The Flowering Plants Handbook

Abstract

This plant book aims to help identify flowering plants to genus and family level anywhere in the world. In 2014 there are very few available works which are both comprehensive and up-to-date for all the flowering plants families and genera of the world. The Flowering Plants Handbook is an easy to use identification guide to the worlds flowering plants designed for both specialists and non-specialists and from beginner to expert. The book contains descriptions of all currently recognised flowering plant families, morphological
notes for 6656 genera (all current genera for 398/413 families) and over 3000 images and illustrations. Flowering plants can be identified using the book to family and much of the world's generic diversity in four 'easy' steps. Some plants will be identified correctly quickly, whilst others may require some retracing of steps and take a little more time. The advantage of this book is that it helps the user learn about the classification system and plant diversity during the identification process. This work was compiled and developed using the living, library and herbarium collections at the University of Aberdeen, Royal Botanic Gardens, Edinburgh and Royal Botanic Gardens, Kew.

Full text

Published by Plant Gateway Ltd., Herord, SG13 7BX, United Kingdom
© Plant Gateway 2014
This book is in copyright. Subject to statutory excepon and to the provision of relevant col-
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ISBN 978-0-9929993-0-8
eISBN 978-0-9929993-1-5
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or third-party internet websites referred to in this book, and does not guarantee that any
content on such websites is, or will remain, