub, Guadeloupe).
KORIBA (Fig. 24.27-c): Psychotria deflexa, (AM-C), P. urbaniana (shrubs, Guadeloupe).
FAGERLIND (Fig. 24.27-d): Duroia aquatica (tree, French Guiana), Genipa americana (small tree, AM), Ixora
montana (shrub, NC, Randia fitzalanii, Rothmannia longiflora (shrubs, AF).
PETIT: Atractogyne bracteata (liana, AF), Chimarrhis cymosa (small tree, West Indies), Morinda citrifolia (small
tree, Pantrop), Rothmannia hispida (tree, AF), Schumanniophyton magnificum (shrub AF-C).
SCARRONE (Fig. 24.27-e): Alseis yucatanensis (tree, Mexico), Gardenia imperialis (tree, AF), Isertia coccinea,
Palicourea guianensis (tree and shrub, AM).
RAUH: Bikkia tubiflora (shrub, NC), Mantalania sambiranensis (shrub, MA).
AUBRÉVILLE: Guettarda speciosa (small tree, Pantrop), Macrocnemum glabrescens (small tree, AM-C).
MASSART: Calycophyllum sp. (tree, AM), Craterispermum caudatum (tree, AF-W), Chiococca alba (liana,
Guianas), Randia ruiziana (tree, Guianas).
ROUX (Fig. 24.27-h): Bertiera sp. (shrub, Guianas), Coffea arabica (shrub, Ethiopia), Lasianthus sp. (shrub,
Indonesia), Neolamarckia cadamba (pionner tree, Indomalesia).
CHAMPAGNAT (Fig. 24.27-g): Catunaregam spinosa (shrub, Indonesia), Nauclea latifolia (shrub, AF).
Fig. 24.27 a Hydnophytum sp., CORNER, shoot modified into a bulb with cavities hosting ants, Solomon Islands.
b CORNER, Pentagonia pinnatifida, pinnate compound leaves, Colombia. c KORIBA, Psychotria sp., Congo.
d PETIT, Morinda citrifolia, detail: modular plagiotropic branch, Marquesas Islands. e SCARRONE, Gardenia
imperialis, equatorial Africa. f LEEUWENBERG, Psychotria poeppigiana, Guianas. g CHAMPAGNAT, Catunar-
egam spinosa, spiny shrub, detail: axillary fruit on a short shoot, Bali. h ROUX, Chapelieria madagascariensis,
Madagascar
24.6 Rutaceae: 21-23-24-40-46-48-49-51-52-57-C 403
24.6 Rutaceae: 21-23-24-40-46-48-49-51-52-57-C
Fig. 24.28 Citrus x aurantium ‘Tarocco’, sterile fruit without seeds
153 genera and 1975 species. Order Sapindales. Plants aromatic and without latex. Bark often bitter
(Angostura, AM). Periderm or young bark often lenticellate (Fig. 24.30 Casimiroa). Inner bark s.t.
with network of fibres (Fig. 24.31, (Clausena), Fig. 24.34 (Harrisonia)). Stems spiny (Fig. 24.33
Zanthoxylum spp.). Leaf arrangement often spiral (Citrus, Clausena, Esenbeckia, Vepris, Zan-
thoxylum) or opposite (Fig. 24.29b Euodia, Flindersia). Stipules absent or spines in stipular position
(Fig. 24.33 Harrisonia). Leaves compound pinnate (Murraya, Fig. 24.36 Zanthoxylum), trifoliolate
(Fig. 24.30 Angostura, Euodia, Monniera), palmate (Casimiroa, Fig. 24.29d Vepris) or even simple
(Esenbeckia, Fig. 24.29e Tetractomia). Leaves or leaflets entire (Atalantia, Murraya) or crenulate
(Citrus, Zanthoxylum). Pinnate leaves often with alternate leaflets (Fig. 24.31 Clausena, Fig. 24.35
Murraya, Zanthoxylum). Leaves s.t. spiny (Zanthoxylum). Petioles or rachis s.t. winged (Angostura,
Fig. 24.30 Citrus, Fig. 24.33 Harrisonia). Leaves are ‘unifoliolate’ when an articulation exists
between the lamina and the petiole (petiole can be also distally pulvinate), (Fig. 24.32 Esenbeckia).
Leaves with numerous translucent dots visible against light. Inflorescences cymose (Fig. 24.35) or
racemose. Flowers with radial symmetry and free petals surrounding a nectariferous disk. Ovary
superior (4-5-n loculate) with axile placentation. Fruits berries (Fig. 24.33), coriaceous in Citrus
Fig. 24.31), (often with an embossed periderm, like orange peel Fig. 24.33), schizocarps (Fig. 24.32
Esenbeckia, Zanthoxylum) or drupes (Skimmia, Spathelia).
Fig. 24.29 RUTACEAE. Leaf forms unusual for Sapindales. a Simple leaves, their petiole pulvinate (p), e.g.
Esenbeckia; b opposite, trifoliolate leaves, e.g. Evodia; c alternate, trifoliolate leaves, e.g. Monniera; d palmate leaves,
e.g. Vepris; d opposite trifoliolate leaves, e.g. Triphasia; e opposite simple leaves, e.g. Tetractomia

Possible confusion with HOLTTUM: Spathelia ulei (small
MELIACEAE, another family of hapaxanthic trees of the Guianas).
the Sapindales. CORNER: Erythrochiton
brasiliensis (pachycaul, AM-S).
LEEUWENBERG: Agathosma
collina (subshrubs, AF-S).
SCARRONE: Calodendrum
capense (tree, AF-S), Flindersia
spp. (trees, AS, AU).
RAUH: Coleonema album (AF-
S), Choisya ternata (shrub, Mex.),
Agathosma ovata (AF-S).
CHAMPAGNAT: Harrisonia
perforata (shrub, AS), Casimiroa
edulis (tree, Mexico).
Fig. 24.30 Angostura trifoliata, leaves trifoliolate, Venezuela. Casimiroa edulis, stem lenticellate, BG Geneva. Citrus
maxima, winged petiole, Thailand
Fig. 24.31 Citrus medica, the genuine citron! BG LC, France. Clausena sp., compound leaf with alternate leaflets,
Thailand. Clausena wallichii, fibrous bark, Thailand
24.6 Rutaceae: 21-23-24-40-46-48-49-51-52-57-C 405
Fig. 24.32 Esenbeckia pilocarpoides, petioles distally pulvinate, Venezuela. Esenbeckia grandiflora, Southern Brazil,
schizocarp sectioned and spiny schizocarp
Fig. 24.33 Glycosmis sp., berries with pericarp like orange peel, Bali. Harrisonia perforata, Thailand. Leaves with
winged rachis, spines in stipular position, and spiny trunk
Fig. 24.34 Harrisonia perforata, fibrous bark, Thailand. Luvunga scandens, liana bearing short shoots modified into
spines, Thailand. Micromelum sp., cymose inflorescence, FRIM, Malaysia
Fig. 24.35 Murraya paniculata, Thailand. Leaves compound, leaflets entire and alternate. Unripe green berry.
Zanthoxylum culantrillo, inflorescence terminal, Venezuela
Fig. 24.36 Zanthoxylum rhetsa, Bali, Indonesia. Trunk bearing large corky spines, Large pinnate leaves. Zanthoxylum
sp., corky spine exhibiting rhythmic growth! Venezuela
Trees (Euodia, Flindersia, Zanthoxylum), shrubs (Agathosma, Atalantia, Choisya), lianas

(Luvunga, Fig. 24.34), herbs (Dictamus, Monniera).
Rutaceae are pantropical, with the bulk of their species in Australasia, represent the most diversified
family of the order Sapindales. Timber: Flindersia spp. (AU-E). Numerous fruit trees: Aegle marmelos
(Bengal quince), Casimiroa edulis (Mexican apple), Citrus maxima (‘pomelo’), Citrus medica (the
genuine citron, but the common lemon fruit is of hybrid origin), C. x aurantium (Orange tree), and many
other Citrus species and their hybrids. Citrus fortunella (‘kumquat’). Essential oils (Aegle, Citrus spp.),
in particular lemon oil (Citrus x limon). Medicinal (‘jaborandi’, Pilocarpus microphyllus).
! Fibrous bark is not infrequent in Rutaceae. A feature justifying the placement of the Sapindales in
the clade Malvids (according to APGs classifications).
! Apomictic orange trees (Citrus x aurantium) develop few or no seeds (Fig. 24.28) and are most
commonly grown by grafting.
! Citrus species exhibit complex architecture with a mixture of orthotropic and plagiotropic lateral
axis, whose growth direction depends on the place of insertion on the principal axis [6].
References 407
References
1. Cuénoud, P., V. Savolainen, L.W. Chatrou, M. Powell, R.J. Grayer, and M.W. Chase. 2002. Molecular
phylogenetics of Caryophyllales based on nuclear 18S RDNA and plastic RBCL, ATPB and MATK DNA
sequences. American Journal of Botany 89: 132–144.
2. Renner, S.S. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic Journal of
Botany 13: 519–540.
3. Mabberley, D.J. 2008. Mabberley’s Plant-Book, 3rd ed, 1022 pp. Cambridge: Cambridge University Press.
4. Schwarzbach, A.E., and R.E. Ricklefs. 2000. Systematics affinities of Rhizophoraceae and Anisophylleaceae, and
intergeneric relationships within Rhizophoraceae, based on chloroplast DNA, nuclear ribosomal DNA, and
morphology. American Journal of Botany 87: 547–564.
5. Potter, D., T. Eriksson, R.C. Evans, and T.A. Dickinson. 2007. Phylogeny and classification of Rosaceae. Plant
Systematics and Evolution 266: 5–43.
6. Demni-Marzouk, S. 1986. Morphogenèse de l’appareil caulinaire du Bigaradier (Citrus aurantium L.). Naturalia
Monspeliensia: Numéro hors série: 351–367
Sabiaceae to Symplocaceae
25
25.1 Sabiaceae: 25-30-48-54
3 genera and 50 species. Order Proteales. Plants not aromatic and without latex. Inner bark with a
network of fibres or a chlorophyllous layer (Fig. 25.2). Spiral leaf arrangement (Meliosma, Sabia).
Stipules absent. Leaves simple (Fig. 25.2 Meliosma AS, Sabia) or pinnately compound (Fig. 25.3
Meliosma AS). Leaves or leaflets entire. Petioles geniculate (Fig. 25.2) at base in Sabia or enlarged
and becoming suberized at base in Meliosma. Venation brochidodromous, embosse on the upper
side. Inflorescences paniculate or cymose (Fig. 25.1). Flowers small with 4–5 petals. Stamens not
numerous. Ovary superior, 2–3 loculate. Fruits drupaceous, unilocular or dicoccous, fleshy or not,
indehiscent. Seeds with coiled up embryo in Ophiocaryon. Trees (Meliosma), shrubs, twining lianas
(Sabia), no herbs.
Sabiaceae grow exclusively in forests (AM: Ophiocaryon, AS: Sabia, AM-AS: Meliosma). They
form a basal group in Eudicots and their vaguely pentamerous flowers probably evolved from a
trimeric state [1].
Similar families:
LECYTHIDACEAE: simple-leaved
Meliosma resemble Barringtonia.
MELIACEAE or SAPINDACEAE: for
compound-leaved Meliosma.
ICACINACEAE: when observing Sabia.
Architectural models poorly known:

KORIBA: Meliosma simplicifolia (AS).
Fig. 25.1 Meliosma simplicifolia, terminal inflorescence, Thailand
https://doi.org/10.1007/978-3-031-05942-1_25
410 25 Sabiaceae to Symplocaceae
Fig. 25.2 Meliosma simplicifolia, Thailand. Inner bark fibrous (f), spiral leaf arrangement, venation brochidodromous
and petiolar bases enlarged. Sabia pauciflora, geniculate petioles and embossed venation (with the courtesy of P.B.
Pelser & J.F. Barcelona, available from Nickrent & al., PhytoImages, http://www.phytoimages.siu.edu)
Fig. 25.3 Meliosma sp., Sumatra. Not rhythmic growth of a sapling, leaves compound pinnate, enlarged petiolar base
and leaflets with brochidodromous venation
! Fossilized remains of Sabiaceae have been discovered in Cretaceous layers in Europe, which
would corroborate the Laurasian origin of the group [2]. Sabiaceae are a “wandering” family, as they
have been placed in Ranunculales by Cronquist [3], then in Sabiales (APG III) and, currently, in the
Proteales (APG IV). In fact, considering only vegetative traits, Sabia (Sabioideae) and Meliosma
(Meliosmoideae) have very few features in common (except somewhat geniculate petiolar bases). In
the end, as suggested by Willis [4], these genera could well fit in two separate families.
25.2 Salicaceae (incl. Lacistemataceae): 6-7-8-9-13-17-18-22-30
54 genera and 1250 species. Order Malpighiales. Plants not aromatic and without latex. Plants s.t.
thorny (Flacourtia, Xylosma), the thorns being modified short shoots (Fig. 25.9). Trunk or twigs
lenticellate (Scolopia, Xylosma). Inner bark s.t. with orange-coloured sclerenchymatous inclusions
(Figs. 25.7 and 25.10). Twigs brittle. Branches with leaf arrangement spiral (Fig. 25.7, Euceraea,
Flacourtia, Salix), but usually two-ranked (Fig. 25.5 Banara, Casearia, Homalium, Laetia,
Xylosma). Stipules present, large in Prockia (Fig. 25.9), s.t. very small. Leaves simple, entire or
25.2 Salicaceae (incl. Lacistemataceae): 6-7-8-9-13-17-18-22-30 411
Fig. 25.4 Casearia sp., fleshy capsules and arillate seeds, Thailand
toothed. Young leaf-folding involute Fig. 25.5a). Leaves have s.t. translucent dots or lines (Casearia,
Samyda) and their lamina has a hyaline margin (Fig. 25.10). Leaves s.t. with glands at the bases
(Fig. 25.5 Banara guianensis, Fig. 25.8 Laetia). Venation pinnate, rarely leaves tripliveined (Has-
seltia) or venation palmate (Fig. 25.9 Prockia). Main vein raised on upper side of the lamina.
Inflorescences lateral (Fig. 25.4 Casearia, Fig. 25.7 Euceraea) or terminal, in lateral catkins in Salix
and Populus. Flowers usually small with parts variable in numbers. Ovary superior, 2-3-6 carpellate
with a parietal placentation (Fig. 25.6). Fruits berries (Flacourtia) or (fleshy) capsules (Fig. 25.4
Casearia, Samyda).
Similar families:
PHYLLANTHACEAE, but their young leaves do Architectural models:
not have an involute folding, almost always no ROUX: Casearia (Pantrop), Laetia (AM), Ryania (AM).
glands, no hyaline leaf margin and no translucent
RAUH: Euceraea (AM-S) Populus, Salix (mainly
dots.
Northern Hemisphere).
VIOLACEAE also look like Salicaceae.
TROLL: Banara (AM), Dovyalis caffra (AF-SE),
APHLOIACEAE: Aphloia theiformis looks like Homalium (Pantrop).
Casearia spp.
Trees (Flacourtia rukam up to 40 m in New Guinea, Homalium, Laetia, Salix), shrubs (Casearia,
Flacourtia), almost no lianas and no herbs. The Flacourtiaceae is a defunct family whose former
members have been scattered to various families, mostly to the Achariaceae and Salicaceae. Sali-
caceae are divided in about ten tribes, Saliceae is dioecious, Flacourtieae is usually dioecious, the
other tribes being hermaphrodite or dioecious. Casearia and Homalium are the only pantropical
genera. A few Salix species extend to the tropics (S. humboldtiana, Andes, S. tetrasperma, Thailand).
Timber, medicinals, small fruit trees: Dovyalis caffra.
! Wind pollination of flowers is an advanced trait in Salicaceae, the willows probably originated
from tropical ancestors. Wind pollination is rare in the lowland tropics, perhaps owing to the fact that
trees are usually not leafless and wind-borne pollen would not be very efficient.
Fig. 25.5 Leaves. a: Glandular and young leaf-folding involute (Banara); b: peculiar venation in Ludia (AF-E);
c: typical leaf. Banara guianensis, leaves two-ranked, with glands at their bases, French Guiana
Fig. 25.6 Casearia sp., glomerulate lateral inflorescences, Venezuela. Casearia sp., lateral small groups of fruits,
Venezuela. Casearia cf. spinescens, fruit showing parietal placentation, Venezuela
Fig. 25.7 Euceraea nitida, inflorescences in lateral panicles, Venezuela. Flacourtia indica, short shoots modified into
spines, Thailand. Flacourtia inermis, sclerenchymatous inclusions in bark, Kebun Raya, Java
Lacistemataceae is a small American family of 2 genera and 44 species, very close to SALI-
CACEAE sharing with it the ROUX’s model. Deciduous stipules leaving an almost annular scar
help to distinguish them from SALICACEAE. Other similar features: leaves with pinnate venation,
hyaline margin and fruit capsular (Fig. 25.11) (cf. Casearia). In Belize, Lacistema aggregatum is
named ironically ‘hairy nose tree’, see the fascicles of small spikes (Fig. 25.11) in leaf axils!
25.2 Salicaceae (incl. Lacistemataceae): 6-7-8-9-13-17-18-22-30 413
Fig. 25.8 Homalium tomentosum, leaves shortly petiolate, Thailand. Homalium racemosum, two-ranked leaf
arrangement, Venezuela. Laetia procera, base of laminas with glands, French Guiana
Fig. 25.9 Prockia crucis, leaves stipulate, venation palmate, Belize. Salix tetrasperma, fissured bark, spiral leaf
arrangement, Chiang Mai, Thailand. Xylosma sp., short shoots modified into spines, Belize
Fig. 25.10 Xylosma sp., plagiotropic branch with 2-ranked leaves, Venezuela. Casearia laetioides (syn. Zuelania
guidonia), Belize, trunk lenticellate, bark with sclerenchymatous inclusions, lamina with hyaline margin and translucent
dots or lines
Fig. 25.11 Lacistema aggregatum, Cockscomb, Belize. Inflorescences in fasciculate spikes and fleshy dehiscent fruits
25.3 Santalaceae (S) + Loranthaceae (L): 12-30-37-44
S: 48 genera, 1025 species. L: 84 genera, 950 species. Order Santalales. Plants not aromatic (except
for the wood of Santalum spp.) and without latex. Usually hemiparasitic plants (chlorophyll is
synthetized). Bark with orange-coloured sclerenchymatic inclusions (S: Okoubaka). Trunk thorny (S:
Scleropyrum). Serial buds could be present (Fig. 25.14d L: Psittacanthus). Leaf arrangement al-
ternate (Fig. 25.14e), two-ranked (S: Okoubaka) or opposite (Fig. 25.14c) (L: Loranthus, S:
Phoradendron). Stipules absent. Leaves simple, entire, s.t. looking like small scales (S: Arceutho-
bium) or somewhat fleshy, but not coriaceous. Young leaf-folding plane. Leaves not glandular.
Venation somewhat indistinct (Fig. 25.14b) or with parallel secondary veins (Fig. 25.12 S: Exo-
carpos). Inflorescences lateral. Flowers small, often greenish (Santalaceae) or flowers showy
(Fig. 25.13), yellow-, orange- or red-coloured (Loranthaceae). Ovary unilocular. Fruits nuts or
drupes (Santalaceae), gelatinous berries or drupes (Loranthaceae).
Fig. 25.12 Exocarpos latifolia, parallel secondary veins, venation indistinct, Bali
Similar families (if arborescent): Architecture models poorly known:

OLACACEAE have also an indistinct venation. LEEUWENBERG (Fig. 25.14 -a): Viscum album
ERYTHROPALACAE: Heisteria is conforming, as (Eurasia).
Okoubaka, to MANGENOT’s model and has also a MANGENOT: Okoubaka.
two-ranked leaf arrangement. ATTIMS?: Santalum album.
25.3 Santalaceae (S) + Loranthaceae (L): 12-30-37-44 415
SANTALACEAE: trees (Fig. 25.12 Exocarpos, AS, AU; Okoubaka, AF; Osyris, AF; Nuytsia,
AU; Santalum, AS-AU; Scleropyrum, AS), shrubs, parasitic plants (Arceuthobium, AM; Den-
drophthora, AM; Fig. 25.13 Psittacanthus, AM; Viscum) or herbs (Fig. 25.13 Thesium, Paleo).
LORANTHACEAE: epiphyte and hemiparasitic herbs or shrubs (Loranthus, Phthirusa).
Fig. 25.13 Exocarpos latifolius, scaly bark, Bali, Indonesia. Psittacanthus ramiflorus, parasitic on Quercus sp. with
tubular showy flowers, Mexico (with the courtesy of Mauricio Bonifacino). Thesium alpinum, herb, Swiss Alps
Santalaceae and Loranthaceae are mainly tropical. The New World is especially rich in parasitic
Santalaceae, and the order Santalales encompasses numerous hemiparasites [5]. Only a few genera
have reached extreme Southern and Northern latitudes, where they seem out of place (Thesium).
Okoubaka aubrevillei (‘oku baku’), the fetish tree of West African healers, parasitizes neighbouring
trees with its roots [6]. The heartwood of Santalum album (India, Sri Lanka, cult. In NW Australia) is
distilled to produce scented essential oils or incense.
Fig. 25.14 Santalaceae-Loranthaceae. a LEEUWENBERG’s model showing a brief monopodial phase, leaf
arrangement opposite, e.g. Loranthus (L), Viscum; b Viscum leaf, venation weakly reticulate; c pair of opposite leaves
with venation weakly reticulate, Psittacanthus sp. (L); d superimposed serial buds or small stems (st-b-st), main stem
(t1), lateral stems (t2), leaf scar (ls), e.g. Phthirusa sp. (L); e alternate leaf arrangement, e.g. Antidaphne sp. (S), Nuytsia
(L)
25.4 Sapindaceae: 23-39-46-51-52-57-C
142 genera and 1850 species. Order Sapindales. Plants not aromatic and usually without latex. Twigs
brittle but wood of adult trees can become very hard and dense (Arytera, Dodonaea, Sarcopteryx).
Lianas are climbing by means of tendrils (Fig. 25.21) (Paullinia, Serjania, Thouinia) or are lying on
their support (Paullinia). Young internodes angular or grooved (Fig. 25.16). Leaf arrangement most
often spiral, rarely opposite (Acer, Aesculus). Stipules absent, but present in Paullinia spp. or small
basal leaflets looking like stipules (e.g. Pometia pinnata Fig. 25.21). Leaves compound pinnate
(Fig. 25.18), s.t. very large (Pometia), 2–3-pinnate (Fig. 25.19 Dilodendron), rarely simple leaves
(Dodonaea, saplings of Cupania). Leaves trifoliolate in Allophylus and several lianas, compound
palmate in Aesculus. Leaflets of pinnate leaves can be alternate (Fig. 25.18), subopposite or opposite
(Fig. 25.20) (s.t. different types in the same individual, and saplings can bear juvenile simple leaves
Fig. 25.17). Leaflets most often entire (crenulate in many lianas and in Cupania). Petiolules without
abscission joint (Fig. 25.23) (a non-Legume feature!). Leaf rachis ending in a leaflet or a mucro. Base
of petiole often enlarged or geniculate (Fig. 25.24). Leaflets with very short, and s.t. suberized,
petiolules. Inflorescences cymose (Fig. 25.19), lateral (Allophylus) or terminal (Sapindus). Flowers
small with petals united at base, with a nectariferous disk. Ovary superior, plurilocular. Fruits dry
(Fig. 25.23) or fleshy (Fig. 25.22 Sapindus saponaria), dehiscent (Fig. 25.21) or not (Fig. 25.22).
Fruit types: trisamaras (Fig. 25.23), angular/winged capsules (Blighia, Fig. 25.21 Serjania), hairy
capsules (Fig. 25.20 Nephelium), inflated capsules (Fig. 25.18 Cardiospermum). Seeds often arillate
(Litchi sinensis, Fig. 25.24 Xerospermum spp.), s.t. with sarcotesta (Nephelium lappaceum).
Fig. 25.15 Lepisanthes fruticosus, sympodial architecture, Thailand
Trees (AM: Cupania, Matayba, Melicoccus, Sapindus, Paleo-AU-OC: Arytera, Blighia, Dimo-
carpus, Lepisanthes (Fig. 25.15), Nephelium, Pometia, Schleichera, Xerospermum), shrubs (AS-AU:
Jagera, Lepisanthes), lianas (AM: Cardiospermum, Paullinia, Serjania), no herbs.
25.4 Sapindaceae: 23-39-46-51-52-57-C 417
Similar families:
ANACARDIACEAE, but usually aromatic.
MELIACEAE, some of their species also have alternate
leaflets, but their branches are stiff and elastic.
Architectures of Sapindaceae are often
sympodial and fuzzy (e.g. Fig. 25. 25).
CORNER: Jagera (AS, AU).
CHAMBERLAIN: Lepisanthes spp. (Paleo).
SCARRONE: Aesculus (AU-AS, AM-N).
RAUH: Acer (maple trees), (EU-AS, AM-N).
Fig. 25.16 Typical angular stems and pulvinate

petiolar bases of Sapindaceae
Sapindaceae are pantropical or warm temperate, they are mainly arborescent, the lianas being all
American. Several heavy and hard woods (Arytera spp., Dodonaea spp.). Allophylus is the only
pantropical genus, consisting of a single species (A. cobbe) which occurs in a range of more than a
hundred varieties. At a vegetative stage, species are very similar, so that, without flowers or fruits it is
difficult to identify the genus. Systematic treatments are essentially based on fruits. Timber: the very
hard wood of several species is used to make tool handles. Fruits eaten for their arils: litchi: Litchi
chinensis, ‘rambutan’ (‘hairy’ in Malayan language): Nephelium lappaceum (Fig. 25.20); ‘akee’:
Blighia sapida (AF-W), but seeds and unripe fruits are highly toxic! [7]; ‘longan’: Dimocarpus
sp. (AS); Senegal cherry: Lepisanthes senegalensis (AF-W); ‘matoa’: Pometia pinnata (AS). Fruits
are often very astringent. Stimulating drinks, ‘guarana’: Paullinia cupana. Soap substances (Sapindus
saponaria). Oil seeds: Schleichera oleosa (Macassar oil). Ornamental trees: Arytera spp., Harpullia
pendula (AU), Xanthoceras (AS) or liana: Cardiospermum halicabanum.
Fig. 25.17 Heterophylly: sapling with simple juvenile leaves and its first compound adult leaf (left, above), Venezuela.
Allophylus cobbe, lateral infructescence, Venezuela. Blighia sapida, 3-carpellate capsules
Fig. 25.18 Cardiospermum spp., liana with membranaceous capsules, Venezuela. Cupania scrobiculata, leaf compound
pinnate, leaflets alternate and crenulate, Belize. Cupania sp., leaflets alternate and camptodromous venation, Belize
Fig. 25.19 Deinbollia sp., lateral inflorescences in panicles, Madagascar. Dilodendron sp., bipinnate leaf, Venezuela.
Dodonaea viscosa, simple leaves and samaras, Southern Brazil
Fig. 25.20 Matayba sp., leaves pinnate and shiny, Belize. Melicoccus bijugatus, leaves paripinnate, Belize. Nephelium
lappaceum, ‘hairy fruit’ or ‘Rambutan’, Ambon, Indonesia
! Children in Latin America used to play marbles with the perfectly spherical seeds of Sapindus
saponaria.
! The origin of Sapindales is somewhat obscure, possibly in the malvacean clade, their sister group
are two herbaceous or shrubby families of arid regions in Central Asia, Africa, Mexico and Australia:
Bierbersteiniaceae and Nitrariaceae [8, 9]. These results do not imply an origin in steppe environ-
ments, however, they do support a Laurasian origin of the order.
25.4 Sapindaceae: 23-39-46-51-52-57-C 419
Fig. 25.21 Paullinia vespertillo, axillary tendril, French Guiana. Paullinia sp., red capsules containing black seeds,
Venezuela. Pometia pinnata, stipuliform basal leaflets, Seram, Indonesia
Fig. 25.22 Sapindus saponaria, indehiscent fruits (drupes?) arranged in a panicle, Venezuela. Schleichera oleosa,
small tree with fleshy capsules, Bali, Indonesia. Serjania yucatanensis, fruits, Belize
Fig. 25.23 Serjania yucatanensis, liana with winged capsules, Belize. Talisia sp., enlarged petiolar bases, Venezuela.
Talisia sp., petiolules without abscission joint, French Guiana
Fig. 25.24 Toulicia guianensis, grooved internodes, enlarged petiolar bases, Venezuela. Xerospermum sp., capsules
with arillate seeds, Sumatra
Fig. 25.25 Nephelium lappaceum.‘Rambutan’, CHAMPAGNAT’s model?, sympodial fuzzy architecture, inflores-
cences terminal (in red), Ambon, Indonesia. Author’s field book
25.5 Sapotaceae: 2-3-4-24
58 genera and 1100 species. Order Ericales. Plants producing latex, not aromatic. Latex usually pure
white (Fig. 25.29) and viscous, rarely yellow, green–blue in New Caledonian nickel accumulators
(Niemeyera spp.) [10]. Many trees with branches developing by apposition growth (Fig. 25.28e).
Short shoots modified into thorns (Fig. 25.31 Sideroxylon spp.). Leaf arrangement spiral; opposite in
25.5 Sapotaceae: 2-3-4-24 421
a few species of Pradosia (AM). Stipules usually absent, present in Ecclinusa (AM), Fig. 25.28d
Mimusops (Paleo), Palaquium, Payena (AS). Leaves simple and entire. Young leaf-folding
conduplicate (Fig. 25.26). Venation, see (Fig. 25.28a–c). Underside of leaves and young internodes
s.t. with a velvety indument (Fig. 25.26) or hairy (Fig. 25.29). Inflorescences lateral (Fig. 25.30), in
contracted cymes (Fig. 25.30), rarely in solitary flowers. Flowers with 4–18 small fused petals
(Fig. 25.27). Ovary superior, plurilocular. Fruits berries (Fig. 25.30), rarely drupes. Seeds shiny with
a distinct lens-shaped hilum (Fig. 25.29).
Fig. 25.26 Chrysophyllum cainito, conduplicate leaf- Fig. 25.27 Manilkara sp., sm: staminode, pt: petal
folding, indument rust-coloured, Yucatan
Trees, often very tall (Baillonella, Chrysophyllum, Delpydora, Manilkara, Micropholis, Mimu-
sops, Palaquium, Payena, Pouteria, Tieghemella, etc. Nickel-accumulators pachycauls (Pycnandra,
NC). Shrubs (Sideroxylon), s.t. geoxylic with subterranean stems (e.g. Pradosia brevipes, Brazil), no
lianas and no herbs (Fig. 25.32).
Sapotaceae are forest trees in all tropics, rarely found in the savannas (Sideroxylon). Timber: AF:
Baillonella toxisperma (‘moabi’), AM: Manilkara bidentata (‘balata franc’). Fruit trees: Manilkara
sapota, (‘sapotiller’), Chrysophyllum cainito (‘Cainito’), Sideroxylon obtusifolium (‘pacurero’),
moreover all species produce edible fruits, which when they drop form a carpet on the ground, an
information to be added to “survival guides” for forest trips! Seeds of shea butter tree (Vitellaria
paradoxa, AF-W) provide an important cooking oil. Latex ‘gutta-percha’, used aforetime for isolation
of first Atlantic cables, was extracted from Palaquium spp. Chicle, tapped from the tree Manilkara
zapota, was harvested for many decades and exported to the USA, where it was used in the pro-
duction of chewing gum. Flowers ‘elengi’ for perfumery (Mimusops elengi). Berries of Synsepalum
dulcificum (AF-W) contain substances which change the perception of acid and salt to sweet taste
(glycoprotein miraculine).
! Sapotaceae and Moraceae, two families that are not closely related, exhibit the same ‘architec-
tural spectrum’, i.e. CORNER’s, RAUH’s, ROUX’s, AUBRÉVILLE’s, CHAMPAGNAT’s, and
TROLL’s models. This architectural convergence is understandable because of a lateral, not terminal,
flowering in both families.
Similar families: Architectural models

(RAUH and AUBRÉVILLE dominant):
APOCYNACEAE, when these have alternate
CORNER: Delpydora spp. (AF), Planchonella
phyllotaxy (Himatanthus, AM), but branches of the
pronyensis (NC).
latter are often lenticellate.
RAUH (Fig. 25.32): Englerophytum
CLUSIACEAE, in relation to opposite-leaved magalismontanum (AF-S). Mimusops elengi (AS),
Sapotaceae. Pouteria (AM).
AUBRÉVILLE (Fig. 25.32): Baillonella spp.,
Faurea saligna, Synsepalum dulcificum, Tieghemella
beckelii, Vitellaria paradoxa (AF), Manilkara spp.
(Pantrop), Palaquium (AS).
ROUX: Chrysophyllum giganteum (AM).
CHAMPAGNAT: Sideroxylon obtusifolium (AM-S).
TROLL: Chrysophyllum cainito, Micropholis
spp.(AM).
Fig. 25.28 SAPOTACEAE. a–d. Venation. a: camptodromous: Madhuca, Palaquium; b: numerous parallel
secondary veins: Mimusops, Micropholis; c: brochidodromous with submarginal veins: Chrysophyllum, Manilkara pp.;
d: stipules enclosing a bud, e.g. Mimusops; e: Apposition growth of a plagiotropic branch, new leader develops from the
axil of a reduced leaf or cataphyll (c), i.e. AUBRÉVILLE’s model
Fig. 25.29 Delpydora macrophylla, Cameroon. Diploknema oligomera, spiral leaf arrangement, Kebun Raya, Java.
Manilkara bidentata, fruits and seed with distinct hilum, Venezuela. Micropholis sp., latex white, inner bark ocre-
coloured, French Guiana
25.5 Sapotaceae: 2-3-4-24 423
Fig. 25.30 Manilkara sp., leaves, flowers and fruit, Venezuela. Palaquium sp., fruits in lateral fascicles, Flores
Fig. 25.31 Pouteria sp., fruits in lateral fascicles, Venezuela. Sideroxylon sp., shrub of back-shore, Belize. Sideroxylon
sp., short shoots modified into thorns
Fig. 25.32 Englerophytum magalismontanum, RAUH’s model (AF-S). Baillonella toxisperma – Moabi, young tree
and adult tree, AUBRÉVILLE’s model (detail: branch), Langoué forest, Gabon. Document available for free access on
Editor’s website [11]
25.6 Schisandraceae: 23
3 genera and 73 species. Order Austrobaileyales. Plants producing essential oils (leaves with
translucent dots). Spiral leaf arrangement (Fig. 25.33c). Prophylls of lateral shoots are facing the
main shoot (Fig. 25.33a), indeed a monocotyledous feature. Flowers with numerous spirally arranged
tepals. Fruits fascicles of folicules. Shrubs (Illicium Fig. 25.33b), lianas (Schisandra).
Ornamentals: Schisandra chinensis. Spices: Illicium verum (star anise, China, Vietnam).
Fig. 25.33 a Prophyll facing the main stem (e.g. Kadsura). b Illicium verum, spiral leaf arrangement, BG LC, France.
c Illicium sp., sympodial architecture and spiral, pseudo-whorled leaf arrangement
25.7 Simaroubaceae (S) + Picramniaceae (P): 11-21-29-46-48-C
24 genera and 200 species. Orders Sapindales-Picramniales. Plants not aromatic and without latex.
Bark thin and papery (Fig. 25.37). Bitter bark (Brucea, Quassia). Spiral leaf arrangement. Stems
are exceptionnally, hollow and inhabited by ants (Picrolemma, AM-S). Stipules usually absent, but
present in American Picrasma. Leaves pinnate compound, rarely simple, entire (S: Quassia indica)
or toothed (S: Picrasma). Leaflets with an abscission joint, usually alternate (Fig. 25.36), opposite in
Simaba (Fig. 25.34-c). Underside of leaves often glaucous (Fig. 25.36). Winged rachis in Quassia
amara. Leaves without translucent dots (except in Quassia indica, syn. Samadera indica). Base of
leaflets glandular (S: Fig. 25.35 Ailanthus spp., Simaba spp.) or glands on the upper side (Odyendea,
AF; Perriera, MA). Indument usually absent, but leaves puberulous in Simaba spp. Inflorescences
racemose or cymose. Flowers small with 3–5-8 free petals (tubular in aspect in Quassia Fig. 25.36).
Ovary superior, plurilocular. Fruits samaras (Fig. 25.35), berries (P: Picramnia), or polydrupes
(Fig. 25.35 Quassia).
Trees (S: Ailanthus, AS; Brucea, Paleo; Castela, AM; Picrasma, AM-AS; Quassia, Pantrop),
shrubs (P: Alvaradoa, Fig. 25.37) no lianas and no herbs.
25.7 Simaroubaceae (S) + Picramniaceae (P): 11-21-29-46-48-C 425
Fig. 25.34 Simarouba glauca, treelet with pinnate leaves, Belize. Simaba sp., young tree with (a) sympodial trunk,
(b) pubescent stem and leaves, (c) leaf odd-pinnate with opposite leaflets
Similar families:
RUTACEAE often have a bitter bark
(Angostura) but are usually aromatic.
ANACARDIACEAE, MELIACEAE,
SAPINDACEAE have leaflets without an
abscission join.
Simaroubaceae are pantropical forest trees, they are rarely found in savannas. Quassia amara (syn.
Simarouba amara) is a source of bitters and poison in fly-papers. Ailanthus is tropical in its distri-
bution except for the warm temperate ‘tree of heaven’ (A. altissima) which has become invasive in
many regions. Ailanthus triphysa (Indian mattipal) provides scented resin and incense. A range of
biological properties has been demonstrated by the quassinoids of Simaroubaceae, including anti-
malarial, antileukemic, antiviral, insecticidal and amoebicidal properties [12].
! A broad circumscription of Quassia is unwarrented given several well-supported clades corre-
sponding to traditional generic limits (i.e. Quassia, Simaruba, Simaba, etc.) [13].
Fig. 25.35 Ailanthus altissima. Glandular (gl) leaves and samaras. Eurycoma sp., sapling, Sumatra
Fig. 25.36 Quassia amara, tubular flowers, Kebun Raya, Java. Quassia amara, polydrupe (one of five fallen), Kebun
Raya, Java. Simarouba glauca, leaflets alternate, glaucous, Belize
Fig. 25.37 Simarouba glauca flaked young bark, Belize. Alvaradoa amorphoides, shrub with alternate glaucous
leaflets, Belize
25.8 Smilacaceae: 55 (Monocotyledons)
A monogeneric family encompassing 260 species. Order Liliales, Monocotyledons. Dioecious rhi-
zomatous (Fig. 25.38), not aromatic plants. No latex. Stems often spiny. Spiral leaf arrangement.
Stipules absent. Leaves simple, entire or lobate. Venation campylodromous (Fig. 25.38). Petioles
bearing a basal pair of tendrils (Fig. 25.39). Plants glabrous. Inflorescences in panicles, terminal or
lateral (rarely uniflorous). Flowers have 3 + 3 small tepals and 6 stamens. Ovary superior, 1- or 3-
locular. Fruits berries (Fig. 25.39) enclosing 1–3-n seeds.
Lianas (Smilax, incl. Heterosmilax).
Smilacaceae can be founds along trails and in clearings. Fortifying beverages: Smilax china.
Ornamentals: Smilax anceps.

DIOSCOREACEAE with same venation but TOMLINSON: probably dominant or exclusive in this
without tendrils and usually twining. family.
25.9 Solanaceae: 11-12-21-30-54 427
Fig. 25.38 Smilax domingensis, Belize. a Leaves with campylodromous venation and b sympodial rhizome
a b
Fig. 25.39 Smilax domingensis, Belize. a Petiole bearing a pair of tendrils and b infructescences of orange-coloured
berries
25.9 Solanaceae: 11-12-21-30-54
88 genera and 2650 species. Order Solanales. Plants s.t. aromatic: crumpled leaves may give out
strange smells. Plants without latex. Stems s.t. spiny (Fig. 25.44), the spines epidermic (Solanum) or
developed from short shoots (Grabowskia, Latua, Lycium). Lateral axes partially fused with the
petiole (Fig. 25.40a concaulescence), this phenomenon associated with anisophylly (Fig. 25.40d).
Leaf arrangement alternate or appearing opposite. Stipules absent. Leaves simple, entire or lobate
leaving s.t. a prominent petiolar scar (Fig. 25.40b). Venation pinnate, often camptodromous, s.t.
secant (Fig. 25.41). Stems or leaves pubescent. Inflorescences cymose or flowers lateral (Fig. 25.42
Cestrum). Flowers showy (Fig. 25.41), (inflated or persistent calyx in Nicandra, Physalis Fig. 25.42
Cestrum), with 5 fused petals (Fig. 25.43) and 5 stamens, often pollinated by moths and scented from
nightfall (Fig. 25.42 Cestrum), or by solitary bees. Ovary superior, 2 or 4-locular. Fruits berries
(Cestrum, Solanum) or capsules (Datura with numerous seeds).
Fig. 25.40 Stem morphology. a Concaulescence (e.g. Datura), a lateral axis is partially fused with a petiole (P) of a
subtending leaf, a-b: prophylls; b prominent petiolar scar (psc); c LEEUWENBERG’s or KORIBA’s models in
Solanum; d Witheringia sp., herb with anisophylly and pseudodistichous leaf arrangement (modules alternatively
represented in black and white); e decumbent branch, CHAMPAGNAT model, e.g. Cestrum
Small trees (Solanum betaceum), shrubs (Brugmansia, Brunfelsia, Cestrum, Lycium, Solanum),
lianas (Markea, Solanum), s.t. with hollow stems inhabited by ants (Markea) or herbs (Solanum).

BORAGINACEAE (also pubescent). LEEUWENBERG (Fig. 25.40 -c): Brugmansia sp.
(shrub, Andes); Solanum betaceum (tree, AM-S).
KORIBA (Fig. 25.40 -c): Lycianthes pauciflora
(liana, Guianas); Solanum auriculatum (tree, AM).
CHAMPAGNAT (Fig. 25.40 -e): Cestrum sp.
(shrub, Guianas), Lycium afrum (shrub, AF-S),
Solanum tampicense (shrub, AM-S).
Solanaceae are mainly American (Brugmansia, Cestrum, Datura, Iochroma, etc. as by far, the
majority of the 1200 Solanum species), they grow along trails and in clearings. Food or condimentary
plants (potato, Solanum tuberosum; ‘aubergine’, Solanum melongena; tomato, Solanum lycoper-
sicum; peppers and red chili, Capsicum annuum). Psychotropic (tobacco, Nicotiana tabacum), often
hallucinogenic (Atropa, Brugmansia, Datura). Ornamentals (Brugmansia, Brunfelsia, Lycium,
Markea, Petunia).
! No known timber tree in Solanaceae, the family has predominantly herbaceous features.
! Flowers of several species (e.g. Solanum) with tubular anthers require buzz pollination by
solitary bees or bumblebees.
25.9 Solanaceae: 11-12-21-30-54 429
Fig. 25.41 Different types of leaves; a cordate, b lanceolate, c dentate or crenate. Brugmansia sp., trumpet flowers BG
Sirikit, Thailand (orig. AM)
Fig. 25.42 Cestrum sp., flowers tubular, pollinated by moths, BG Sirikit, Thailand. Physalis sp., inflated calyxs, BG
Sirikit, Thailand. Solanum betaceum, fruits drooping, Southern Brazil
Fig. 25.43 Solanum subinerme, spiny stem, French Guiana. Solanum wrightii, hairy leaves, Venezuela. Solanum sp.,
cymose infructescences, Belize
Fig. 25.44 Nicandra physaloides, winged persistent calyxs, BG Geneva, Solanum sp., spiny stems, Venezuela
25.10 Styracaceae: 12-22
11 genera and 170 species Leaves entire or toothed. Venation brochidodromous or camptodromous.
Leaves often discoloured in American Styrax. Indument of scaly-peltate hairs (Fig. 25.45a–c) or
stellate hairs (Fig. 25.45b). Fruits dehiscent capsules (Asian Styrax Fig. 25.45d), sunken in a per-
sistent cupular calyx in American Styrax (Fig. 25.45e), or drupes (Parastyrax, Burma).
Benzoin resin is extracted from Styrax benzoin, a tree native to Sumatra and, to a lesser extent from
S. benzoides, native to Tailand [14]. Benzoin is used in perfumery and body care and in past times by
the Malays in many ceremonies. Considering their American-Asiatic distribution, Styracaceae
probably have a Laurasian origin, like Symplocaceae, a closely related family.
Fig. 25.45 a Styrax leprosum, numerous minute hyaline peltate hairs, leaf entire, Paraguay. b Styrax serrulatus,
scattered stellate hairs, leaf serrulate, Assam, India. c Styrax parallelodromus, scattered minute lepidote hairs, Sumatra.
Styrax spp. d Asian Styrax with dehiscent fruits; e American Styrax with indehiscent fruits inserted in a cupule. (Drawn
from herbarium specimens, CBG H)
25.11 Surianaceae: 21
A very small family containing 5 genera and 8 species. Order Fabales. Branches sympodial. Spiral
leaf arrangement. Leaves linear, entire. Venation indistinct. Flowers with parts in five, axillary
(Fig. 25.46). Mainly Australian, except for a frequent littoral shrub (Fig. 25.46) of all tropical coasts:
Suriana maritima.
Fig. 25.46 Suriana maritima, back-shore, Belize. Flowering stem, back-shore, short shoots with clustered leaves
25.12 Symplocaceae: 30 431
Similar families:
SAPOTACEAE:
Eusideroxylon, also with KORIBA: Suriana maritima.
clustered leaves, but latex!
25.12 Symplocaceae: 30
Family encompassing a single genus and 320 species. Order Ericales. Plants not aromatic and without
latex. Several species are aluminium accumulators (i.e. leaves turn yellow after drying). Spiral leaf
arrangement. Stipules absent. Leaves simple, entire or toothed, glabrous or pubescent. Venation
pinnate, brochidodromous (Fig. 25.47). Inflorescences lateral (Fig. 25.48), racemose or cymose.
Flowers small, with 3–5 fused petals and 5-n stamens. Ovary inferior, plurilocular. Fruits are
drupes, often ripening blue (Fig. 25.47). Small trees or shrubs, no lianas and no herbs.
Fig. 25.47 Symplocos guianensis, brochidodromous venation, French Guiana. Symplocos racemosa, flowering lateral,
Thailand. Symplocos paniculata, blue drupes (with the courtesy of Blake Willson)
Fig. 25.48 Symplocos racemosa, rhythmic growth, racemes of fruits, Thailand (author’s field book)

THEACEAE (Gordonia, Schima) have
also entire leaves with a brochidodromous RAUH probably exclusive in this
venation. monogeneric family.
Symplocaceae have a disjunct Asian-American distribution, they grow in savannas and open
forests. Few uses are known (S. cochinchinensis, medicinal).
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(Simaroubaceae) based on chloroplast and nuclear markers. International Journal of Plant Sciences 168: 1325–
1339.
14. Burkill, I.H. 1935. A dictionary of the economic products of the Malay peninsula, Vols. I and II, 2402 pp. Oxford:
University Press.
Tetramelaceae to Torricelliaceae
26
26.1 Tetramelaceae: 20
A small family of 2 genera and 2 species. Order Cucurbitales, separated here from Datiscaceae. Big
buttressed trees (Fig. 26.1) with soft wood. Spiral leaf arrangement. Leaves large, simple
(Fig. 26.2a), not stipulate, entire (Octomeles) or toothed (Tetrameles). Petiole decurrent on stem
(Fig. 26.2). Indument of simple (Fig. 26.2) or peltate (Fig. 26.2) hairs. Inflorescences in spikes or
panicles of reduced flowers. Flowers unisexual, with 6–8 petals and numerous stamens. Fruits
capsules. Winged seeds in Tetrameles.

MALVACEAE, but fibrous MASSART (Fig. 26.2):
bark. Octomeles sumatrana (AS).
Fig. 26.1 Octomeles sumatrana, impressive buttresses, Seram, Indonesia
https://doi.org/10.1007/978-3-031-05942-1_26
434 26 Tetramelaceae to Torricelliaceae
Fig. 26.2 Octomeles sumatrana, a: petioles decurrent on stem, peltate hairs; b: Tetrameles nudiflora, indumentum of
thick hairs. Octomeles sumatrana (‘Pohon tabu’), young tree conforming to MASSART’s model, Seram, Indonesia.
Octomeles sumatrana, petioles decurrent on stem, Indonesia
! Roots embracing Angkor Wat temples (Cambodia) are not those of fig trees but of Tetrameles
nudiflora (AS-SE, AU).
26.2 Theaceae: 12-29-30
7–8 genera and 225 species. Order Ericales. Several species are aluminium accumulators. Plants not
aromatic and without latex. Spiral leaf arrangement. Stipules absent. Leaves simple, entire or
toothed. Young leaf-folding plane (Schima Fig. 26.4) or somewhat convolute (Camellia). Venation
brochidodromous (Fig. 26.3). Lateral inflorescences of solitary flowers (Fig. 26.4). Flowers with 5-n
petals, somewhat large in Camellia and Gordonia (Fig. 26.4), white, pink or red. Stamens very
numerous. Ovary superior, plurilocular with axil placentation. Fruits capsules (Fig. 26.4), (Camellia,
Gordonia incl. Polyspora) or drupes.
Fig. 26.3 Camellia sinensis, venation brochidodromous, capsule, Thailand

26.3 Thymelaeaceae: 10-25-44-54-56 435
Fig. 26.4 Camellia sinensis in cultivation, fruits, Azerbaijan. Polyspora (syn. Gordonia) amboinensis. a: flower with
numerous stamens; b: leaves slightly crenulate, lateral, subterminal flowering (cp) capsule, New Guinea, CBG H
specimen; c: Capsule with winged seeds. Schima wallichii, young leaves flat, Thailand
Small trees or shrubs (Camellia, Gordonia, Schima), no lianas and no herbs.

SYMPLOCACEAE and PENTAPHYLACACEAE, two other RAUH: Gordonia, Schima wallichii (AS-SE. TROLL:
families of the order Ericales. Camellia japonica, Stewartia (China, Japan).
Theaceae are absent in Africa, they share with Symplocaceae an Asian-American discontinuous
distribution. Emblematic are tea plant, Camellia (*Thea) sinensis, and hundreds of ornamental
cultivars of Camellia japonica. Gordonia spp. are often grown in tropical gardens for their large
white flowers. Schima wallichii is used for afforestation and as a source of tannins.
! Formerly Theaceae and Pentaphylacaceae were treated as the subfamily rank (resp. Camellieae
and Ternstroemieae in the Theaceae) [1].
26.3 Thymelaeaceae: 10-25-44-54-56
46 genera and 860 species. Order Malvales. Plants not aromatic and without latex. Bark fibrous
(Fig. 26.5, Fig. 26.6). Leaf arrangement spiral (Daphne, Fig. 26.7 Lasiosiphon), two-ranked
(Aquilaria, Fig. 26.6 Dicranolepis, Gonystylus), or opposite (Fig. 26.7 Phaleria). Stipules absent.
Leaves simple, entire. Venation pinnate, brochidodromous. Leaves without glands, with translucent
dots in Gonystylus. Indument absent (Daphne) or appressed hairs (Aquilaria, Gonystylus). Venation
s.t. with fimbrial veins (Fig. 26.6 Aquilaria). Inflorescences lateral, s.t. born directly on trunk
(Fig. 26.7 Phaleria). Flowers small, with 4-5-6 petals. Ovary superior, plurilocular. Fruits berries
(Fig. 26.7), drupes or capsules.
436 26 Tetramelaceae to Torricelliaceae
Similar families:
When leaves opposite: with Oleaceae but
these not fibrous.
When leaves spiral: with Pentaphylacaceae,
but these not fibrous.
When leaves two-ranked: with Annonaceae,
but these with aromatic bark.
LEEUWENBERG: Edgeworthia (AS).
RAUH: Daphne (EU).
MASSART: Aquilaria (AS).
MANGENOT: Dicranolepis (AF).
Fig. 26.5 Fibrous bark of the Thymeleaceae
Fig. 26.6 Aquilaria crassna, BG Sirikit, Thailand. Internal bark fibrous. Fimbrial vein. Dicranolepis cf. persei, lateral
fruit, Cameroon
Fig. 26.7 Lasiosiphon sp., a Malagasian shrub resembling Daphne species. Phaleria sp., KORIBA’s model, leaf and
fruits inserted on trunk. Phaleria macrocarpa, in cultivation, Bali
Reference 437
Trees: Aquilaria (AS), Gonystylus (AS, OC). Shrubs: Craterosiphon (AF), Daphne, Edgeworthia
(Eurasia), Gnidia (Paleo), Lophostoma (AM), Phaleria (AS, OC). Several lianas: Craterosiphon and
Dicranolepis (AF), Enkleia (AS).
Thymelaeaceae are pantropical and warm temperate in their distribution. Timber (Gonystylus).
Bark fibre for paper (Daphne, Gnidia, Thymelaea, Wikstroemia) or ropes (Dais, Daphnopsis). Incense
(Aquilaria). Ornamentals (Daphne, Dirca, Edgeworthia, etc.). Medicinals: Phaleria macrocarpa
(‘mahkota dewa’, God’s crown).
! The most remarkable species belong to the Indomalayan genus Aquilaria. Due to the effect of
fungal pathogens, the inner bark of Aquilaria malaccensis and A. crassna produces scented sub-
stances during the rotting process, which are used as incense (gaharu). However harvesting is a lottery
as the harvested tree may not host the precious pathogen!
26.4 Torricelliaceae: 26
A very small family with 3 genera and 11 species. Order Apiales. Aralidium pinnatifidum (West
Malesia) is a small tree of the understorey conforming to CHAMBERLAIN’s model. In fact, Ara-
lidium, with their sheathing petioles (Fig. 26.8) and deeply incised leaves (Fig. 26.8) will perfectly
match with Araliaceae. Inflorescences cymose. Fruits white berries or drupes.
Fig. 26.8 Aralidium pinnatifidum, Khao Chong, Thailand. Sheathing petioles and pinnatifid leaves
Reference
1. Willis, J.C. 1973. A dictionary of the flowering plants and ferns, 8th ed, 1246 + 66 pp. Cambridge: Cambridge
University Press.
Ulmaceae s.l. to Urticaceae
27
27.1 Ulmaceae (U) + Cannabaceae (C): 5-6-8-31
7 genera, 50 species (U) and 9 genera, 90 species (C). Order Rosales. Plants not aromatic and without
latex. Trunk bark with scars of fallen branches (Fig. 27.2). Bark fibrous, often with a chlorophyllous
layer plastered to bark (Fig. 27.1). Ulmaceae have a continuous branching in trunk or even in
branches (Fig. 27.3). Short twigs modified into spines (C: Celtis iguanaea, AM-S). Leaf arrangement
two-ranked (C: Fig. 27.3 Celtis, Trema, Holoptelea, U: Ulmus, Zelkova), rarely opposite (C:
Lozanella). Stipules present, enclosing bud (C: Chaetacme, Gironniera). Leaves simple, entire or
toothed. Venation pinnate (U: Holoptelea, Ulmus, Fig. 27.2 Zelkova) or tripliveined (C: Fig. 27.3
Celtis, Trema). Leaves without glands and usually pubescent (glabrous Celtis philippensis). Flowers
small, wind-pollinated Fig. 27.2. Ovary superior, plurilocular (U) or unilocular (C). Fruits samaras
(U: Fig. 27.3 Holoptelea, Ulmus), drupes (C: Fig. 27.3 Celtis). Trees (U: Ampeloceras, Holoptelea,
Ulmus; C: Celtis), herbs (C: Cannabis).
Ulmaceae have a single tropical genus: Holoptelea (two species, one in India and one in Africa).
Ulmaceae and Cannabaceae grow in open sites and along trails, Trema orientalis (AS) and
T. micrantha (AM) are very frequent pioneer trees. Uses: charcoal, fibres for weaving and fishing nets.
! Ulmaceae are very close to Cannabaceae and the segregation of these two families is not a
definitive issue.
https://doi.org/10.1007/978-3-031-05942-1_27
440 27 Ulmaceae s.l. to Urticaceae
Fig. 27.1 Trema orientalis, branch with continuous ramification, Cameroon. Trema orientalis, fibrous bark and
chlorophyllous layer, Thailand. Trema orientalis, ROUX’s model, Bali, Indonesia
Fig. 27.2 Ulmus laevis, male flowers, Switzerland. Holoptelea integrifolia, trunk with ‘eye-marks’, an Indian tree,
Thailand. Zelkova carpinifolia, leaves crenulate, pinnate venation, Azerbaijan
Fig. 27.3 a: Celtis philippensis, continuous branching and leaves tripliveined, Seram, Indonesia (author’s field book);
b: Celtis australis, toothed leaves and drupe, in cultivation; c: Holoptelea sp., leaves two-ranked, samaras, x: abortion
of an apical meristem, herbarium specimen, CBG H
27.2 Urticaceae: 2-6-14-15-31-47 441
27.2 Urticaceae: 2-6-14-15-31-47
48 genera and 1600 species. Order Rosales. Plants not aromatic and usually without latex. Bark
producing a white latex in Broussonetia papyrifera (Fig. 27.7) or a brownish exudate in Cecropia
spp. Small stilt-roots in Pourouma spp. (Fig. 27.11). Trunk or branches usually lenticellate. Cecropia
(AM) have hollow stems inhabited by Azteca ants. Petiolar bases bear Muellerian bodies intended for
attracting insects (Fig. 27.8). Inner bark s.t. fibrous (Fig. 27.6). Leaf arrangement spiral (Cecropia,
Coussapoa, Dendrocnide, Pourouma), two-ranked (Fig. 27.5 Elatostema, Leucosyke), s.t. opposite
(Fig. 27.6 Boehmeria). Stipules present (Fig. 27.6), s.t. hood-like and leaving an annular scar
(Fig. 27.8 Cecropia, Musanga). Stipules of the same pair are often asymmetrically disposed (Fig. 27.6
Dendrocnide) (Fig. 27.4). Leaves often variable in shape according to individual stage (Fig. 27.7
Broussonetia, Fig. 27.11 Pourouma). Leaves simple, entire, lobate (Cecropia) or s.t. (almost?)
compound palmate (Fig. 27.11 Musanga cecropioides, Myrianthus, Pourouma cecropiifolia), s.t.
with its base distinctly asymmetric (Fig. 27.5 Elatostema), rarely peltate (Pilea peperomioides,
herb, Southern China). Petioles s.t. slightly enlarged distally (Fig. 27.8 Coussapoa spp.). Venation
tripliveined (Debregeasia, Dendrocnide, Fig. 27.9 Laportea), palmate (Fig. 27.11 Cecropia) or
camptodromous and scalariform (Fig. 27.11 Coussapoa, Pourouma). Plants pubescent, often with
stinging hairs (Dendrocnide, Laportea, Urera, Urtica). Inflorescences lateral (Fig. 27.9), often
cymose (Fig. 27.10) or umbels of spikes (Fig. 27.7 Cecropia). Flowers reduced. Ovary superior,
unilocular. Fruit achenes, nuts or drupes (Fig. 27.10 Dendrocnide, Pourouma). Trees (Cecropia,
Coussapoa, Dendrocnide, Musanga, Myrianthus), shrubs (Boehmeria, Leucosyke), epiphytes
(Coussapoa pp.) or herbs (Elatostema, Laportea, Pilea, Urtica).
Fig. 27.5 Elatostema sp., asymmetrical leaves, Flores,

Indonesia
Fig. 27.4 Cecropia peltata, Belize

Fig. 27.6 Fibrous bark and lenticels, e.g. Broussonetia, Dendrocnide. Stipules (s1, s2) asymmetrically disposed (e.g.
Dendrocnide). Leaves opposite and toothed and small stipules (e.g. Boehmeria)
Fig. 27.7 Broussonetia papyrifera, cult., Switzerland, trunk producing latex and variation in leaf form, from simple
(1–2) to lobate (3–6). Cecropia sp., male inflorescence consisting of an umbel of spikes, Sierra de Lema, Venezuela
Fig. 27.8 Boehmeria tsaratananensis, lateral inflorescences, Madagascar. Cecropia peltata, base of petioles with
Muellerian bodies, Belize. Coussapoa oligocephala, petioles slightly enlarged distally, camptodromous venation,
Belize
27.2 Urticaceae: 2-6-14-15-31-47 443
Fig. 27.9 Coussapoa sp., inflorescences in heads, Sierra de Lema, Venezuela. Debregeasia longifolia, lateral
flowering on older growth units, Thailand. Dendrocnide stimulans, sapling, leaves tripliveined, Bali, Indonesia
Fig. 27.10 Laportea aestuans, herb, palmate venation, Guadeloupe. Dendrocnide moroides, compound infructescence
with small drupes, BG LC, France. Laportea sp., inflorescences lateral and cymose, palmate venation, Madagascar
Fig. 27.11 Stem and deeply incised, almost compound, leaf of Musanga cecropioides or Pourouma cecropiifolia. (to
the left: trunk of a strangler fig tree, Moraceae). Pourouma sp., stilt-roots, Lema, Venezuela. Pourouma bicolor,
sapling, heterophylly: basal leaves oval-shaped, distal leaf lobate, Belize

ULMACEAE-CANNABACEAE and MORACEAE. TOMLINSON: Boehmeria cylindrica (AS).
Both groups have stipulate, pubescent tripliveined RAUH: Laportea stimulans (Indonesia); Pourouma
leaves. However, URTICACEAE never conform to cecropiifolia (Amazonia); Cecropia peltata, (AM);
ROUX’s model and MORACEAE produce latex, Musanga cecropioides (AF).
although s.t. scanty at the end of the dry season.
AUBREVILLE: Dendrocnide microstigma (Indonesia);
Pourouma minor (Guianas).
CHAMPAGNAT: Boehmeria macrophylla (tree, AF-E);
Myriocarpa longipes (shrub, AM-C); Urera
caracassana (shrub, AM).
TROLL: Leucosyke capitellata (Indonesia); Elatostema
repens (herb, AS); Oreocnide rubescens (AS).
Urticaceae are pantropical, but Musanga and Myrianthus are African and Cecropioideae
(Cecropia, Coussapoa, Pourouma) American. Usually pioneer trees in secondary forests, clearings
and along trails: ‘yagrumos’ or Cecropia (AM), ‘parasolier’ or Musanga cecropioides (AF). Fruits of
Pourouma cecropiifolia taste of grapes (Guianas). Source of fibres: Boehmeria nivea, Oreocnide,
Urera, etc.), Broussonetia papyrifera was used for the manufacture of paper and clothes. Several
species are extremely urticant and harmful. In Queensland, the terrible ‘gympie-gympie’ (Den-
drocnide moroides) can be identified (before rubbing against it!) by its slightly peltate leaves (see also
D. peltata in New Guinea). Other urticant plants: Dendrocnide spp. and Laportea spp. (AS).
! Urticaceae provide a nice example of evolution starting from trees and leading to herbs [1].
Reference
1. Hallé, F., & R. Keller. 2019. Mais d’où viennent les plantes? 180 pp. Actes Sud.
Verbenaceae to Vochysiaceae
28
28.1 Verbenaceae + Callicarpa: 23-35-38-40-41-44-56
32 genera and 1000 species. Order Lamiales. Plants s.t. aromatic (Aloysia, syn. Lippia), without latex.
Branches usually lenticellate. Internodes quadrangular. Opposite leaf arrangement. Stipules absent.
Leaves simple, entire or toothed, s.t. with glands (Citharexylum). Indument of simple or stellate hairs
(Fig. 28.1 Callicarpa). Inflorescences usually terminal. Flowers with a radial symmetry or weak
bilateral symmetry (Fig. 28.3), 4–5 fused petals. Ovary at first bilocular then dividing into 4
uniovulate compartments. Fruits drupes (Fig. 28.3) enclosing 1-2-4 stones or groups of free
mericarps.
Similar families:
ATTIMS: Lantana camara (AM,
LAMIACEAE (Gmelina), but their invasive). CHAMPAGNAT (2):
branches are not lenticellate. Callicarpa arborea (AS).
Small trees (Fig. 28.2 Callicarpa, Fig. 28.2 Citharexylum), shrubs (Fig. 28.3 Duranta, Lantana,
AM, cult., naturalized, even invasive), subshrubs (Aloysia, Stachytarpheta), twining lianas (Fig. 28.3
Petrea volubilis) or herbs (e.g. Stachytarpheta).
Verbenaceae are very close to Lamiaceae and the status of several genera has been debated
(Callicarpa and Tectona ought to be placed in Lamiaceae). Both families grow in dry deciduous
forests, savannas or secondary forests. Duranta erecta, Lantana camara and Stachytarpheta
cayenensis has become invasive almost everywhere in the tropics. Timber: Citharexylum (fiddle
wood, this name being a distortion of ‘bois-fidèle’). Plants aromatic: Aloysia. Ornamentals: Duranta
erecta, Lantana camara, Callicarpa spp. Invasives: Lantana camara, Stachytarpheta cayenensis.
! Aloysia citrodora (syn. A. triphylla, Lippia citrodora), lemon verbena plant, can be considered a
cultigen, that is, a plant cultivated from time immemorial that should not be considerate as a true
‘wild’ species.
! Lamiaceae and Verbenaceae were placed by John Hutchinson [1] at the very top of two diverging
phyla: Herbaceae for the first and Lignosae for the latter. Molecular systematics led to a drastically
different conclusion, almost uniting these two families in a single one.
https://doi.org/10.1007/978-3-031-05942-1_28
446 28 Verbenaceae to Vochysiaceae
Fig. 28.1 Callicarpa sp., indument of stellate hairs, Thailand
Fig. 28.2 Callicarpa arborea, CHAMPAGNAT’s model, Thailand. Citharexylum sp., small tree in the savanna,
Venezuela. Duranta erecta, infructescences of drupes, naturalized, Bali, Indonesia
Fig. 28.3 Lantana camara, shrub, flowers with a weak bilateral symmetry and infructescence of drupelets, Belize.
Lantana involucrata, fruit or group of drupelets, Guadeloupe. Petrea volubilis, liana, Shipstern, Belize
28.2 Violaceae: 8-9-18-33-34-54 447
28.2 Violaceae: 8-9-18-33-34-54
29 genera and 815 species, of which 225 Rinorea. Order Malpighiales. Plants not aromatic, without
latex. Sympodial (Leonia, American Rinorea Fig. 28.7), s.t. with abortion of trunk apical meristems
(Fig. 28.6 Rinorea) or monopodial (Paypayrola, paleotropical Rinorea, Viola). Leaf arrangement
spiral (Leonia, Fig. 28.5 Paypayrola), two-ranked (Fig. 28.7b Rinorea) or opposite (Fig. 28.7c
Rinorea, AM). Stipules present. Leaves simple, entire or toothed, without glands. Underside of
lamina satiny pale green (Fig. 28.7, a feature existing also in European violets!). Young leaf with
involute folding. Indument of simple, not stellate hairs. Inflorescence lateral (Fig. 28.4) or terminal
(Fig. 28.5). Flowers bilateral, in their symmetry (Fig. 28.4) or almost with a radial symmetry
(Fig. 28.5 Paypayrola), perfectly radial in Melicytus, with 5 free petals imbricate or convolute in bud
and 3 carpels. Ovary with parietal placentation. Fruits capsules (Fig. 28.6).
Small trees (Leonia, Melicytus, Paypayrola), shrubs (Hybanthus, Rinorea), lianas (Anchietea,
Corynostylis) or herbs (Hybanthus, Viola).
In spite of the many different vernacular names refering to Rinorea species in Colombia-
Venezuela: ‘canilla de viejo’, ‘pata de grulla’, in Peru: ‘majaras caspi’, ‘yurac varilla’, AF-W:
‘olobohoro’, etc. [2, 3], there are few traditional uses for this genus. Ornamentals are extratropical
(Melicytus, Viola), essential oils (Viola odorata).
! In the Caura basin (Venezuela), fruits of Leonia glycycarpa (Fig. 28.5) have a scent of violets, so
the origin of the herbaceous genus Viola may be found in their tropical woody ancestors !
Similar families: MASSART: Paypayrola sp. (Guianas).
EUPHORBIACEAE-PHYLLANTHACEAE, RAUH: Melicytus macrophyllus (New Zealand).
however with conduplicate leaf-folding.
FAGERLIND (Fig. 28.7a): Rinorea riana (Guianas).
SALICACEAE (e.g. Casearia) often have
capsules with parietal placentation. TROLL: Hybanthus phyllanthoides (shrub, Guianas).
Fig. 28.4 Hybanthus phyllanthoides, solitary axilar flowers with bilateral symmetry, Venezuela
Fig. 28.5 Leonia cymosa, infructescence terminal, Venezuela. Leonia glycycarpa, fleshy capsules, Venezuela.
Paypayrola sp., lateral spike, flowers have almost a radial symmetry, Venezuela
Fig. 28.6 Rinorea flavescens, 3-valvate capsules, Venezuela. Rinorea sp., trunk apex with parenchymatized meristem
and bearing three plagiotropic branches, Venezuela
Fig. 28.7 Architectural variation in American Rinorea. a: FAGERLIND’s model; b: sympodial branch and two-
ranked leaf arrangement; c: sympodial branch and opposite leaf arrangement. Underside of leaves satiny pale green,
Rinorea deflexiflora (Belize), Viola biflora (Swiss Alps)
28.3 Vitaceae: 5-6-14-46-47-55-57
16 genera and 900 species. Order Vitales. Plants not aromatic, without latex. Trunk cross-section of
lianas with radiating large rays (Fig. 28.10, Fig. 28.12a) (Tetrastigma, Vitis). Trunk s.t. spiny in
Leea (Fig. 28.10) or lenticellate (Fig. 28.12g). Lianas are climbing by means of oppositifoliate
tendrils (Fig. 28.12b). Inner bark fibrous for some species (Fig. 28.10). Leaf arrangement spiral
28.3 Vitaceae: 5–6-14-46-47-55-57 449
(Leea) or two-ranked (Vitis, AS). Stipules present, large and appressed one to another in Leea
(Fig. 28.10) Leaves simple (Fig. 28.12e), entire, toothed or lobate (Fig. 28.9 Cissus) or leaves
compound (Fig. 28.12d Leea, Fig. 28.11 Tetrastigma). Leaves without glands. Venation pinnate
(Fig. 28.9 Cyphostemma) or palmate (Fig. 28.9 Cissus, Vitis). Inflorescences cymose (Fig. 28.8),
terminal and oppositifoliate, thyrses or corymbs. Flowers small, with a radial symmetry and 4–5
petals. Ovary 1-2-4-locular. Fruits berries.
Small trees (Leea spp.), with bottle-like trunks in arid environments (Fig. 28.9 Cyphostemma,
AF), lianas (Cissus, pantropical; Tetrastigma, AS, AU; Vitis, Northern Hemisphere), herbs (Leea).
Fig. 28.8 Cissus sp., inflorescence cymose (thyrse), Southern Brazil
Fig. 28.9 Cyphostemma juttae, pachycaulous shrub, BG Geneva. Cissus sp., lobate leaves with palmate venation,
Belize. Leea sp., leaves 3 x compound, inflorescence a corymb of flower buds, Madagascar
Fig. 28.10 Leea sp., large stipules appressed one to another, Madagascar. Leea sp., spiny trunk, in cultivation, Bali,
Indonesia. Leea sp., inner bark of a small tree with a network of fibres or rays?, Seram, Indonesia
Fig. 28.11 Tetrastigma sp., liana with palmate leaves, Sumatra. A giant flower of Rafflesia arnoldi, Bukittingi,
Sumatra
Before APG classifications, Vitaceae have been placed near Rhamnaceae (Rosales), then in Vitales
(APG III) and actually in Saxifragales (APG IV). Their origin remains a mystery. The ‘rosoid’ teeth
of the family [4] is a character which deserves more attention. Vitaceae occur mainly in Asia and
Africa. Lianescent Tetrastigma species host the enormous flowers of Rafflesia (Fig. 28.11). Fruiting
lianas (vines): Vitis labrusca, Vitis vinifera. Ornamentals (Cissus, Leea).
! Are shoot-tendrils of Vitis vinifera terminal or lateral? This subject caused passionate debates in
the middle of the last century [5], however the homology of shoot-tendrils with inflorescences has
been accepted for a very long time [6].
28.4 Vochysiaceae: 33-42 451
Fig. 28.12 Vitaceae a Multi-layered bark and wood with large rays. b Heterophylly in Tetrastigma sp., leaves simple
or trifoliolate. d Pinnate leaves (Leea spp.). e Simple leaf (Leea spp.). f Rosoid teeth [4], (typical in Vitaceae).
g Lenticellate stem (Leea spp.)
28.4 Vochysiaceae: 33-42
7 genera and 200 species. Order Myrtales. Plants not aromatic, without latex. Bark often scaly.
Branching monopodial (Fig. 28.16f), rhythmic (Fig. 28.15), or sympodial (Fig. 28.16g). Opposite
leaf arrangement. Stipules present, s.t. with a gland nearby on the stem (Fig. 28.16b Qualea). Leaves
simple, entire, without glands. Young leaf-folding conduplicate (Fig. 28.13). Venation pinnate, with
numerous secondary veins in Qualea (Fig. 28.16e) and Ruizterania, s.t. an intramarginal vein
(Fig. 28.16c–d, Vochysia spp.). Inflorescences in racemes or thyrses. Flowers showy, with a strong
Fig. 28.13 Vochysia pygmaea, raceme of yellow flowers, most of them still flower buds, calyx spur-like (sp), open
flowers with 3 petals. Vochysia sp., conduplicate young leaves, Venezuela. Capsules 3-locular: Vochysia (V), Qualea (Q)
Fig. 28.14 Erisma uncinatum, samara, Venezuela. Ruizterania ferruginea, large abaxial petal, Venezuela. Qualea sp.,
leaves with numerous parallel veins, detail: cupuliform glands (gl) by the stipules (st), Guianas
Fig. 28.15 Erisma uncinatum, flushing stem (rhythmic growth), Venezuela. Qualea dinizii, emergent reiterated trees
exhibiting crown shyness, Gran Sabana, Venezuela
bilateral symmetry, yellow (Vochysia), white or pink (Qualea, Ruizterania), with 3 petals or only one
developed petal (Fig. 28.14). Ovary 3-locular. Fruits winged samaras (Fig. 28.14 Erisma) or 3-
locular capsules (Fig. 28.13 Vochysia, Qualea). Trees (Qualea, Ruizterania, Vochysia, AM) and
shrubs (Callisthene, Brazil).
Vochysiaceae are all American, except two West African genera: Erismadelphus (2 species) and
Korupodendron (1 species), which corroborates the hypothesis of continental drift. The family can be
compared, from the functional point of view, to the Dipterocarpaceae [7], because of several common
traits: tall trees, somewhat gregarious, exhibiting crown shyness (Fig. 28.15), (see also Chrysobal-
anaceae), with architectural metamorphosis, and whose fruits are samaras. Vochysiaceae are
emblematic of sparsely exploited forests of the Guianas and Brazilian cerrados: ‘mandioqueira’
(Qualea paraensis), ‘bananheira do campo’ (Salvertia convallariidora), ‘pau novo’ (Vochysia
magnifica), ‘mureillo’ (Erisma uncinatum), ‘guarapo amarillo’ (Qualea dinizii), ‘saladillo’ (Vochysia
surinamensis), etc. Their ecomomic importance is linked to construction wood.
Similar families:
SCARRONE: Vochysia meridensis (AM).
MYRTACEAE, but these are not stipulate and have
dotted leaves. RAUH: Vochysia crassifolia (AM).
CLUSIACEAE, however producing a sticky MASSART: Erisma uncinatum (AM).
exudate. MANGENOT: Qualea dinizii, Ruizterranea spp. (AM).
References 453
Fig. 28.16 VOCHYSIACEAE. a Young tree with monopodial rhythmic branching and orthotropic branches
(Vochysia); b cauline glands (Qualea); c–d leaves with submarginal veins (sv), the secondary veins not numerous
(Vochysia spp.); e submarginal vein and numerous V II (Qualea) f base of a lateral stem with scale-leaves (Callisthene,
Southern Brazil); g stems orthotropic, then becoming plagiotropic, hence branching sympodial, sapling of Ruizterania
ferruginea, Gran Sabana, Venezuela
! The name of this family seems to be based on Ancient Greek, but in fact the name originates from
‘Voki’, or ‘Wachi wachi’ [8], the vernacular name of some Guianan species. Hence Auguste de Saint-
Hilaire should have named the family ‘Voquisiaceae’ or, if a German-speaking botanist got ahead of
him, ‘Wokisiaceae’ or ‘Wockisiaceae’ would have been more logical, but at that time, ‘Vochysiaceae’
sounded better.
References
1. Hutchinson, J. 1973. The families of flowering plants, arranged according to a new system based on their probable
phylogeny, 3rd ed, 968 pp. Oxford University Press.
2. Schnee, L. 1984. Plantas comunes de Venezuela. Colección ciencias biológicas VIII, 822 pp. Universidad Central de
Venezuela.
3. Soukup, J. 1970. Vocabulario de los nombres vulgares de la flora peruana, 384 pp. Lima: Colegio Salesiano.
4. Hickey, L.J. 1973. Classification of the architecture of Dicotyledonous leaves. American Journal of Botany 60:
17–33.
5. Bugnon, F. 1953. Recherches sur la ramification des Ampelidacées. Publications de l’Université de Dijon 11.
Presses Universitaires de France.
6. Gerrath, J.M., U. Posluszny, S.M. Ickert-Bond, and J. Wen. 2017. Inflorescence morphology and development in the
basal rosid lineage. Journal of Systematics and Evolution 55: 542–558.
7. Oldeman, R.A.A., and J.M. Fundter. 1986. On Diptero-Vochysian tree strategies. Naturalia Monspeliensia—
Colloque international de l’Arbre (Montpellier 9–14 septembre 1986), 191–208.
8. Den Outer, R.W. 2001. Vernacular names of Surinam woody plants, 356 pp. Wageningen University, The
Netherlands.
Zygophyllaceae s.l.
(incl. Balanitaceae) 29
29.1 Zygophyllaceae (incl. Balanitaceae): 48–50
27 genera and 300 species. Order Zygophyllales. Wood very dense and hard, often containing resins
or essential oils (Bulnesia, Guaiacum). In Larrea (AM subtropical), leaves are sticky with a strong
resin smell. Short twigs modified into spines (Balanites). Sympodial and modular architecture
(Fig. 29.2). Opposite leaf arrangement. Stipules present, but very small. Leaves compound pinnate
(Fig. 29.2), (Bulnesia, Guaiacum), bifoliolate for Balanites and Larrea spp. Leaf margin entire
(leaves notched in Larrea). Flowers showy, yellow (Bulnesia) or blue (Guaiacum), with a radial
symmetry. Ovary 2-5-locular. Fruits capsules (Fig. 29.3 Guaiacum), schizocarps (Tribulus), rarely
drupes (Balanites, winged in Bulnesia).
Trees (Fig. 29.1 Bulnesia, Guaiacum, AM; Balanites, AF), shrubs (Guaiacum, Fig. 29.2 Larrea,
AM), subshrubs or herbs (Kallstroemia, AM-N; Fig. 29.4 Tribulus, mainly AF).
Similar families:
Leguminosae-Caesalpinioids, but these with
leaves alternate and petioles pulvinate at base.
Architectural models badly known:

TROLL: Guaiacum (AM-S).
TROLL (erect form of the model?): Larrea
(subtropical N and S AM).
Fig. 29.1 Bulnesia arborea, flowering tree, Northern

Venezuela
https://doi.org/10.1007/978-3-031-05942-1_29
456 29 Zygophyllaceae s.l. (incl. Balanitaceae)
Zygophyllaceae grow in subtropical steppes and savannas, in America (Bulnesia, Guaiacum) and
Africa (Balanites). Bulnesia arborea (‘Arbol de vera’) is used in carpentry to make bowling balls,
barrels, toys (Fig. 29.5), etc. The wood of Guaiacum officinale and G. sanctum (lignum vitae), is
extremely strong and behaves as a natural lubricant, also used for traditional shipbuilding and
chandlery. Essential oils produced by distillation of the wood of Guaiacum officinale or Bulnesia
sarmientoi (‘palo santo’, Paraguay), creosote extracted from Larrea tridentata (Southern USA) used
as wood preservative.
Fig. 29.2 Guaiacum coulteri, leaves compound pinnate, BG LC, France. Guaiacum officinale, sympodial branch, each
module consisting of a single growth unit, x: abortion of an apical meristem. Larrea cuneifolia, fan-shaped branches
adapted to minimize evapotranspiration, Mendoza, Argentina
Fig. 29.3 Guaiacum officinale, open capsule with red seed, Venezuela. Guaiacum officinale, sympodial and modular
branches, BG Sirikit, Thailand
Similar families: LEGUMINOSAE, but Architectural models:

with petioles pulvinate at their base. TROLL: Guaiacum officinale, Larrea spp. (AM-S)
! Larrea is a subtropical genus, present in two widely separate areas, one in southern North
America, the other in Argentina, along the Andean range. As an adaptation to the arid and hot climate
of this region, stems of Larrea cuneifolia (‘jarilla’) are disposed in fans (Fig. 29.2), their edges facing
the direction of the zonda, a warm west wind.
29.1 Zygophyllaceae (incl. Balanitaceae): 48–50 457
Fig. 29.4 Tribulus terrestris, prostrate herb with a sympodial and modular architecture (branching looking
monopodial), Bali (widely naturalized)
Fig. 29.5 Dominos made of ‘arbol de vera’ wood (Bulnesia arborea)

Herbaceous Versus Arborescent
Habit in the Families and Orders 30
of Flowering Plants—Evolutionary
Trends
30.1 A Diagrammatic Representation of Angiosperm Evolution
This chapter is aimed to go one step further in the description of the families, it is based on
evolutionary trends (i.e. to the arborescent or to the herbaceous state). With this end in view, habit is
mapped onto family phylogeny and is shown here in a very simplified linear manner by coloured bars
(Fig. 30.1). Trees and woody lianas are represented in brown, shrubs or subshrubs in yellow, and
“herbs” (i.e. small rhizomatous or rosette plants in addition to annual or biennial herbs) in green. The
extent of a given colour in a bar represents the relative proportion of that biotype in the family and
arrays of colours represent the evolutionary trends (see Figs. 30.2, 30.3, 30.4, 30.5, 30.6, 30.7, 30.8,
30.9 and 30.10).
Conversion of a cladogram into a simplified representation of evolutionary trends

Figure 30.1 shows how the traditional cladogram is represented in a simplified linear manner by a
coloured bar. For example in the cladogram of the Apocynaceae [1], the Aspidospermeae (Aspi-
dosperma, etc.) show basal traits (i.e. the arborescent habit), whereas the Asclepiadoideae (Ascle-
piadeae, Marsdenieae, etc.) show derived traits (i.e. the herbaceous habit). «Traits» are emphasized
here, and «basal», or «derived», should refer to traits. In contrast to Aspidospermeae, Dyera (tribe
Melodineae) shows a derived arborescent trait embedded in the cladogram within shrubby-
herbaceous traits, this situation is referred to as a “reversal”.
If one considers the right-hand end of each bar as the “head” and the left-hand end as the “tail”,
then a brown bar with a small green head (right) represents a group that is mainly arborescent with
some derived herbaceous taxa, whereas a brown bar with a small yellow tail (left) represents an
arborescent group where shrubs are basal (strictly speaking, in phylogenetic trees, traits, not clades,
can be ‘basal’ [2]). Evolution goes from left to right. Yellow/green following brown is for evolution
from trees to shrubs/herbs. Brown following yellow/green represents the reverse process.
Simplified representations (i.e. coloured bars) are given for the families treated in this book and
also for some other families. Numbers in brackets [R1], [R2], etc., are for references concerning
phylogenies (see Sect. 30.2). Empty bars are where data is missing.
https://doi.org/10.1007/978-3-031-05942-1_30
460 30 Herbaceous Versus Arborescent Habit in the Families …
Fig. 30.1 A simplified representation of phylogenies
AUSTROBAILEYALES - Myristicaceae
- Schisandraceae - Annonaceae
CANELLALES LAURALES
PIPERALES - Monimiaceae
- Piperaceae - Lauraceae
- Aristolochiaceae - Hernandiaceae
MAGNOLIALES CHLORANTHALES
- Magnoliaceae - Chloranthaceae [R1]
Fig. 30.2 Basal angiosperms

30.1 A Diagrammatic Representation of Angiosperm Evolution 461
MONOCOTS [R2] POALES

ALISMATALES - Poaceae [R3]
DIOSCOREALES LILIALES
- Dioscoreaceae - Smilacaceae
PANDANALES ASPARAGALES [R4]
- Pandanaceae - Asparagaceae
- Cyclanthaceae
Fig. 30.3 Monocots
BUXALES RANUNCULALES [R5]

- Buxaceae - Eupteleaceae
PROTEALES - Menispermaceae
- Sabiaceae - Berberidaceae [R6]
- Proteaceae - Ranunculaceae [R7]
Fig. 30.4 Buxales–Proteales–Ranunculales
SAXIFRAGALES [R8] VITALES

- Hamamelidaceae - Vitaceae [R9]
- Altingiaceae DILLENIALES
- Grossulariaceae - Dilleniaceae
- Saxifragaceae
Fig. 30.5 Dilleniales–Saxifragales–Vitales
CUCURBITAL. [R10] ROSALES [R13]

- Anisophylleaceae - Rosaceae [R14]
- Tetramelaceae - Rhamnaceae [R15]
- Begoniaceae - Ulmaceae
FABALES - Cannabaceae [R16]
- Leguminosae [R11] - Moraceae [R17]
- Surianaceae - Juglandaceae
- Polygalaceae [R12] - Urticaceae [R18]
FAGALES ZYGOPHYLLALES
- Fagaceae - Zygophyllaceae
- Myricaceae
Fig. 30.6 Cucurbitales–Fabales–Fagales–Rosales–Zygophyllales

CELASTRALES - Euphorbiaceae [R23]

- Celastraceae [R19] - Phyllanthaceae [R24]
MALPIGHIAL. [R20] - Hypericaceae [R25]
- Lacistemataceae - Rhizophoraceae
- Salicaceae - Erythroxylaceae
- Achariaceae - Humiraceae
- Violaceae [R21] - Linaceae [R26]
- Passifloraceae - Ixonanthaceae
- Irvingiaceae - Malpighiaceae
- Pandaceae OXALIDALES [R27]
- Picrodendraceae - Huaceae
- Ochnaceae [R22] - Oxalidaceae
- Bonnetiaceae - Connaraceae
- Clusiaceae - Cunoniaceae
- Hypericaceae - Elaeocarpaceae
- Rhizophoraceae - Brunelliaceae
- Putranjivaceae
Fig. 30.7 Celastrales–Malpighiales–Oxalidales
BRASSICALES [R28] - Lythraceae [R33]

- Brassicaceae - Vochysiaceae
- Moringaceae - Myrtaceae
- Caricaceae - Crypteroniaceae
- Capparaceae - Melastomataceae
MALVALES [R29] PICRAMNIALES
- Bixaceae [R30] - Picramniaceae
- Malvaceae SAPINDALES [R34]
- Muntingiaceae - Sapindaceae
- Thymelaeaceae [R31] - Burseraceae
- Dipterocarpaceae - Anacardiaceae
- Sarcolaenaceae - Simaroubaceae
MYRTALES [R32] - Meliaceae
- Combretaceae - Rutaceae
Fig. 30.8 Brassicales–Malvales–Myrtales–Picramniales–Sapindales

30.1 A Diagrammatic Representation of Angiosperm Evolution 463
CARYOPHYLLAL. [R35] SANTALALES [R38]

- Rhabodendraceae - Erythropalaceae
- Ancistrocladaceae - Olacaceae
- Polygonaceae [R36] - Opiliaceae
- Physenaceae - Santalaceae [R39]
- Cactaceae - Loranthaceae
- Phytolaccaceae
- Nyctaginaceae [R37]
Fig. 30.9 Caryophyllales–Santalales
CORNALES [R40] - Gelsemiaceae

- Cornaceae - Apocynaceae [R46]
- Nyssaceae BORAGINALES
- Loasaceae - Boraginaceae [R47]
ERICALES [R41] LAMIALES [R48]
- Marcgraviaceae - Oleaceae
- Lecythidaceae - Acanthaceae [R49]
- Theaceae - Verbenaceae
- Pentaphylacaceae - Bignoniaceae
- Ebenaceae - Lamiaceae [R50]
- Symplocaceae SOLANALES
- Primulaceae [R42] - Solanaceae [R51]
- Styracaceae - Convolvulaceae [R52]
- Sapotaceae APIALES [R53]
- Actinidiaceae - Torricelliaceae
- Clethraceae - Pittosporaceae
- Cyrillaceae - Araliaceae
- Ericaceae [R43] - Apiaceae
ICACINALES AQUIFOLIALES
- Icacinaceae s.l. - Aquifoliaceae
GENTIANALES [R44] ASTERALES
- Rubiaceae - Goodeniaceae
- Loganiaceae [R45] - Asteraceae [R54]
Fig. 30.10 Apiales–Aquifoliales–Asterales–Boraginales–Cornales–Ericales–Gentianales–Icacinales–Lamiales–

Solanales
30.2 References to Phylogenetic Studies
[R1] » Chloranthaceae: Ascarina, shrubby or arborescent, Chloranthus japonicus herbaceous and

derived [3].
[R2] » Monocots: Alismatales herbaceous and basal [4].
[R3] » Poaceae: Bambusoideae basal and mainly woody [5].
[R4] » Asparagales: mainly herbaceous, some “reversals”, Cordyline and Agave arborescent [4].
[R5] » Ranunculales: Eupteleaceae arborescent and basal [5].
[R6] » Berberidaceae: Nandina arborescent and basal, Epimedium herbaceous and derived [6].
[R7] » Ranunculaceae: Xanthorhiza woody and basal Pulsatilla, Ranunculus, herbaceous and derived [7].
[R8] » Saxifragales: Daphniphylllum, Hamamelis arborescent and basal, Saxifraga, herbaceous and
derived [8, 9].
[R9] » Vitaceae: almost all woody, Leea basal, arborescent to herbaceous [10, 11].
[R10] » Cucurbitales: Anisophylleaceae arborescent and basal, Cucurbitaceae herbaceous and
derived [12].
[R11] » Leguminosae: Caesalpinioids arborescent and basal, Papilionoideae trees, shrubs and many
herbaceous derived [5].
[R12] » Polygalaceae: Xanthophyllum arborescent and basal, Monnina, Polygala shrubby-
herbaceous and derived [13].
[R13] » Rosales: several «reversals», Rosaceae basal, Moraceae derived [14].
[R14] » Rosaceae: Dryadoideae shrubs and basal, Alchemilla, Sanguisorba herbaceous and derived,
possible «reversals» from herbs (Acaena) to trees (Polylepis) [15].
[R15] » Rhamnaceae: mainly arborescent, small shrubs derived (e.g. Spyridium) [16].
[R16] » Cannabaceae: Trees basal, herb Cannabis derived [17].
[R17] » Moraceae: almost all trees, Dorstenia herbaceous and derived, Castilla arborescent, derived
[18].
[R18] » Urticaceae: Cecropia, trees basal, Pouzolzia herbaceous and derived, Broussonetia, trees
derived, many «reversals» [19].
[R19] » Celastraceae: Almost all arborescent or shrubby, Parnassia herbaceous and derived, [20].
[R20] » Malpighiales: many «reversals», Euphorbiaceae derived [21, 22].
[R21] » Violaceae: several «reversals» (e.g. arborescent Melicytus), Rinorea arborescent-shrubby
and basal, Hybanthus not monophyletic, shrubby or herbaceous, derived [23].
[R22] » Ochnaceae: almost all arborescent, Sauvagesia shrubby or woody herbaceous, derived [24].
[R23] » Euphorbiaceae + Peraceae: Peraceae (Chaetocarpus, Pera), Suregada, arborescent and
basal, Euphorbia spp. herbaceous and derived, but many «reversals» [25, 26].
[R24] » Phyllanthaceae: many trees, some shrubs and herbs (Phyllanthus spp.) [27].
[R25] » Hypericaceae: Vismia arborescent and basal, Hypericum herbaceous and derived [28].
[R26] » Linaceae: Roucheria arborescent and basal, Linum herbaceous and derived [29, 30].
[R27] » Oxalidales: mainly arborescent, herbaceous Oxalis derived [31].
[R28] » Brassicales: Carica, Moringa arborescent and basal, Brassica herbaceous and derived [32, 33].
[R29] » Malvales: basally arborescent (Theobroma, etc.), Gossypium, Malva, shrubby or herbaceous
and derived [34].
[R30] » Bixaceae: trees basal (Cochlospermum), herbs derived (Amoreuxia) [35].
[R31] » Thymelaeaceae: trees basal (Gonystylus, etc.), shrubs derived (Daphne, Wikstroemia, etc.)
[36].
[R32] » Myrtales: Combretaceae arborescent and basal, herbaceous Melastomataceae derived [37].
30.3 Discussion and Hypothesis 465
[R33] » Lythraceae: Duabanga, Sonneratia arborescent and basal, Cuphea herbaceous and derived
[38].
[R34] » Sapindales: almost all arborescent but Nitraria, Biebersteinia shrubby-herbaceous basal,
Ruta herbaceous and derived [39].
[R35] » Caryophyllales: Rhabdodendraceae arborescent and basal, many «reversals» [40, 41].
[R36] » Polygonaceae: derived from herbaceous Plumbaginaceae, Symmiria shrubby and basal,
Ruprechtia arborescent and derived, many herbs, several «reversals» [42].
[R37] » Nyctaginaceae: Pisonia arborescent and basal, Mirabilis herbaceous and derived [43].
[R38] » Santalales: Erythropalum, Strombosia arborescent and basal, Viscum, Phoradendron
herbaceous and derived [44], parasitic Balanophora derived [45].
[R39] » Santalaceae: many «reversals», parasitic shrubs (Comandra), trees (Exocarpos, Santalum),
herbs (Thesium) [46].
[R40] » Cornales: Cornus, Alangium arborescent and basal, Loasa herbaceous and derived [47].
[R41] » Ericales: Impatiens, Pyrola, Primula herbaceous and derived [48].
[R42] » Primulaceae: Deherainia arborescent and basal, Anagallis herbaceous and derived [49].
[R43] » Ericaceae: mainly arborescent or shrubby, several «reversals», Pyrola herbaceous and
derived [50].
[R44] » Gentianales: mainly woody, Cinchonoideae, arborescent and basal, but Rubioideae herba-
ceous and basal, Asclepiadoideae herbaceous and derived, frequent «reversals» [51].
[R45] » Loganiaceae: Mitrasacme shrubby-herbaceous and derived [52].
[R46] » Apocynaceae: Aspidosperma arborescent and basal, Vinca herbaceous and derived [1].
[R47] » Boraginaceae: Ehretia arborescent and basal, Heliotropium, Tournefortia herbaceous and
derived [53].
[R48] » Lamiales: Oleaceae arborescent and basal, but families of core Lamiales with a complicated
phylogeny [54], frequent «reversals».
[R49] » Acanthaceae: Nelsonia herbaceous and basal, Avicennia arborescent and somewhat basal,
Bravaisia in derived Ruellieae which are mostly herbaceous [55].
[R50] » Lamiaceae: arborescent basal, (Tectona, Gmelina), herbs derived (Salvia) [56].
[R51] » Solanaceae: Schizanthus herbaceous and basal, Solanum, Cestrum, Goetzea shrubby and
derived, several «reversals» [57].
[R52] » Convolvulaceae: Humbertia, large trees, basal, Ipomoeeae, herbaceous lianas derived [58].
[R53] » Apiales: Aralidium arborescent and basal, herbaceous Apiaceae derived [59].
[R54] » Asteraceae: shrubby and arborescent Barnadesioideae sister group of all other Asteraceae [60].
30.3 Discussion and Hypothesis
Figures 30.11, 30.12, 30.13 and 30.14 represent a summary of the different patterns of evolution:
Frequencies of the different patterns of evolution
Fig. 30.11 Type 1: from trees to trees (e.g. Ebenaceae, Salicaceae, Sapotaceae, Vochysiaceae, etc.)
Fig. 30.12 Type 2: from trees to herbs (e.g. Apiales, Leguminosae, Malvales, Myrtales, Polygonaceae, Primulaceae,
etc.)
Fig. 30.13 Type 3: from herbs to trees, only a few examples (e.g. basal Angiosperms, some special cases like
Sanguisorba to Polylepis in Rosaceae [61], s.t. derived arborescent taxa in tropical highlands or islands: Echium,
Senecio, etc.)
Fig. 30.14 Type 4: from herbs to shrubs and trees, with several “reversals” (e.g. Chloranthaceae, Ranunculales
without Eupteleaceae, Gentianales)
Type 1: Evolution starts from trees and continues as trees (bar entirely brown). This occurs frequently
and one of the best examples is the Sapindales, an order without true herbs (i.e. annual or biennial
herbs).
Type 2: Evolution starts from trees and leads to herbs. (brown tail and green head). A very frequent
case, see also [62] and one of the best examples is Malvales, a group with a well-balanced mixture of
trees, shrubs and herbs. Some authors have suggested that past climatic fluctuations causing
expansion and contraction of rainforest could also have offered ecological opportunities for small
shrubs and herbs and contributed to the process of divergent evolution [63]. Obviously, shrubs
represent links between trees and herbs. In the tropics, savannas, campo cerrados, and miombo
woodlands host many shrubs, often with subterranean stems or geoxylic Kalahari Bassin, Sudano-
Guinean region [64].
Type 3: Evolution starts from herbs and leads to trees (green tail and brown head). This is an
uncommon pattern but is exemplified by the arborescent Dendrosenecio which has evolved from the
smaller Senecioneae [65], the giant Lobelia which has developed from herbaceous ones [66] and the
arborescent Phytolacca dioica coming from herbaceous representatives of the genus. On the evolu-
tionary scale, arborescent Monocotyledons (Arecaceae, Pandanaceae) provide striking examples of
groups which have originated from the herbaceous Alismatales (see the Angiosperm Phylogeny
Groups website).
Type 4: Evolution starts with herbs or shrubs and develops lineages with shrubs or trees. This is quite
rare but examples exist in the Monocotyledons and the Chloranthaceae.
30.3 Discussion and Hypothesis 467
The patterns illustrated in Figs. 30.2, 30.3, 30.4, 30.5, 30.6, 30.7, 30.8, 30.9 and 30.10, for
different orders of flowering plants and summarized in Figs. 30.11, 30.12, 30.13 and 30.14 show that
evolution from trees (Types 1 and 2) appears to be much more common than evolution from herbs
(Types 3 and 4). In addition, within the pattern described in Type 2, radiation can be profuse leading
to many herbaceous taxa (e.g. Papilionoideae), or very limited and leading to isolated taxa, for
example Cassytha filiformis (Lauraceae), or Parnassia palustris (Celastraceae). As this field guide
concerns principally trees, it is evident that entirely brown bars are overrepresented in Figs. 30.2,
30.3, 30.4, 30.5, 30.6, 30.7, 30.8, 30.9 and 30.10. However, when considering the groups where
herbs are relatively frequent, there is a same asymmetry in the distribution: the herbaceous habit
appears more often as derived and rarely basal. Even for strongly herbaceous families (e.g.
Caryophyllaceae, Plantaginaceae, Ranunculaceae), the few existing arborescent taxa seem to be rarely
derived. The following discussion will attempt to explain why these divergent evolutionary patterns
appear to be unevenly distributed within the angiosperms.
Evolution rates
How fast does evolution proceed? Plants with a short generation time, (i.e. the time from germination
to production of a seed which germinates) generally show more rapid rates of molecular evolution.
Longer-lived trees and shrubs, in contrast, evolve more slowly and show less variability in their rates
of evolution, [67, 68] and the same trend holds for tall plants with respect to smaller ones [69]. In
addition, age at maturity appears to be negatively correlated with the rate of diversification [70]. It is
interesting to note that this difference in the rate of evolution between herbs and woody plants has
been maintained throughout time [68].
However, evolution rates alone do not explain why a process starting from trees and leading to
herbs seems much more frequent than the reverse process.
Missing links in extant plants
Returning to Figs. 30.2, 30.3, 30.4, 30.5, 30.6, 30.7, 30.8, 30.9 and 30.10 which illustrates the
patterns of evolution in the major groups, it is apparent that several orders are quite isolated and seem
to appear from nowhere. Incomplete lineage sorting (the consequence of extinction of several
arborescent taxa), could be the explanation for the missing links in the classification of extant plants
(it is not obvious to see a connection between Monocotyledons and ‘Dicotyledons’ with respect to
morphology). Similarly, one must admit that there are significant gaps surrounding the orders
Ranunculales, Proteales, Dilleniales, Caryophyllales, Vitales, Cornales, Icacinales, Gentianales,
Asterales, and their relatives. This is not an absolute rule, as in certain cases the gap is less marked.
For example, the Connaraceae form a bridge between the Oxalidales-Fabales and the Gentianales-
Solanales-Lamiales have certain features in common (gamopetalous flowers, flower parts in four or
five and simple leaves). Isolated groups can also be found within the same order: the Rhizophoraceae
do not resemble any other family in the Malpighiales, and the Thymelaeaceae are quite atypical in the
order Malvales.
Revisiting Dollo’s law
Dollo’s law states that, following loss, a complex trait cannot re-evolve in an identical manner [71]. In
this case, the “complex trait” is the arborescent habit and the “less complex trait” is the herbaceous
habit. It is obvious that trees are, in their vegetative parts, more complex organisms than herbs. For
example, the formation of bark relies on a particular form of cambium, the phellogen. Phellogen
produces phellem and phelloderm, which are tissues characteristic of the complex structure of bark.
Trees are also more complex in other ways, such as the specific rhythms of growth for each kind of
axis (trunk, branch, twig) as well as the elaborate tropisms and the specific reiteration processes (e.g.
delayed or sequential). In contrast, herbs have no bark and their growth rhythms, tropisms and
reiteration processes are much less diversified. Consequently, for a tree to evolve from a herb the
complex traits must be re-invented, relying on transference of function [72], secondary woodiness
[73, 74] or a resurgence of similar traits existing in pre-existing groups (see Chap. 3 - Preliminary
remarks to architectural models). Hence a new woody lineage is unlikely to resemble an ancient one.
For this reason, the bars in Types 3 and 4 have been drawn farther apart than in Types 1 and 2.
In contrast, in the majority of cases (i.e. the evolution from trees to herbs), the ancient processes
would have been relatively well preserved and this could explain why profusely diversifying groups
such as the Asteraceae, have evolved to give a huge number of species which, vegetatively, closely
resemble each other. In fact most Asteraceae represent many variations on the same theme and do not
show any major vegetative differences, despite considerable diversification.
The possible herbaceous origin of trees
It is now generally accepted that angiosperms have a herbaceous origin [75–77]. This hypothesis was
proposed by Agnes Arber [78] already a century ago. Faster evolution in herbs and small plants
probably resulted in divergence i.e. radiation followed by new orders. An expansion phase would
then have occurred among the Paleoherbs. For example, Montsechia vidalii is a herbaceous aquatic
plant which resembles the oldest known fossil angiosperm, the Ceratophyllum species [64].
If the first angiosperms were herbs, and perhaps aquatic ones, their remains would be much less
likely to be fossilized and preserved than those of shrubs or trees. Fossils of Paleoherbs are already
extremely rare compared to those of ligneous plants and this also applies to herbs of more recent
origin. As Crepet & Feldman [79] argue, ‘the paucity of fossil grasses (Poaceae and Cyperaceae) is
particularly vexing, as these are some of the largest families of flowering plants’. So perhaps this
fossil paucity may also exist for groups of more recent origin than the paleoherbs. One could imagine
that history has been repeated, and on many occasions, an arborescent group has evolved from a
herbaceous ancestor which has disappeared leaving no fossil evidence.
Perhaps the best examples are certain Monocotyledons which, evolving from herbaceous
Alismatales-like ancestors [80] have “reinvented” trees of a very different kind from those of the
‘Dicotyledons’. In Monocotyledons, the cambium, when it exists, originates from a primary thick-
ening of the meristem. Hence, stem thickening does not result from a vascular cambium as is the case
for ‘Dicotyledons’ and Gymnosperms [81]. Nymphaeales-Nelumbonales-Proteales could well pro-
vide another example if basal taxa in these orders were also herbaceous and older than Platanaceae
and Proteaceae. Species of Nymphaea and Nelumbo are aquatic herbs of recent origin and their
ancestors might be also herbaceous. Actually, Nymphaeaceae is considered to have appeared slightly
earlier than Platanaceae [82] and this argues for a derived arborescent habit in the above mentioned
orders.
The consequence of higher mutation rates in herbs
Assuming that evolution rates are higher for herbs than for trees, it is not surprising that, originating
from herbs (Types 3 and 4) arborescent lineages will diverge more than for Type 1. This is in
accordance with Dollo’s law. In summary, the fast-evolving herbaceous ancestors have become
extinct and have led to well-separated woody groups. Consequently this would provide an expla-
nation for the asymmetrical distribution of the frequencies, i.e. the relative abundance of brown bars
with a green head, but without a green tail. In other words, one could say that radiation from trees to
herbs is a cohesive process because, as in the Asteraceae, the many derived taxa are kept in the
family. In contrast, evolution from herbs to trees would appear to be a more divergent process
because the taxa with basal herbaceous traits have vanished, as is the case for the emergence of
Monocotyledons from basal angiosperms.
References 469
Final words
As explained in the introduction, this field guide concentrates mainly on trees and large lianas.
However, herbaceous plants also merit some attention, particularly in view of the strange asymmetry
observed in the evolutionary patterns of angiosperm families.
Tropics are a prolific source of diversification where trees dominate herbs, whereas higher latitudes
are the realm of herbs. If the tendency of evolution was to produce small plants, all the vegetation on
Earth would consist of herbs. This is obviously not the case, especially in the tropics, therefore some
large plants may have evolved from smaller ones and missing links would be explained, at least in
part, by the scarcity of fossil herbs. We are aware that the argumentation is weak, because it is based
upon negative evidence. On the other hand, our hypothesis is supported by the impressive gaps
between orders, the relatively high mutation rates in herbs, and by the principles of Dollo’s Law,
which would have provided the mechanisms for this “inverted” evolution.
Evolution from herbs to trees and trees to herbs represent two diametrically opposite processes.
However, the interest in these two processes has been inversely proportional to the frequency of their
occurrence. In the case of tree-like herbs, Western botanists have been attracted to the exotic flora of
high tropical mountains and were fascinated by the phenomenon of pachycauly [66, 83–88], etc.
Depending on taxa and methodology, the ancestors of giant Lobelias, Frailejones or arborescent
Ragworts might be a pachycaul for some authors or a rosette plant for others, “true trees” being
discarded.
The conjecture exposed above is based on the evolution from small plants to trees, a process often
accepted for pachycauls in tropical highlands. However this is much less common than the reverse
process and it is paradoxical that botanists have been relatively indifferent to the much more frequent
examples of herbaceous plants evolving from trees!
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List of Identified Families
Families identified by the keys (at least partly)

and listed as in the APG IV system of classifi-
cation
BASAL ANGIOSPERMS Monocotyledons

Alismatales
Austrobaileyales
Araceae
Schisandraceae
Trimeniaceae
Arecales
Arecaceae
MESANGIOSPERMS
Magnoliidae Asparagales
Asparagaceae
Canellales
Canellaceae
Dioscoreales
Winteraceae
Dioscoreaceae
Laurales
Liliales
Hernandiaceae
Ripogonaceae
Lauraceae
Smilacaceae
Monimiaceae
Siparunaceae
Pandanales
Pandanaceae
Magnoliales
Annonaceae
Poales
Magnoliaceae
Poaceae-Bambusoideae
Myristicaceae
EUDICOTYLEDONS
Piperales
Aristolochiaceae Buxales
Piperaceae Buxaceae
Chloranthales Proteales
Chloranthaceae Proteaceae
Sabiaceae
© The Editor(s) (if applicable) and The Author(s), under exclusive license 473
to Springer Nature Switzerland AG 2023
https://doi.org/10.1007/978-3-031-05942-1
474 List of Identified Families
Ranunculales Zygophyllales
Menispermaceae Zygophyllaceae
Berberidaceae
Ranunculaceae (Clematis) Celastrales
Celastraceae
Dilleniales
Dilleniaceae Malpighiales
Achariaceae
Superrosidae Bonnetiaceae
Calophyllaceae
Saxifragales
Caryocaraceae
Daphniphyllaceae Centroplacaceae
Hamamelidaceae Chrysobalanaceae
Medusandraceae Clusiaceae
Ctenolophonaceae
Rosidae Dichapetalaceae
Vitales Erythroxylaceae
Vitaceae Euphorbiaceae
Euphroniaceae
Fabidae Goupiaceae
Humiriaceae
Cucurbitales Hypericaceae
Anisophylleaceae Irvingiaceae
Corynocarpaceae Ixonanthaceae
Tetramelaceae Linaceae
Malpighiaceae
Fabales Ochnaceae
Leguminosae Pandaceae
Polygalaceae Passifloraceae
Surianaceae Peraceae
Phyllanthaceae
Fagales Picrodendraceae
Casuarinaceae Putranjivaceae
Fagaceae Rhizophoraceae
Juglandaceae Salicaceae
Myricaceae Lacistemataceae
Trigoniaceae
Rosales Violaceae
Elaeagnaceae
Moraceae Oxalidales
Rhamnaceae Brunelliaceae
Rosaceae Connaraceae
Ulmaceae Cunoniaceae
Cannabaceae Elaeocarpaceae
Urticaceae Huaceae
Oxalidaceae
List of Identified Families 475
Malvidae Superasteridae
Brassicales
Caryophyllales
Capparaceae Ancistrocladaceae
Caricaceae Cactaceae
Moringaceae Dioncophyllaceae
Nyctaginaceae
Crossosomatales Physenaceae
Staphyleaceae (Turpinia) Phytolaccaceae
Polygonaceae
Geraniales Rhabdodendraceae
Melianthaceae
Santalales
Huerteales Erythropalaceae
Tapisciaceae (Huertea) Loranthaceae
Olacaceae
Malvales Schoepfiaceae
Bixaceae Opiliaceae
Dipterocarpaceae Santalaceae
Malvaceae
Muntingiaceae Asteridae
Sarcolaenaceae
Thymeleaceae Cornales
Cornaceae
Myrtales Hydrangeaceae (Hydrangea)
Combretaceae Nyssaceae
Crypteroniaceae
Lythraceae Ericales
Melastomataceae Actinidiaceae
Myrtaceae Clethraceae
Vochysiaceae Cyrillaceae
Ebenaceae
Picramniales Ericaceae
Picramniaceae Lecythidaceae
Marcgraviaceae
Sapindales Pentaphylacaceae
Anacardiaceae Primulaceae
Burseraceae Sapotaceae
Meliaceae Styracaceae
Rutaceae Symplocaceae
Sapindaceae Tetrameristaceae
Simaroubaceae Theaceae
476 List of Identified Families
Lamiidae Bignoniaceae
Lamiaceae
Icacinales
Oleaceae
Icacinaceae
Scrophulariaceae
Verbenaceae
Metteniusales
Metteniusaceae
Campanulidae
Gentianales Apiales
Apocynaceae Araliaceae
Gentianaceae Pittosporaceae
Loganiaceae Torricelliaceae
Gelsemiaceae
Rubiaceae Aquifoliales
Aquifoliaceae
Boraginales Cardiopteridaceae
Boraginaceae Stemonuraceae
Solanales Asterales
Convolvulaceae Goodeniaceae
Montiniaceae Asteraceae
Solanaceae
Dipsacales
Lamiales Viburnaceae
Acanthaceae
Index
A Aglaia, 108, 110, 111, 324, 326

Abroma, 313 Agonandra, 55, 355, 356
Abrus, 115, 297, 298 Agrostistachys, 59, 241, 243
Abuta, 60, 328, 329 Ailanthus, 109, 110, 424, 425
Abutilon, 313–315 Alangium, 54, 215, 216
Acacia, 8, 32, 33, 71, 73, 113, 116, 138, 291–294, 296, Albizia, 117, 293, 294
383 Alcea, 314
Acaena, 396 Alchornea, 58, 64, 97, 98, 99, 241–243
Acalypha, 60, 98, 99, 241, 242 Aleurites, 101, 241–243
ACANTHACEAE, 10, 34, 77–79, 121, 132, 142, 476 Alexa, 90, 115, 297, 298
Acanthus, 141, 142 Allamanda, 155, 156
Acer, 80, 92, 106, 107, 109, 416 Allanblackia, 42, 202
Aceratium, 76, 231, 232 Allophylus, 90, 91, 109, 416, 417
Acharia, 143–145 Aloysia, 65, 445
ACHARIACEAE, 31–33, 56–59, 61, 62, 68, 143, 145, Alphitonia, 47, 390, 391
411, 474 Alseis, 402
Acoelorrhaphe, 163 Alsodeiopsis, 270
Acosmium, 115 Alstonia, 43, 154–156
Acridocarpus, 58, 125, 310, 311 Altingia, 70, 261, 262
Acrocomia, 167 Alvaradoa, 109, 424, 426
Acrotrema, 222 Amanoa, 95
Actinidia, 145, 146 Amasonia, 277, 278
ACTINIDIACEAE, 32, 33, 65, 66, 69, 145, 475 Amherstia, 113, 289
Actinodaphne, 66, 280, 281 Amicia, 116
Actinostemon, 102, 243 Amoreuxia, 39, 176
Adansonia, 106, 313–315 Ampelocera, 48, 439
Adelobotrys, 322 Amphilophium, 176
Adenanthera, 116, 117, 292 Amyris, 106
Adenia, 130, 361–363 ANACARDIACEAE, 7, 29, 32, 33, 35, 41, 43, 65, 66, 69,
Adenium, 155 84, 90, 106–110, 147–149, 475
Adenocalymma, 135, 173 Anacardium, 41, 69, 107, 147, 148
Adina, 404 Anacolosa, 127, 352
Adinandra, 52, 363, 364 Anadenanthera, 116, 291, 294
Aegiphila, 133, 277 Anaxagorea, 151, 152, 154
Aegle, 90, 406 Anchietea, 125, 447
Aeglopsis, 62 ANCISTROCLADACEAE, 121, 127, 149, 475
Aesculus, 92, 416 Ancistrocladus, 127, 149, 150
Afrobrunnichia, 127, 378 Ancistrophyllum, 138
Afrostyrax, 49, 264 Ancistrothyrsus, 127, 361–363
Afzelia, 287 Andira, 114, 297
Agapetes, 128, 234 Androsace, 382
Agarista, 234, 235 Aneulophus, 75, 239
Agathis, 38, 84, 162, 163 Angostura, 107, 403, 404
Agathosma, 406 Aniba, 9, 63, 66, 280–282, 297
Agelaea, 89, 90, 124, 136, 209–212 Anisophyllea, 53, 62, 150
© The Editor(s) (if applicable) and The Author(s), under exclusive license 477
to Springer Nature Switzerland AG 2023
https://doi.org/10.1007/978-3-031-05942-1
478 Index
ANISOPHYLLEACEAE, 31, 32, 53, 61, 62, 150, 461 Asclepias, 43, 155, 156
Anisoptera, 225, 228 ASPARAGACEAE, 29, 37, 38, 124, 473
Anneslea, 65, 363, 364 Asparagus, 124
Annona, 50, 151, 152, 154 Aspidosperma, 41–43, 154–156, 459
ANNONACEAE, 30, 50, 121, 124, 151, 262, 315, 337, Asplundia, 38, 138, 219, 220
460 ASTERACEAE, 32, 34, 35, 62–65, 79, 81, 121, 132, 168,
Anogeissus, 83, 206, 207 169, 171, 468, 476
Anopyxis, 76, 393 Astronium, 107–109, 147
Anthocleista, 79, 253–255 Asystasia, 141
Anthodiscus, 88 Atalantia, 62, 107, 403, 406
Anthonotha, 113 Ateleia, 114
Anthostema, 103 Atractogyne, 132, 398
Antiaris, 331–333 Atropa, 428
Antidaphne, 415 Attalea, 166, 167
Antidesma, 95, 96, 242, 365, 366 Atuna, 44, 200
Antigonon, 126, 378 Aucoumea, 107, 182
Antonia, 79, 303, 304 Aulax, 386
Apeiba, 44, 104, 313–315 Auranticarpa, 373
Aphanamixis, 110, 324 Austrobuxus, 370
Aphelandra, 141, 142 Averrhoa, 18, 89, 357, 358
Aphloia, 50 Avicennia, 78, 141, 142
APHLOIACEAE, 30, 49, 50 Azadirachta, 90, 109, 110, 324–326
APOCYNACEAE, 29, 41–43, 121, 123, 154, 155, 459,
476
Apodocephala, 168 B
Apodytes, 73, 269 Baccaurea, 57, 96, 365, 366
Aporosa, 46, 57, 96, 365, 366 Baccharis, 168–170
Aptandra, 351 Bactris, 38, 166, 167
AQUIFOLIACEAE, 8, 10, 31, 58, 59, 158, 476 Baeckea, 84
Aquilaria, 50, 435–437 Bagassa, 42, 331
ARACEAE, 29, 32, 37, 121, 219, 473 Baikiaea, 299
Arachis, 297 Baillonella, 41, 421, 423
Aralia, 138, 159 Balakata, 102
ARALIACEAE, 32, 33, 36, 60, 67, 87–90, 121, 138, 159, Balanites, 89, 455, 456
160, 162, 437, 576 Balfourodendron, 106
Aralidium, 68, 437 Baloghia, 100
Araucaria, 38, 162 Balthasaria, 363
ARAUCARIACEAE, 29, 35, 37, 38, 83, 84, 162, 163 Bambusa, 38
Arceuthobium, 414, 415 Banara, 46, 410–412
Archidendron, 293 Bandereia, 111, 130
Archytaea, 66, 202, 205 Banisteriopsis, 75, 132–134, 311
Ardisia, 51, 65, 380, 382, 383 Banksia, 384
Areca, 163, 166, 372 Baphia, 112, 297
ARECACEAE, 4, 17, 29, 37, 38, 121, 138, 163, 164, 167, Barleria, 141, 143
219, 360, 466, 473 Barnadesia, 169
Arenga, 163, 166, 167 Barringtonia, 57, 67, 70, 282–285
Argocoffeopsis, 398 Barteria, 46, 361–363
Argostemma, 400 Bauhinia, 45, 111, 112, 121, 124, 130, 137, 138,
Argylia, 173 286–289
Argyreia, 213 Beauprea, 386
Argyrodendron, 312 Beilschmiedia, 65, 84, 280
Aristolochia, 52, 129, 167, 168 Bejaria, 73, 234–236
ARISTOLOCHIACEAE, 31, 51, 52, 121, 129, 167, 473 Benstonea, 360
Aristotelia, 231, 232 BERBERIDACEAE, 36, 89, 473
Arrabidaea, 135, 173 Berberis, 89
Artabotrys, 124, 151 Berchemia, 76, 390, 391
Artocarpus, 40, 331–333 Berlinia, 113
Arytera, 110, 416, 417 Berrya, 105
Ascarina, 74, 197 Bersama, 87
Index 479
Bertholletia, 282–285 Buchholzia, 71, 187, 188

Bertiera, 402 Bucida, see Terminalia, 66
Bhesa, 57, 195, 196 Buddleja, 80
Biancaea, 288 Bulnesia, 91, 455–457
Bidens, 168 Bunchosia, 81, 311, 312
Bignonia, 173 Burasaia, 136, 328, 329
BIGNONIACEAE, 32, 35, 36, 62, 63, 81, 84, 90–92, 121, Burkea, 118, 286
134–136, 370, 476 Bursera, 108, 181–183
Bikkia, 402 BURSERACEAE, 29, 35, 41, 86, 87, 90, 106–108, 110,
Billardiera, 373 181, 475
Billia, 92 Burttia, 63, 209, 212
Biophytum, 357, 358 Bussea, 118
Bischofia, 88, 365, 368 Butea, 16, 114, 297
Bixa, 39, 56, 176–178 Butia, 167
BIXACEAE, 3, 29, 31, 36, 39, 56, 87, 88, 176, 178, 475 BUXACEAE, 35, 80, 84, 86, 184, 473
Blachia, 99, 101 Buxus, 80, 86, 184
Blakea, 322 Byrsonima, 75, 310–312
Blepharistemma, 76, 77, 393, 394 Byttneria, 104, 129, 313–316
Blighia, 110, 416, 417
Bocageopsis, 151
Bocoa, 112 C
Boehmeria, 74, 441, 442, 444 CACTACEAE, 32, 62, 63, 124, 187, 188, 202, 463
Boerhavia, 78, 346, 347 Cadia, 297
Bombax, 88, 105, 106, 312, 313, 315 Caesalpinia, 113, 118, 286–289
Bonamia, 214 Cajanus, 297, 299
Bonnetia, 70, 202, 205 Calamus, 138, 164, 166
BONNETIACEAE, 32, 33, 65, 66, 69, 70, 202, 205, 474 Calea, 79
Bonyunia, 303 Calliandra, 113, 292–294, 383
BORAGINACEAE, 32, 33, 61–63, 65–67, 71, 72, 178, Callicarpa, 80, 83, 85, 277, 445, 446
179, 476 Callistemon, 339, 341
Borassus, 38, 166 Callisthene, 339, 341
Borreria, 398 Callitris, 38, 78
Boscia, 187–189 Calodendrum, 418
Boswellia, 181, 182 Caloncoba, 59, 143, 144
Bouea, 43, 84, 147, 148 CALOPHYLLACEAE, 29, 30, 35, 39–43, 51, 83, 202,
Bougainvillea, 345, 346 204, 474
Bourreria, 63, 67, 179 Calophyllum, 43, 151, 202, 204, 205
Bowdichia, 115, 303 Calostephane, 169
Bowringia, 114 Calotropis, 43
Brachystegia, 287, 288 Calpocalyx, 291, 294
Brassaia, 161 Calycolpus, 341
Bravaisia, 79, 141, 143 Calycophyllum, 397, 398, 400
Brazzeia, 54, 284, 285 Calyptranthes, 82, 83, 339, 341
Bredemeyera, 126, 376, 377 Camellia, 53, 71, 197, 434, 435
Brexia, 72 Cameraria, 156
Breynia, 47, 242, 366 Campnosperma, 66, 148
Bridelia, 47, 96, 365, 366 Campomanesia, 86, 341
Brosimum, 40, 331–333 Campsiandra, 115, 289
Broussonetia, 40, 333, 441, 442, 444 Cananga, 151
Brownea, 113, 286, 289 Canarium, 87, 106, 107, 181, 182
Brownlowia, 105 Canavalia, 297, 299
Brucea, 424 Canella, 69, 72, 186, 187
Brugmansia, 428, 429 CANELLACEAE, 31, 33, 53, 54, 69, 71, 72, 187
Bruguiera, 75, 394 CANNABACEAE, see ULMACEAE
Brunellia, 91, 217 Cannabis, 439
BRUNELLIACEAE, 36, 91, 217, 562 Cansjera, 72, 355, 356
Brunfelsia, 73, 428 Canthium, 119, 134, 400
Buchanania, 107, 148 Capirona, 398
Buchenavia, 66, 206
480 Index
CAPPARACEAE, 30–33, 36, 46–48, 57, 58, 62–64, 68, Cephalosphaera, 337
69, 71, 73, 87, 88, 128, 187, 475 Ceratozamia, 39
Capparis, 8, 47, 48, 58, 63, 64, 71, 73, 128, 187–190 Cerbera, 42, 154, 155, 157
Capsicodendron, 54, 186 Cereus, 185
Capsicum, 428 Ceriops, 394
Caraipa, 42, 51, 202, 204 Ceroxylon, 166
Carallia, 392–394 Cespedesia, 349, 350
Carapa, 324, 325, 327 Cestrum, 51, 52, 73, 427–429
CARDIOPTERIDACEAE, 123, 476 Chadsia, 299
Cardiopteris, 123, 270 Chaetachme, 9, 44
Cardiospermum, 416–418 Chaetocarpus, 48, 96, 241, 243
Careya, 70, 71, 282–284 Chamaecrista, 112, 286, 287, 289
Carica, 41, 61, 191, 192 Chamaedorea, 165–167
CARICACEAE, 29, 32, 36, 41, 61, 90, 91, 191, 475 Chamaesyce, 42, 103, 241, 245
Cariniana, 283, 284 Chapelieria, 402
Carissa, 43, 154, 155, 157 Cheiloclinium, 195, 196
Carludovica, 39, 219 Chenopodium, 239
Carpolobia, 51 Chimarrhis, 398, 400
Carpotroche, 143–145 Chiococca, 402
Carya, 276 Chionanthus, 85, 86, 353–355, 370
Caryocar, 91 Chisocheton, 108, 110, 324, 326
CARYOCARACEAE, 36, 87, 88, 91, 474 CHLORANTHACEAE, 34, 74, 196, 197, 330, 466, 473
Caryodaphnopsis, 84 Chloranthus, 197
Caryodendron, 97 Chlorocardium, 281
Caryota, 163, 167 Chloroleucon, 117
Casearia, 47, 49, 50, 60, 410–413 Choisya, 406
Casimirella, 270, 271 Chondrodendron, 328
Casimiroa, 106, 410–413 Chorisepalum, 254, 255
Cassia, 112, 116, 286, 287, 289 Chrysalidocarpus, 38
Cassinopsis, 86 CHRYSOBALANACEAE, 29–31, 39, 40, 43, 44, 46, 48,
Cassipourea, 74, 76, 77, 393–394 53, 54, 198, 452, 474
Cassytha, 281, 282, 467 Chrysobalanus, 46, 198–200
Castanea, 49, 249, 250 Chrysophyllum, 41, 421, 422
Castanopsis, 47, 49, 58, 250, 251 Chukrasia, 324–326
Castela, 63, 424 Chusquea, 374
Castilla, 332, 333 Chydenanthus, 67, 282, 285
Casuarina, 78, 192, 193, 371 Cinchona, 398, 400
CASUARINACEAE, 34, 78, 192, 371, 474 Cinnamodendron, 54, 72, 186, 187
Catha, 195, 196 Cinnamomum, 61, 80, 280–281
Catharanthus, 155 Cinnamosma, 54, 186, 187
Cathedra, 54, 352 Cissampelos, 130
Catostemma, 105, 106, 313, 314, 316 Cissus, 45, 131, 137, 449, 450
Catunaregam, 119, 398, 402 Citharexylum, 77, 79, 81, 445, 446
Cavanillesia, 105 Citronella, 54, 269, 270
Cavendishia, 55, 62, 234, 235 Citrus, 107, 403, 404, 406
Cecropia, 18, 39, 56, 88, 331, 441, 442, 444 Cladogynos, 99, 243
Cedrela, 110, 111, 324, 325 Claoxylon, 60, 98
Cedrelinga, 117, 291 Clappertonia, 105
Ceiba, 88, 106, 312–316 Clarisia, 40, 332, 333
CELASTRACEAE, 8, 29–35, 42, 46, 49, 53, 55, 57–59, Clathrotropis, 115
62, 71, 72, 75–77, 86, 121, 123, 125, 134, Clausena, 403, 404
194–196, 467, 474 Clavija, 70, 380–382
Celastrus, 125, 194, 195 Cleidion, 96
Celtis, 9, 44, 45, 261, 439, 440 Cleistanthus, 95, 96, 365
Cenostigma, 289 Cleistopholis, 50, 154
Centrolobium, 114, 297–299 Clematis, 136, 474
CENTROPLACACEAE, 57, 196, 474 CLEOMACEAE, 187, 188
Cephalaralia, 138 Cleome, 188, 190
Cephalomappa, 97 Clerodendron, 80, 85, 133
Index 481
Clethra, 69, 71, 200, 201 Coursetia, 113

CLETHRACEAE, 33, 69, 71, 201, 202, 475 Coussapoa, 39, 56, 441–444
Clidemia, 322 Coussarea, 399
Clitoria, 114, 297, 299 Coutoubea, 79, 254
Clusia, 42, 43, 200, 201 Craibiodendron, 73
CLUSIACEAE, 29, 31, 34, 35, 42, 43, 51, 75, 76, 79, 82, Craterispermum, 402
83, 121, 202–204, 474 Craterosiphon, 84, 134, 437
Cnesmone, 98 Crateva, 88, 187, 189
Cnestis, 209 Cratoxylum, 43, 82, 202, 204
Cnidoscolus, 242, 243 Crepidospermum, 181
Cobaea, 137 Crescentia, 63, 90, 173, 175
Coccocypselum, 398 Crossopetalum, 77, 86, 195, 196
Coccoloba, 43, 55, 125, 127, 378, 379 Crossostylis, 395
Cocculus, 60, 328 Crotalaria, 297, 299
Cochlospermum, 39, 88, 176–178 Croton, 58, 60, 61, 64, 97, 99, 102, 241, 242, 244
Cocos, 163, 165–167 Crotonogyne, 100, 244
Codiaeum, 68, 99, 242 Crotonoides, 96
Coffea, 75, 119, 398, 400 Crudia, 112, 286, 287
Cojoba, 113 Crumenaria, 390
Cola, 88, 105, 106, 313, 315, 316 Crypteronia, 79
Colea, 92 CRYPTERONIACEAE, 34, 79, 475
Coleonema, 404 Cryptocarya, 52, 280, 281
Colletia, 390 Ctenolophon, 75
Colona, 104, 313 CTENOLOPHONACEAE, 34, 74, 75, 474
Colophospermum, 112, 113, 288 CUCURBITACEAE, 121, 131
Colubrina, 47, 58, 60, 74, 390 CUNONIACEAE, 34, 36, 74, 75, 91, 217, 232, 474
COMBRETACEAE, 32, 33, 35, 62–66, 71, 72, 81–83, Cupania, 109, 416, 418
121, 133, 135, 205, 475 Cuphea, 305–307
Combretum, 81–83, 133, 135, 205–207 CUPRESSACEAE, 29, 34, 37, 38, 78
Commiphora, 108, 182 CuraDilleniatella, 223
Comocladia, 147 Curatella, 68, 69, 222, 223, 383
Comoranthus, 86 Cuscuta, 212
Conceveiba, 98, 241 Cusparia, 90
Congea, 80, 278 Cussonia, 90, 160
CONNARACEAE, 31, 32, 36, 51, 52, 62, 63, 89, 90, 121, Cybianthus, 381, 382
124, 136, 209, 467, 574 Cybistax, 92
Connarus, 89, 124, 136, 209–212 CYCADACEAE, 29, 37, 39, 218
Conocarpus, 64, 206–208 Cycas, 39, 218
CONVOLVULACEAE, 121, 123, 126, 130, 136, 225, 476 CYCLANTHACEAE, 29, 37–39, 121, 124, 138, 166, 219
Copaifera, 287, 288, 290 Cyclanthus, 219
Coprosma, 397, 399 Cyclea, 328
Corchorus, 315 Cylicomorpha, 41, 191
Cordia, 61, 63, 66, 67, 178–180 Cynanchum, 155
Cordyla, 111 Cynometra, 112, 113, 286
Cordyline, 37, 38 Cyphostemma, 137, 449
CORNACEAE, 31, 53, 69, 71, 201, 215, 475 Cyrilla, 71, 200, 201
Cornus, 215 CYRILLACEAE, 33, 69, 71, 200, 475
Corylopsis, 261
CORYNOCARPACEAE, 31, 58, 59, 216, 474
Corynocarpus, 59, 216, 258 D
Corynostylis, 125, 447 Dacrydium, 375
Corypha, 163, 165 Dacryodes, 107, 110, 181–183
Cossinia, 109 Dactyladenia, 46, 200
Cotylelobium, 225, 227 Dahlia, 170
Couepia, 48, 198–200 Dalbergia, 48, 111, 112, 114, 128, 297, 299
Coula, 41, 237, 238, 351, 352 Dalechampia, 59, 98, 241, 242, 244
Coumarouna, 116, 297 Daniellia, 287, 288
Couratari, 50, 54, 284 Dapania, 129, 357
Couroupita, 59, 293, 294 Daphne, 435–437
482 Index
DAPHNIPHYLLACEAE, 33, 69, 70, 474 Diospyros, 51, 52, 229–231

Daphniphyllum, 70 Diplotropis, 115, 297
Daphnopsis, 67, 437 DIPTEROCARPACEAE, 29–31, 39, 40, 43, 44, 46–48,
Dasyphyllum, 168–170 56, 58, 60, 225, 226, 452, 475
Datura, 427, 428 Dipterocarpus, 44, 225–227
Davilla, 125, 223 Dipteryx, 90, 116, 297, 300
Debregeasia, 74, 441, 443 Dirca, 437
Dehaasia, 66, 281 Disciphania, 136
Deherainia, 380, 383 Discoglypremna, 98
Deinbollia, 418 Discophora, 53, 269, 271
Delonix, 118, 286–288 Dissotis, 322
Delpydora, 421, 422 Distylium, 48
Dendrobangia, 52, 269–271 Dodonaea, 65, 416–418
Dendrocalamus, 38, 374 Dolichandra, 135, 173
Dendrocnide, 441–444 Doliocarpus, 125, 222
Dendrolobium, 300 Dombeya, 56, 104, 105, 313
Dendropanax, 60, 67, 160 Domohinea, 101
Dendrophthoe, 78 Dorstenia, 332, 333
Dendrophthora, 415 Dracaena, 37
Denhamia, 77 Dracontomelon, 10, 147, 148
Deplanchea, 81, 173 Drimys, 65, 186, 187
Derris, 114, 136, 297 Dryobalanops, 225, 227
Desbordesia, 272 Drypetes, 49, 50, 96, 386, 487
Desmodium, 297 Duabanga, 76, 82, 306, 307
Desmoncus, 138, 163, 165 Dubouzetia, 232
Desmos, 151, 152 Duguetia, 50
Desmostachys, 126, 127, 269, 270 Dulacia, 55, 352
Desplatsia, 104 Duranta, 445, 446
Detarium, 286, 287 Durio, 44, 47, 103, 313–316
Dialium, 286 Duroia, 397
Dichaeanthera, 77, 322 Dussia, 297, 300
DICHAPETALACEAE, 8, 30, 46, 48, 121, 125, 127, 222, Dyera, 154, 155, 459
474 Dysoxylum, 92, 324–326
Dichapetalum, 46, 48, 125, 127, 221, 222
Dichostemma, 103, 244
Dichrostachys, 113, 117, 291, 292, 294 E
Diclidanthera, 376, 377 EBENACEAE, 30, 31, 35, 51, 52, 81, 229, 230, 463, 465
Dicorynia, 111, 112, 115, 286 Ecclinusa, 40, 421
Dicoryphe, 48, 261 Echium, 178, 179, 466
Dicranolepis, 50, 129, 435–437 Edgeworthia, 474
Dicranopygium, 220 Ehretia, 72, 180, 181
Dicranostyles, 126, 212, 213 ELAEAGNACEAE, 128, 502
Didymopanax, 159–161 Elaeagnus, 128
Dillenia, 55, 68, 223, 224 Elaeis, 38, 166
DILLENIACEAE, 31, 33, 55, 67–69, 121, 125, 146, 222, ELAEOCARPACEAE, 31, 33, 34, 56–60, 68, 76, 231,
223, 474 474
Dilodendron, 110, 416, 418 Elaeocarpus, 56, 57, 59, 60, 68, 231–233
Dimocarpus, 416, 417 Elaeodendron, 77
Dimorphandra, 118, 287, 290 Elateriospermum, 39, 100, 244, 248
Dimorphanthera, 234 Elatostema, 441
Dinizia, 293 Ellipanthus, 52, 209, 212
Dioclea, 88, 114, 136, 297, 300 Elmerrillia, 309
Diocrescis, 399 Elvasia, 349
DIONCOPHYLLACEAE, 121, 127, 475 Embelia, 51, 382, 383
Dioncophyllum, 127 Embothrium, 386
Dionycha, 322 Emmotum, 53, 67, 269–271
Dioscorea, 129, 133, 136, 224, 225 Empetrum, 234
DIOSCOREACEAE, 121, 129, 133, 136, 224, 225, 328, Encephalartos, 218, 219
473 Endodesmia, 42
Index 483
Endospermum, 101 F
Engelhardtia, 90, 275, 276 FAGACEAE, 30, 31, 46, 47, 49, 58, 192, 250, 251, 474
Enicostema, 255 Fagopyrum, 379
Enkleia, 134, 437 Fagraea, 79, 254, 255
Entada, 116, 137, 292–294 Fagus, 249, 251
Entandrophragma, 111, 325 Faidherbia, 294
Enterolobium, 113, 117, 291–293 Falcataria, 292
Ephippiandra, 329, 330 Fallopia, 379
Epinetrum, 328, 329 Faramea, 119, 398, 399
Epiphyllum, 124 Fatoua, 332
Eranthemum, 141, 143 Faurea, 67, 384
Erica, 234 Feijoa, 297, 339–341, 339–341
ERICACEAE, 31–33, 51, 53, 55, 61, 62, 69–71, 73, 128, Fernandoa, 175
234, 475 Ficus, 39, 40, 42, 331–335
Eriolaena, 105 Firmiana, 56, 105
Eriotheca, 106 Flabellaria, 85, 132, 133
Erisma, 10, 452 Flacourtia, 59, 410–412
Erismadelphus, 75, 452 Flaveria, 169
Eritrichium, 179 Flemingia, 45, 112, 297
Ernodea, 399 Flindersia, 91, 403, 406
Erycibe, 212–214 Flueggea, 98
Eryobotrya, 56, 396 Foetidia, 70, 285
Erythrina, 88, 114, 286, 297, 300 Fordia, 297
Erythrochiton, 389, 404 Forsteronia, 123, 155
ERYTHROPALACEAE, 29, 31, 41, 51–54, 131, 237, 352, Forsythia, 86, 353, 355
475 Fraxinus, 92, 353–355
Erythropalum, 131, 237 Fremontodendron, 104
Erythrophleum, 117, 118 Freycinetia, 128, 360, 361
Erythrostemon, 289 Freziera, 52, 363, 364
ERYTHROXYLACEAE, 30, 34, 46, 47, 74, 75, 239, 302, Froesia, 91, 349
474 Fuchsia, 83
Erythroxylum, 47, 239, 240 Funtumia, 43, 155
Eschweilera, 49, 50, 54, 282–285
Esenbeckia, 65, 66, 107, 403, 405
Espeletia, 169 G
Euadenia, 88, 188 Gaertnera, 398, 399
Eucalyptus, 64, 339–341 Gagnebina, 113
Euceraea, 410–412 Galearia, 95
Euclea, 81, 229, 230 Garcinia, 18, 42, 43, 76, 202–205
Eugenia, 81–83, 339–341 Gardenia, 75, 119, 400, 402
Euodia, 81, 106, 403, 406 Gardneria, 133
Euonymus, 76, 194–196 Garuga, 90, 106, 181
Euphorbia, 42, 103, 241, 242, 245 Gastonia, 160
EUPHORBIACEAE, 3, 7, 8, 10, 29–34, 36, 39, 40, 42, Gaultheria, 234
44–48, 52, 55–62, 64, 68, 74, 76, 77, 87, 88, 95, Gaussia, 164
121, 123, 129, 241, 242, 248, 365, 474 Gaylussacia, 234, 236
Euphronia, 48 Geissois, 74
EUPHRONIACEAE, 48, 474 Geissospermum, 154, 156
Eurya, 363, 364 GELSEMIACEAE, 75, 80, 133, 303, 304, 476
Eurycoma, 109, 425 Gelsemium, 133
Eurypetalum, 112, 113 Geniostoma, 303–305
Eusideroxylon, 53, 280 Genipa, 397, 402
Euterpe, 38, 163, 165–167 Gentiana, 254, 255
Evodianthus, 38, 124, 138, 219 GENTIANACEAE, 34, 35, 78, 79, 255, 476
Evolvulus, 212–214 Geonoma, 37, 164, 166
Exbucklandia, 44, 261, 262 Geum, 395
Excoecaria, 103, 241, 242 Gigantochloa, 38, 374
Exocarpos, 72, 414, 415 Gilbertiodendron, 113, 286, 288
484 Index
Gironniera, 9, 44, 439 HAMAMELIDACEAE, 30, 32, 33, 43–46, 48, 60, 69, 70,
Givotia, 101, 242 262, 474
Gleditsia, 115, 286 Hamamelis, 261
Gliricida, 114 Hamelia, 397, 399, 400
Glochidion, 95, 366 Hampea, 104
Glossocalyx, 86, 329, 330 Hancea, 74, 99
Gluta, 107, 108, 147, 148 Hancornia, 155
Glycosmis, 405 Handroanthus, 173, 174
Glycydendron, 45 Harmsiopanax, 160
Gmelina, 77, 80, 277–279 Harpullia, 109, 110, 417
GNETACEAE, 34 Harrisonia, 87, 106, 109, 403, 405
Gnetum, 78, 80, 123, 131, 256, 257 Harungana, 43, 202
Gnidia, 437 Hasseltia, 58, 411
Gochnatia, 65 Hebepetalum, 59, 302
Godmania, 92, 173, 175 Hedycarya, 330
Gomphandra, 53, 269, 270 Hedyosmum, 74, 197
Gongylolepis, 169, 170 Heisteria, 41, 54, 237, 238, 352
Goniothalamus, 151, 153 Helicia, 3, 67, 384, 385
Gonocaryum, 53, 269–271 Heliciopsis, 385
Gonystylus, 9, 50, 435, 437 Helicteres, 104, 313, 317
Goodenia, 258 Helinus, 131
GOODENIACEAE, 32, 33, 65, 66, 69, 70, 258, 476 Heliocarpus, 105
Gordonia, 71, 72, 434, 435 Heliotropium, 179, 180
Gossweilerodendron, 288 Hemandradenia, 52
Gossypium, 105, 313–316 Henriettea, 322
Gouania, 45, 131, 390, 391 Heritiera, 9, 104, 105, 312, 313, 315
Goupia, 18, 50, 258, 259 Hernandia, 62, 262, 263
GOUPIACEAE, 30, 50, 258, 474 HERNANDIACEAE, 32, 61, 62, 121, 130, 131, 136, 137,
Grabowskia, 427 262, 473
Grevea, 85, 331 Herrania, 313, 317
Grevillea, 90, 384 Hesperomeles, 396
Grewia, 44, 104, 313, 314, 317, 336 Heteropsis, 32, 37, 129
Grias, 67, 70, 283, 284 Heteropterys, 75, 311
Griffonia, 111 Heterosmilax, 426
Grossera, 100 Hevea, 88, 100, 101, 241, 242, 245
Guadua, 374 Hexalobus, 151
Guaiacum, 91, 455, 456 Hibbertia, 125, 222
Guapira, 345, 346 Hibiscus, 17, 55, 104, 313–315, 317–319
Guarea, 108, 110, 324, 326 Hieronyma, 60, 99, 365, 366
Guatteria, 50, 151, 153 Hildegardia, 105
Guazuma, 104, 313, 314, 317 Hillia, 134
Guettarda, 75, 119, 399 Himatanthus, 42, 154
Guibourtia, 287 Hippocratea, 194, 196
Guiera, 206 Hippomane, 99, 102, 103, 242, 245
Gustavia, 67, 70, 282–285 Hiptage, 132, 311
Gymnanthes, 40, 99, 103 Hiraea, 310–312
Gymnocladus, 118, 287 Hirtella, 48, 198–200
Gymnopodium, 378–380 Hoffmannia, 119
Gymnosporia, 62, 72, 194–196 Holarrhena, 155
Gynocardia, 143, 144 Holigarna, 107
Gynotroches, 76, 394 Holoptelea, 439, 440
Gyranthera, 106, 312, 314, 317 Homalanthus, 102, 241, 245
Gyrocarpus, 62, 262, 263 Homalium, 48, 50, 410, 411, 413
Homonoia, 250
Hopea, 47, 48, 225, 228
H Horsfieldia, 337, 338
Haematoxylum, 115, 286, 288, 290 Hovenia, 391
Hagenia, 87, 395, 396 Hoya, 43, 155
Hakea, 384 Hua, 264, 265
Index 485
HUACEAE, 30, 46, 49, 264, 474 J

Huertea, 87, 475 Jacaranda, 92, 173, 175
Hugonia, 127, 302 Jacaratia, 41, 91, 191, 192
Humiria, 64, 72, 265, 266 Jacquinia, 70, 380, 382, 383
HUMIRIACEAE, 30–33, 49, 50, 53–55, 64, 71, 72, 265, Jagera, 416
266, 302, 474 Jasminum, 132, 134–136, 353, 354
Humiriastrum, 50, 265, 266 Jatropha, 100–102, 241, 242, 246
Hura, 102, 241, 242, 246 Joannesia, 88
Hybanthus, 50, 447 Johannesteijsmannia, 37
Hydnocarpus, 57, 143, 144 Jollydora, 212
Hydnophytum, 400, 402 JUGLANDACEAE, 36, 89, 90, 275, 474
Hydrangea, 79, 475 Juglans, 90, 275, 474
HYDRANGEACEAE, 79, 475 Justicia, 141
Hydrocotyle, 160
Hydrophyllum, 178
Hygrophila, 141 K
Hylocereus, 112, 113, 186 Kadsura, 125, 424
Hymenaea, 286 Kageneckia, 396
Hymenocardia, 98, 365 Kallstroemia, 455
Hymenosporum, 114, 373 Kalopanax, 159, 161
Hyophorbe, 164, 167 Kandelia, 394
Hypenia, 278, 279 Karomia, 277
Hyperbaena, 328 Karwinskia, 392
HYPERICACEAE, 43, 79, 82, 202, 205, 474 Khaya, 110, 324, 325, 327
Hypericum, 202, 205 Kibara, 329, 330
Hyphaene, 38, 163, 165 Kielmeyera, 42, 202
Hypodaphnis, 280 Kigelia, 92, 173, 174
Hyptidendron, 283 Kiggelaria, 62, 143–145
Hyptis, 278, 279 Kingiodendron, 112, 286, 288
Kirkia, 90
KIRKIACEAE, 89, 90
I Kleinhovia, 105, 314, 318
ICACINACEAE, 31, 33, 35, 51–54, 67, 71, 73, 84, 86, Knema, 337, 338
121, 123, 126–128, 130, 131, 269, 271, 476 Kokoona, 196
Ilex, 8, 59, 158, 159 Koompassia, 288
Illicium, 65, 424 Korupodendron, 452
Illigera, 121, 136, 137, 262, 263 Kostermansia, 312
Incarvillea, 173 Krugiodendron, 392
Indigofera, 297, 298 Kydia, 105, 315
Indorouchera, 127, 302
Indovethia, 350
Inga, 112, 286, 291–295 L
Inocarpus, 39, 297 Labisia, 382, 383
Intsia, 287, 297 Lablab, 297
Iochroma, 428 Labordia, 303–305
Iodes, 131, 269–271 Lacistema, 30, 50, 412, 414
Ipomoea, 123, 212–214 LACISTEMATACEAE, 49, 50, 410, 474
Iriartea, 166, 167 Lacmellea, 154
Irlbachia, 254, 255 Lacunaria, 349, 350
Irvingia, 44, 272, 273 Ladenbergia, 399, 400
IRVINGIACEAE, 30, 43, 44, 271, 302, 474 Laetia, 46, 47, 410, 411, 413
Iryanthera, 337, 338 Lafoensia, 83, 305, 306
Isertia, 398 Lagerstroemia, 8, 76, 82, 83, 305–307
Isolona, 152 Laguncularia, 81, 206, 208
IXONANTHACEAE, 31, 33, 53, 54, 58, 59, 69, 71, 72, Lamanonia, 75
302, 474 LAMIACEAE, 34–36, 77, 79, 80, 84–86, 91, 92, 121,
Ixonanthes, 59, 71, 302, 303 133, 134, 288, 459, 504
Ixora, 119, 399, 400 Landolphia, 123, 155
486 Index
Lannea, 110 Livistona, 38

Lansium, 326 Lobelia, 258, 466, 469
Lantana, 445, 446 Logania, 304
Laportea, 441, 443, 444 LOGANIACEAE, 34, 74, 75, 77, 79, 80, 121, 133–135,
Larrea, 455, 456 303, 476
Lasianthera, 54, 269, 271 Lonchocarpus, 297, 300
Lasianthus, 119, 399 Lophanthera, 42
Lasiodiscus, 74, 390 Lophira, 349, 350
Lasiosiphon, 435, 436 Lophopetalum, 76, 194–196
Latania, 165 LOPHOPYXIDACEAE, 126
Latua, 427 Lophopyxis, 126
LAURACEAE, 9, 31–33, 35, 51–53, 61–66, 69, 71, 73, Lophostoma, 84, 437
80, 84, 280–282, 467, 473 LORANTHACEAE, 34, 78, 414, 415, 475
Laurelia, 330 Loranthus, 414, 415
Laurus, 280 Loreya, 322
Lawsonia, 306 Loxopterygium, 108
Leandra, 333 Lozanella, 74, 439
Lecointea, 112 Ludia, 73, 412
LECYTHIDACEAE, 30, 31, 33, 49, 50, 53, 54, 56–59, 67, Ludovia, 38, 219
69–71, 282, 285, 475 Ludwigia, 83
Lecythis, 49, 54, 282, 284, 285 Luehea, 104, 313, 314, 318
Leea, 43, 55, 86, 448–451 Lueheopsis, 104
LEGUMINOSAE, 7, 29, 30, 33, 35, 36, 39, 44–46, Lumnitzera, 72, 206, 208
48–50, 71, 73, 86–91, 121, 123, 124, 128, 130, Luvunga, 138, 405, 406
136–138, 285, 286, 291, 294, 297, 466, 474 Luxemburgia, 350
Leonardoxa, 286, 287, 290 Lycianthes, 442
Leonia, 48, 60, 447, 448 Lycium, 63, 427, 428
Lepianthes, 372 Lysiloma, 113, 295
Lepidagathis, 142 LYTHRACEAE, 8, 34, 35, 75, 76, 78, 79, 82, 83, 305,
LEPIDOBOTRYACEAE, 33, 68 308, 475
Lepidobotrys, 68 Lythrum, 306
Lepidotrichilia, 110, 324
Lepidozamia, 218
Lepionurus, 54, 72, 352, 355, 356 M
Lepisanthes, 109, 416, 417 Mabea, 103, 246
Leptaulus, 53, 269, 271 Maburea, 352
Leptolaena, 44 Macadamia, 384, 385
Leptospermum, 64, 339, 342 Macaranga, 56, 58, 59, 99, 241, 242, 246
Leretia, 73, 270 Machaerium, 111, 112, 123, 138, 297, 300, 301
Leucadendron, 386 Macleania, 234
Leucaena, 117, 292–295 Maclura, 332, 333
Leucospermum, 386 Macphersonia, 110
Leucosyke, 44, 455 Macrocnemum, 402
Licania, 40, 48, 54, 198–200 Macrolobium, 113, 286, 287, 290
Licaria, 71 Macropsychanthus, 297, 300
Licuala, 164, 165 Macroptilium, 301
Ligustrum, 353 Madhuca, 422
LINACEAE, 31, 58, 59, 121, 127, 302, 474 Maerua, 187, 188
Lindackeria, 57, 68, 143, 145 Maesa, 380, 382, 383
Linociera, 85, 86, 354, 355 Maesobotrya, 96, 365, 367
Linum, 302, 303 Maesopsis, 49, 390, 310
Lippia, 445 Magnolia, 43, 56, 309, 310
Liquidambar, 261 MAGNOLIACEAE, 30, 31, 43, 55, 56, 310, 337, 473
Liriodendron, 309, 310 Mahonia, see Berberis, 89
Lisianthus, 78, 253–255 Mahurea, 40, 202, 204
Litchi, 416, 417 Maieta, 321–323
Lithocarpus, 47, 111, 114, 249–251 Maingaya, 48, 261
Lithraea, 147 Mallotus, 58, 60, 74, 98, 99, 241, 242, 246
Litsea, 71, 280–282 Malpighia, 311, 312
Index 487
MALPIGHIACEAE, 8, 29, 31, 34, 35, 42, 58, 75, 81, 84, Meriania, 322
85, 121, 125, 132–134, 310, 311, 474 Merremia, 136, 212, 214
Malus, 395 Meryta, 160
Malva, 314, 318 Mesechites, 155, 157
MALVACEAE, 7, 9, 30, 31, 36, 43, 44, 46, 47, 55, 56, 87, Mesua, 43, 83, 202, 204, 205
88, 103, 312, 315, 336, 475 Metopium, 147–149
Mammea, 43, 202, 204, 205 Metrodorea, 81, 107
Manettia, 132, 400 Metrosideros, 339
Mangifera, 41, 69, 107, 147–149 Metroxylon, 163, 166
Manihot, 101, 242, 246 METTENIUSACEAE, 52, 53, 67, 128, 476
Manilkara, 41, 421–423 Mezoneuron, 118, 138, 286, 287
Maniltoa, 113 Michelia, 309, 310
Manniophyton, 100, 123 Miconia, 77, 80, 321–323
Manotes, 52, 124, 209 Micrandra, 101
Mansonia, 105 Microcos, 314, 318
Mantalania, 402 Microdesmis, 95, 359
Maprounea, 102 Micromelum, 405
Maranthes, 200 Micropholis, 421, 422
Marcetia, 323 Mikania, 132, 168–170
Marcgravia, 55, 128, 320 Milicia, 332, 333
MARCGRAVIACEAE, 31, 53, 55, 121, 128, 320, 475 Millettia, 114, 115, 137, 297, 301
Mareya, 57, 97 Millingtonia, 92
Marila, 43 Mimosa, 113, 116, 117, 291–293, 295
Maripa, 126, 214 Mimusops, 40, 421, 422
Markea, 129, 428 Minquartia, 54, 351, 352
Markhamia, 92, 173, 175 Mirabilis, 77, 78, 345, 346
Marlierea, 341 Mischocarpus, 110
Marquesia, 46, 226 Mischodon, 76, 85, 98, 370
Mastixia, 71, 215, 216 Mitracasme, 304
Matayba, 92, 109, 416, 418 Mitragyna, 400
Matelea, 157 Mitreola, 304
Matisia, 104 Mollia, 103, 313
Matthaea, 329, 330 Mollinedia, 85, 86, 329–331
Matudaea, 45, 261 Monimia, 330
Mauritia, 38 MONIMIACEAE, 35, 81, 82, 84–86, 197, 329, 330, 473
Maytenus, 49, 195, 196 Monniera, 106, 403, 406
Medinilla, 77, 80, 321, 322 Monnina, 377
Medusandra, 57 Monodora, 50
MEDUSANDRACEAE, 31, 57, 474 Monosis, 169
Melaleuca, 340 Monotes, 46, 225–227
Melanophylla, 68 Montanoa, 169
Melastoma, 80, 322, 324 MONTINIACEAE, 35, 84, 85, 331, 476
MELASTOMATACEAE, 34, 35, 77, 78, 80, 82, 84, 85, Montrouziera, 202
321, 322, 339, 377, 475 Mora, 116, 286, 288, 290
Melia, 89, 108, 324, 325, 327 MORACEAE, 3, 29, 39, 40, 42, 331, 333, 421, 443, 474
MELIACEAE, 7, 31, 32, 36, 51, 52, 65, 66, 89–92, Morella, 72, 336
106–111, 324, 325, 327, 475 Morinda, 119, 397, 399, 400, 402
MELIANTHACEAE, 86, 87, 475 Moringa, 89, 335
Melianthus, 87 MORINGACEAE, 3, 36, 89, 335, 475
Melicoccus, 110, 416, 418 Morisonia, 68, 71, 73, 187, 188
Melicytus, 447 Moronobea, 202, 203
Melientha, 72, 355, 356 Morus, 332, 334
Meliosma, 67, 90, 409, 410 Mostuea, 75, 80, 303, 304
Melochia, 313, 318 Mouriri, 78, 82, 321–324, 339
Melodorum, 124 Moutabea, 126, 376, 377
Memecylon, 78, 82, 85, 322, 323 Mucuna, 114, 123, 136, 297, 301
Mendoncia, 132, 141 Muehlenbeckia, 380
MENISPERMACEAE, 32, 60, 121, 130, 136, 328, 329, Muntingia, 44, 336
473 MUNTINGIACEAE, 30, 44, 336, 475
488 Index
Murraya, 106, 403, 406 Nyctanthes, 77, 353, 354

Musanga, 88, 331, 441, 443, 444 Nymphaea, 197, 468
Mussaenda, 75, 119, 397, 398, 400 Nyssa, 71
Myllanthus, 106 NYSSACEAE, 33, 215, 475
Myosotis, 178
Myrcia, 339, 341
Myrcianthes, 341 O
Myrciaria, 339, 341 Ochanostachys, 54, 352
Myrianthus, 56, 88, 441, 444 Ochna, 47, 349–351
Myrica, 336 OCHNACEAE, 30, 31, 34, 36, 46, 47, 49, 58, 76, 91, 302,
MYRICACEAE, 33, 71, 72, 336, 474 350, 474
Myriocarpa, 56 Ochroma, 56, 105, 313–315
Myristica, 41, 52, 337, 338 Ochthocosmus, 72, 302, 303
MYRISTICACEAE, 29, 31, 41, 51, 52, 337, 473 Ocimum, 278
Myrmecodia, 400 Ocotea, 66, 73, 281, 282
Myrocarpus, 297 Octoknema, 52, 237, 238
Myrospermum, 297 Octolobus, 314
Myroxylon, 111, 297 Octomeles, 61, 62, 433, 434
Myrsine, 65, 380, 382, 383 Odyendea, 109, 424
MYRTACEAE, 8, 29, 32, 35, 42, 43, 64, 80–84, 86, 339, Okoubaka, 10, 52, 414, 415
341, 475 OLACACEAE, 31, 32, 51, 53–55, 62, 127, 351, 352, 356,
Myrteola, 341 475
Mystroxylon, 59, 72, 196 Olax, 55, 352
Myxopyrum, 77, 132, 353, 354 Oldfieldia, 93, 370
Olea, 16, 82, 86, 353–355
OLEACEAE, 34–36, 77, 81, 82, 84–86, 91, 92, 121, 132,
N 134–136, 353, 354, 370, 476
Napoleonaea, 50, 283, 284 Omphalea, 100, 129, 242, 246
Natsiatum, 130 ONAGRACEAE, 35, 82, 83
Nauclea, 400 Oncostemum, 65, 380, 382, 383
Nectandra, 61, 280–282 Ongokea, 352
Neea, 84, 85, 345, 346 Ophiocaryon, 90, 409
Neesia, 105 Ophiocolea, 174
Nemuaron, 330 Opilia, 54, 356
Neoboutonia, 101 OPILIACEAE, 31, 33, 53–55, 71, 72, 352, 355, 356, 475
Neolamarckia, 402 Opuntia, 186
Neolitsea, 66 Oreocallis, 384
Neoscortechinia, 98 Oreocnide, 56, 444
Neostachyanthus, 126 Oreomunnea, 275
Nephelium, 110, 416–418, 420 Ormosia, 115, 297
Neptunia, 293 Oroxylum, 173
Nerium, 155 Osmanthus, 354
Nesiota, 392 Osmoxylon, 88, 160
Neuburgia, 75, 303–305 Ostodes, 40, 99
Neurolaena, 169 Osyris, 415
Neuropeltis, 126, 212, 214 Ouratea, 47, 49, 58, 302, 349–351
Newbouldia, 173 OXALIDACEAE, 18, 33, 36, 68, 89–91, 129, 357, 474
Newtonia, 113 Oxalis, 91, 357
Nicandra, 427, 429 Oxera, 278
Nicotiana, 428 Oxygonum, 378, 380
Niemeyera, 420 Ozoroa, 69, 107, 147, 148
Norantea, 55, 128, 320
Noronhia, 85, 353, 354
Nothapodytes, 270 P
NOTHOFAGACEAE, 251 Pachira, 106, 314, 318
Notopora, 236 Pachyelasma, 118
Nuytsia, 415 Pachylobus, 110
NYCTAGINACEAE, 34, 35, 77, 78, 82–85, 121, 135, Pachypodium, 154, 155
346, 475 Pachystroma, 103
Index 489
Paederia, 132 Peronema, 92, 277

Pagamea, 398 Perriera, 109, 424
Pakaraimaea, 226 Persea, 71, 281
Palaquium, 40, 202, 421–423 Persicaria, 378, 380
Palicourea, 398, 400, 401 Persoonia, 384
Palmeria, 330 Petersianthus, 70, 285
Panax, 160 Petiveria, 369
Panda, 95, 359 PETIVERIACEAE, 369
PANDACEAE, 30, 46, 95 Petrea, 133, 445, 446
PANDANACEAE, 29, 37, 121, 128, 219, 360, 361, 466, Petunia, 428
473 Peumus, 330
Pandanus, 37, 219, 360, 361 Phacelia, 178
Pandorea, 136 Phainantha, 322, 324
Pangium, 56, 61, 143, 144 Phaleria, 84, 435–437
Panopsis, 84, 86, 384, 385 Phaseolus, 297
Paranephelium, 92 Philenoptera, 115, 301
Parashorea, 60 Philgamia, 311
Parastyrax, 430 Phoebe, 66, 280
Parinari, 46, 198, 200 Phoenix, 38, 163, 164, 166
Parishia, 147, 148 Phoradendron, 414
Parkia, 117, 291, 295 Phthirusa, 78, 415
Parkinsonia, 118 PHYLLANTHACEAE, 7, 17, 29, 30, 33, 36, 39–42,
Parmentiera, 92, 173, 175 46–48, 68, 69, 87, 88, 95, 365, 474
Parnassia, 467 Phyllanthus, 17, 47, 89, 95, 96, 365–368
Paropsia, 46, 361, 362 Phyllarthron, 174
Parrotia, 261 Phyllobotryon, 59, 143–145
Parsonsia, 123 Phyllocosmus, 54, 302, 303
Passiflora, 45, 58, 126, 130, 137, 361–363 Phyllostachys, 374
PASSIFLORACEAE, 30–32, 44–46, 58, 64, 121, 126, Physalis, 427, 429
127, 130, 137, 361, 474 Physena, 73
Patagonula, 180 PHYSENACEAE, 33, 71, 73, 368, 475
Paullinia, 87, 106, 124, 137, 416, 417, 419 Physocalymma, 83
Pausandra, 64, 97, 100, 241, 247 Phytocrene, 130, 269, 270
Pavetta, 75, 421 Phytolacca, 73, 369, 370, 466
Payena, 421 PHYTOLACCACEAE, 33, 73, 121, 128, 369, 475
Paypayrola, 447, 448 Picramnia, 89, 109, 424
Pedilanthus, see Euphorbia PICRAMNIACEAE, 36, 89, 106, 109, 424, 475
Pellacalyx, 76 Picrasma, 87, 89, 106, 424
Pelliciera, 70 PICRODENDRACEAE, 34–36, 75, 76, 84, 85, 90, 92,
Peltogyne, 112, 286–288 93, 95, 98, 99, 370, 474
Peltophorum, 118, 288 Picrodendron, 90, 370
Pemphis, 82 Picrolemma, 424
Pentace, 326 Pilea, 441
Pentaclethra, 117, 118, 286, 287, 291, 295 Pilocarpus, 65, 106, 406
Pentadesma, 205 Pimelodendron, 102
Pentagonia, 402 Pimenta, 83, 339, 340, 342
Pentalinon, 157 PINACEAE, 17, 29, 34, 37, 78, 163, 371
PENTAPHYLACACEAE, 31–33, 51, 52, 64, 65, 69, 70, Pinanga, 166
72, 363, 435, 475 Pinus, 37, 78, 371
Pentaspadon, 107 Pinzona, 125, 222–224
Peperomia, 371, 372 Piper, 53, 54, 61, 371, 372
Pera, 52, 96, 98, 241, 242, 248 PIPERACEAE, 31, 32, 53, 54, 61, 371, 372, 473
PERACEAE, 31, 51, 52, 96, 98, 241, 242, 474 Piptadenia, 116, 117, 291–293, 296
Perebea, 39, 331 Piptadeniastrum, 116
Pereskia, 63, 185, 186 Piptocoma, 168
Pericopsis, 297 Piptostigma, 151
PERIDISCACEAE, 44, 45 Piranhea, 99, 370
Peridiscus, 45 Pisonia, 83, 85, 135, 345, 346
Peristrophe, 142 Pithecellobium, 117, 286, 291, 296
490 Index
PITTOSPORACEAE, 33, 69, 70, 373, 476 Prunus, 46, 51, 395, 396
Pittosporum, 70, 373 Pseuderanthemum, 141, 143
Planchonella, 422 Pseudobombax, 105
Planchonia, 67 Pseudoconnarus, 136
Platonia, 42, 202, 204 Pseudolmedia, 39, 331, 334
Platymiscium, 91, 114, 297, 301 Pseudomonotes, 226
Platysepalum, 297 Pseudopiptadenia, 117
Plectranthus, 278 Pseudoprosopis, 116, 137
Plenckia, 195 Pseudosamanea, 117, 292
Pleurostylia, 76 Pseudospondias, 149
Ploiarium, 66, 202 Psidium, 8, 339, 340, 342
Pluchea, 168, 169 Psittacanthus, 414, 415
Plukenetia, 98, 242, 247 Psophocarpus, 297
Plumeria, 42, 154–156 Psychotria, 75, 119, 397, 398, 400–402
POACEAE, 29, 37, 38, 373, 468, 473 Psydrax, 119
Podandrogyne, 188 Pterocarpus, 112, 114, 297, 298, 301
PODOCARPACEAE, 29, 37, 38, 375 Pterocarya, 275
Podocarpus, 38, 114, 375, 376 Pterocymbium, 105, 318
Poecilandra, 58, 349–351 Pterospermum, 44, 104, 313, 314, 318
Poecilanthe, 112, 115 Pterygota, 314
Poeppigia, 112 Ptychopyxis, 97, 241, 247
Pogonophora, 98 Putranjiva, 54, 96, 387
Pogostemon, 278 PUTRANJIVACEAE, 30, 31, 46, 49, 50, 53, 54, 95, 96,
POLEMONIACEAE, 137 386, 474
Polyalthia, 50, 151, 153 Pycnandra, 421
Polygala, 377 Pycnanthus, 41, 52, 337, 338
POLYGALACEAE, 30, 51, 121, 126, 376, 377, 474 Pycnocoma, 97
POLYGONACEAE, 30, 31, 43, 55, 125–127, 378, 379, Pyrenacantha, 126, 127, 130, 270, 271
466, 475 Pyrola, 234
Polylepis, 395, 396, 466 Pyrostegia, 173
Polyscias, 89, 160, 161 Pyrus, 395
Polyspora, 434, 435
Pometia, 416, 417, 419
Poraqueiba, 269–271 Q
Porophyllum, 81 Qualea, 75, 451–453
Posoqueria, 119, 401 Quararibea, 104, 313
Potalia, 254, 255 Quassia, 51, 107, 109, 424–426
Pothomorphe, 371, 372 Quercus, 58, 249–252, 415
Poulsenia, 331, 333 Quiina, 76, 349–351
Pourouma, 39, 331, 441, 443, 444 Quisqualis, 206
Pouteria, 41, 421, 423
Pradosia, 42, 66, 421
Prainea, 332–334 R
Premna, 80, 86, 278 Radermachera, 173, 176
Prestonia, 155 Rafflesia, 450
Primula, 381–383 Randia, 119, 398, 401
PRIMULACEAE, 30, 32, 33, 51, 64, 65, 69, 70, 380, 382, RANUNCULACEAE, 121, 136, 197, 467, 474
383, 466, 475 Raphia, 163
Prionostemma, 42, 123, 194, 196 Raputia, 92
Pritchardia, 164 Rauvolfia, 43, 155, 157
Prockia, 45, 410, 411, 413 Rauwenhoffia, 124
Protea, 384 Reissantia, 77
PROTEACEAE, 3, 33, 35, 36, 67, 84, 86, 89, 90, 384, Retrophyllum, 375
385, 468, 473 Reynosia, 392
Protium, 41, 107, 181–183 Rhabdadenia, 157
Protomegabaria, 96 RHABDODENDRACEAE, 32, 64, 65, 475
Protorhus, 43 Rhabdodendron, 65, 389
Prumnopitys, 375 Rhacoma, 195, 196
Index 491
RHAMNACEAE, 30, 31, 34, 44–47, 49, 58, 60, 74–76, Salacia, 123, 134, 194–196
121, 126, 128, 131, 134, 390, 450, 474 SALICACEAE, 7, 30–32, 44–50, 53, 54, 58, 59, 64, 71,
Rhamnus, 391 73, 410–412, 465, 474
Rhaphiostylis, 128, 270 Salix, 410, 411, 413
Rhapis, 164 Salvertia, 452
Rheum, 379 Samadera, 51, 107, 424
Rhigozum, 174 Samanea, 18, 116, 287, 294
Rhipsalis, 124, 185, 186 Sambucus, 92
Rhizophora, 75, 392–394 Sandoricum, 109, 324–326
RHIZOPHORACEAE, 34, 74–77, 392, 394, 467, 474 Sanguisorba, 396, 466
Rhodamnia, 80 SANTALACEAE, 31, 33, 35, 51, 52, 71, 72, 84, 85, 356,
Rhododendron, 70, 234–236 414, 415, 475
Rhodoleia, 60, 261, 262 Santalum, 85, 414, 415
Rhoicissus, 137 Santiria, 182
Rhus, 108, 109, 147, 148 SAPINDACEAE, 7, 32, 35, 36, 64, 65, 80, 86, 87, 90–92,
Rhynchanthera, 322 106, 108–110, 121, 124, 137, 416, 417, 475
Rhyticaryum, 271 Sapindus, 416–419
Richeria, 57, 96, 365, 367, 368 Sapium, 102, 241, 247
Ricinodendron, 88, 100, 242 SAPOTACEAE, 29, 32, 39–42, 65, 66, 202, 420–422,
Ricinus, 99, 242 465, 475
Rinorea, 48, 60, 76, 77, 447, 448 Saraca, 287, 290
RIPOGONACEAE, 133, 473 Sararanga, 360
Ripogonum, 133 Sarcandra, 74, 197
Ritchiea, 188 Sarcolaena, 44
Rivina, 369, 370 SARCOLAENACEAE, 30, 43, 44, 475
Rochefortia, 63, 178–180 Sarcomelicope, 107
Rollinia, 153, 154 Sarcopteryx, 416
Rondeletia, 398 Sarcostigma, 126, 269–271
ROSACEAE, 30, 31, 35, 46, 51, 56, 86, 87, 395, 466, 474 Sarcotheca, 68, 90, 357, 358
Rotala, 306 Satyria, 128, 234, 237
Rothmannia, 402 Saurauia, 66, 69, 145, 146
Roucheria, 59, 302 Sauropus, 95, 365–367
Roupala, 67, 90, 384–386 Sauvagesia, 350
Rourea, 209–212 Savia, 48, 96
Roystonea, 38 Scaevola, 66, 70, 258
Rubia, 400 Scaphopetalum, 9, 104, 313, 314, 318
RUBIACEAE, 34, 74, 75, 119, 121, 132, 134, 372, 397, Scheelea, 167
400, 476 Schefflera, 90, 159–162
Rubus, 395, 396 Schima, 71, 434, 435
Ruellia, 141, 143 Schinopsis, 147
Ruizterania, 83, 451–453 Schinus, 108, 148, 149
Rumex, 379 Schisandra, 125, 424
Ruprechtia, 43, 378 SCHISANDRACEAE, 32, 64, 65, 125, 424, 473
RUTACEAE, 32, 35, 36, 62, 64–66, 81, 86, 87, 89–92, Schizolobium, 118, 288
106, 107, 121, 138, 403, 406, 475 Schizostemon, 266
Ryania, 47, 60 Schlegelia, 135
Ryparosa, 68, 143–145 SCHLEGELIACEAE, 135
Schleichera, 109, 416, 417, 419
Schoepfia, 53, 351
S SCHOEPFIACEAE, 55, 351, 475
Sabal, 164, 167 Schoutenia, 104
Sabia, 73, 126, 409, 410 Schrebera, 85, 92, 353
SABIACEAE, 33, 36, 67, 71, 73, 89, 90, 121, 126, 409, Schumanniophyton, 400
410, 473 Schuurmansia, 349
Sabicea, 134, 401 Sciadodendron, 89
Saccifolium, 254 Sclerocarya, 148
Sacoglottis, 49, 54, 55, 265, 266 Scleropyrum, 414, 415
Sagotia, 100 Scolopia, 54, 64, 73, 410
Salacca, 163, 166
492 Index
Scorodocarpus, 237 Steganotaenia, 88

SCROPHULARIACEAE, 35, 80, 476 Stelechocarpus, 151, 153
Scutia, 76, 134, 390, 391 STEMONURACEAE, 53, 54, 476
Scyphiphora, 119 Stemonurus, 53
Scytopetalum, 54, 284, 285 Stenocarpus, 385, 386
Sebastiania, 40, 99 Stenomeris, 224, 225
Securidaca, 51, 126, 376, 390 Stephanopodium, 221
Securinega, 95 Sterculia, 105, 313, 314, 319
Seguieria, 128, 369, 370 Stereospermum, 174
Selenicereus, 186 Sterigmapetalum, 76, 392, 394
Semiramisia, 62 Steriphoma, 58, 68, 188
Senefeldera, 103 Stewartia, 435
Senefelderopsis, 247 Stifftia, 170
Senegalia, 291, 294 Stigmaphyllon, 75, 132, 311, 312
Senna, 112, 116, 286, 287, 291 Streblus, 40, 332
Serianthes, 118 Strephonema, 64
Sericolea, 76, 231, 232 Strobilanthes, 142
Serjania, 137, 416, 419 Strombosia, 54, 237, 238
Sesbania, 113, 115, 297, 301 Strongylodon, 301
Sherbournia, 132 Strophanthus, 155
Shirakiopsis, 242, 247 Strophioblachia, 62, 101
Shorea, 46–48, 60, 225–228 Strumpfia, 402
Sida, 314 Strychnos, 77, 133–135, 303–305
Sideroxylon, 41, 420, 421, 423 Stryphnodendron, 117, 292, 296
Simaba, 109, 424, 425 Styloceras, 184
Simarouba, 109, 110, 181, 182, 425, 426 STYRACACEAE, 31, 32, 51, 52, 64, 430, 475
SIMAROUBACEAE, 30, 32, 33, 35, 51, 62, 63, 69, 70, Styrax, 52, 64, 430
86, 87, 89, 106, 107, 109, 110, 424, 425, 475 Sumbaviopsis, 97
Simira, 75 Suregada, 95, 96
Siparuna, 82, 86, 329–331 Suriana, 63, 430
Siphoneugena, 341 SURIANACEAE, 32, 62, 63, 430, 474
Siphonodon, 55, 195, 196 Swartzia, 111, 112, 286, 297, 301
Skimmia, 403 Swietenia, 111, 324, 325, 327
Sloanea, 57, 68, 231–233 Swintonia, 147–149
SMILACACEAE, 121, 131, 426, 473 Syagrus, 164
Smilax, 131, 426, 427 Symbolanthus, 254, 256
Socratea, 166, 167 Symphonia, 42, 43, 202, 204
SOLANACEAE, 30–33, 51, 52, 62, 63, 71, 73, 121, 127, SYMPLOCACEAE, 33, 71–73, 201, 430–432, 435, 475
129, 427, 428, 476 Symplocos, 72, 73, 431
Solandra, 129 Synadenium, 103
Solanum, 63, 73, 127, 427–429 Syncarpia, 43
Sonchus, 168 Synsepalum, 421
Sonerila, 321, 322, 324 Syntriandrium, 328
Sonneratia, 78, 305, 306, 308 Syringa, 354, 355
Sophora, 115, 297 Syzygium, 81, 83, 339–342
Sorindeia, 108, 109, 147, 148
Sorocea, 40, 332
Soulamea, 70 T
Spachea, 42, 310–312 Tabebuia, 84, 92, 174, 176, 370
Sparattanthelium, 130, 131, 262–264 Tabernaemontana, 43, 154–157
Spathelia, 403 Tachia, 78, 254, 256
Spathodea, 92, 173 Tachigali, 115, 287
Spermacoce, 400 Takhtajania, 65
Spigelia, 304, 305 Talauma, 309
Spondianthus, 64, 97, 241 Talisia, 419
Spondias, 109, 147–149 Tamarindus, 287, 288, 291
Stachytarpheta, 445 Tambourissa, 330
Stapelia, 155 Tamus, 224
STAPHYLEACEAE, 35, 36, 86, 87, 91, 475 Tannodia, 242
Index 493
Tapeinosperma, 65, 382 Touroulia, 91

Tapirira, 108, 147 Tovomita, 79, 202
Tapura, 46, 48, 221, 222 Tragia, 98, 242
Taralea, 91, 114 Trapa, 317, 306
Tarenna, 119, 400 Trattinnickia, 182
Tasmannia, 187 Treculia, 39
Tecoma, 92, 173 Trema, 45, 439, 440
Tectona, 80, 277–279, 445 Trevesia, 161
Teijsmanniodendron, 84, 92 Tribulus, 455, 457
Tephrosia, 115 Trichilia, 107–111, 324, 326, 327
Terminalia, 18, 63, 64, 66, 81, 83, 205, 206, 209 Trichoscypha, 108
Ternstroemia, 65, 70, 72, 363–365 Trichostigma, 369
Tetracera, 125, 222–224 Trigonia, 75, 132
Tetractomia, 107, 403 TRIGONIACEAE, 34, 48, 74, 75, 121, 132, 474
Tetragastris, 107, 108, 181–183 Trigoniastrum, 48
TETRAMELACEAE, 3, 32, 61, 62, 474 Trigonobalanus, 249, 250
Tetrameles, 61, 62, 433, 434 Trigonostemon, 100, 247
Tetrameranthus, 151 Trimenia, 81
TETRAMERISTACEAE, 70, 475 TRIMENIACEAE, 35, 81, 473
Tetrapanax, 160 Triplaris, 43, 378–380
Tetrapleura, 117 Triplochiton, 44, 104, 315
Tetrapterys, 85, 311 Tristania, 64
Tetrastigma, 88, 131, 137, 448–451 Triumfetta, 104
Tetrorchidium, 77, 102 Trophis, 331, 332, 335
Thea, 435 Turnera, 64, 362, 363
THEACEAE, 31, 33, 51, 53, 69, 71, 72, 145, 201, 434, Turpinia, 91, 475
435, 475 Turraea, 52, 324, 325, 327
Thecacoris, 95 Tyleria, 350
Theobroma, 9, 17, 102, 313–315, 319
Theophrasta, 70, 381, 382
Thesium, 415 U
Thespesia, 315, 319 Uapaca, 40, 42, 69, 97, 365, 367, 368
Thevetia, 155 ULMACEAE, 8, 9, 30, 43–46, 48, 74, 439, 474
Thiloa, 82 Ulmus, 439, 440
Thoracocarpus, 124, 219, 220 Uncaria, 134, 372, 397, 398, 400
Thottea, 167 Unonopsis, 151, 153
Thouinia, 416 Uraria, 297, 300
Thryallis, 311 Urena, 314
Thunbergia, 132, 141, 142 Urera, 441, 444
Thymelaea, 437 Urtica, 441
THYMELAEACEAE, 9, 30, 33, 35, 50, 67, 84, 121, 129, URTICACEAE, 29–31, 34, 36, 39, 40, 44, 55, 56, 74, 87,
134, 435, 437, 467 88, 331, 441, 444, 474
Thyrsodium, 108, 109, 147 Uvaria, 124, 151, 153
Tibouchina, 322 Uvariopsis, 152
Tieghemella, 421
Tilesia, 170
Tiliacora, 328 V
Timonius, 119 Vaccinium, 53, 234–237
Tinospora, 329 Vachellia, 291, 293, 296
Tipuana, 114, 297, 302 Vallea, 232
Tococa, 321, 322, 324 Vantanea, 54, 55, 265, 266
Toddalia, 65, 106, 107, 138 Vasconcellea, 191
Toechima, 109, 110 Vatairea, 114
Tontelea, 196 Vateria, 225, 226
Toona, 111, 324, 325, 327 Vatica, 39, 46, 56
Topobea, 322 Vaupesia, 100
TORRICELLIACEAE, 67, 68, 437, 476 Vavaea, 66, 107, 324, 325
Toulicia, 109, 420 Veitchia, 38, 167
Tournefortia, 180 Ventilago, 45, 126, 128, 390–392
494 Index
Vepris, 65, 90, 106, 403 Wollemia, 162, 163

VERBENACEAE, 32, 34, 35, 64, 65, 77, 79–83, 85, 121, Woodfordia, 79
133, 277, 278, 354, 445, 463, 476 Wrightia, 154, 155
Verbesina, 79, 168, 170 Wulffia, 168, 170
Vernonia, 170
Viburnum, 77, 79
Vigna, 297 X
Viguiera, 168, 169 Xanthium, 169, 170
Viola, 447, 448 Xanthoceras, 417
VIOLACEAE, 8, 30, 31, 34, 46, 48–50, 58, 60, 75–77, Xanthophyllum, 51, 230, 376, 377
121, 125, 242, 411, 447, 462, 474 Xenostegia, 212, 214
Virola, 41, 52, 337 Xerospermum, 416, 420
Viscum, 414, 415 Ximenia, 62, 237, 351, 352
Vismia, 42, 43, 79, 202–205 Xylia, 118, 292, 293, 296
VITACEAE, 30, 31, 35, 36, 43–45, 55, 86–88, 121, 131, Xylocarpus, 89, 111, 324, 325
137, 363, 448, 450, 451, 461, 474 Xylopia, 50, 151–153
Vitellaria, 421, 422 Xylosma, 50, 179, 410, 413
Vitex, 92, 174, 277–279 Xymalos, 82, 330, 331
Vitis, 92, 174, 277–279
Vochysia, 75, 451–453
VOCHYSIACEAE, 34, 35, 75, 83, 451–453, 462, 465, Y
475 Yucca, 37
Vouacapoua, 115, 286, 291
Voyria, 254, 256
Voyriella, 254 Z
Zamia, 39, 218, 219
ZAMIACEAE, 29, 37, 39, 218
W Zanthoxylum, 87, 89, 106, 403, 406
Walsura, 107 Zapoteca, 113
Waltheria, 314 Zelkova, 439, 440
Warburgia, 72, 186, 187 Zinowiewia, 76, 195
Wedelia, 168 Ziziphus, 8, 45, 390–392
Weinmannia, 74, 91, 217 Zollernia, 50, 112, 297, 302
Wendlandia, 400 Zuccarinia, 119
Wigandia, 178–180 Zygia, 117, 118, 291, 295, 296
Wikstroemia, 437 Zygogynum, 65
WINTERACEAE, 32, 64, 65, 186, 473 ZYGOPHYLLACEAE, 36, 89, 91, 455, 456, 461, 474
Witheringia, 428

A Field Guide To Tropical Plant Families (Roland Keller)

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A Field Guide To Tropical Plant Families (Roland Keller)

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Roland Keller

A Field Guide to Tropical

ISBN 978-3-031-05941-4 ISBN 978-3-031-05942-1 (eBook)

Botanists, formerly (and with differing motives), collected, described and

requiring publication of papers of theoretical interest, in ‘high-impact’ jour-

As an ecologist, I like keys for determination of plants according to vege-

My humble thanks to David Mabberley for his flamboyant preface; his

Part I User Manual and Architectural Concepts

Part II Identiﬁcation Keys

4 Key to the Groups of Families of Trees and Shrubs . . . . . . . . 29

Part III Description of the Families

9 Acanthaceae to Asteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

12 Dichapetalaceae to Dipterocarpaceae . . . . . . . . . . . . . . . . . . . . 221

19.9 Loganiaceae + Gelsemiaceae . . . . . . . . . . . . . . . . . . . . . . 303

23.16 Primulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380

30 Herbaceous Versus Arborescent Habit in the Families

List of Identiﬁed Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473

Roland Keller is an independent botanist living in Switzerland. He was

1.1 Scope of the Book

1.2 Why Consider Families?

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 3

1.3 Photos or Drawings?

1.4 Sources and Tests

1.5 Structure of the Book

The book consists of three parts:

Explanations relative to keys

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 7

a Spiral leaf arrangement. Leaves simple.

Thus items a and b, each with two propositions, are complementary.

Explanations relative to the descriptions of the families

3.1 Concepts and Statements

Fig. 3.1 Basal (b) and distal

Monopodial versus sympodial

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 13

Fig. 3.3 Rhythmic and

Branching (or ramiﬁcation), axillary buds and prophylls

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

Immediate or delayed branching

Fig. 3.5 Immediate

Ramiﬁcation (i.e. branching) and reiteration

Fig. 3.6 (a) Branching,

Fig. 3.7 Orthotropism, plagiotropism and plagiotropism by apposition growth

Reorientation of a main axis

Fig. 3.9 Orientation of

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

Preliminary remarks to architectural models

Nevertheless, a pagoda tree (Terminalia catappa), a mangosteen (Garcinia mangostana) and a

3.2 Key to the Architectural Models

1b Trunk polycaulous (plant appearing branched at ﬁrst sight). 3

See also groups 53 to 57 if lianescent in habit. Unillustrated families written in italics.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 29

Group 5: CHRYSOBALANACEAE, DIPTEROCARPACEAE, HAMAMELIDACEAE, IRVIN-

10a Leaf venation palmate or tripliveined (Fig. 4.2a).

Group 12: ARISTOLOCHIACEAE, CONNARACEAE, EBENACEAE, ERICACEAE, ERY-

Group 13: ANISOPHYLLEACEAE, CANELLACEAE, CELASTRACEAE, CHRYSOBALANA-

7b All stems with spiral leaf arrangement (Fig. 4.3a). 16

18b Inner bark different. 19

Group 19: ARALIACEAE, EUPHORBIACEAE, HAMAMELIDACEAE, MENISPERMACEAE

Group 21: ASTERACEAE, BIGNONIACEAE, BORAGINACEAE, CACTACEAE, CAPPAR-

Group 25: BORAGINACEAE, ICACINACEAE, LECYTHIDACEAE, PROTEACEAE, SABIA-

31a Branches with growth strongly rhythmic denoted by shorter

Group 31: CANNABACEAE, EUPHORBIACEAE, RHAMNACEAE, URTICACEAE

38a Nodes or base of internodes swollen (Fig. 4.1a).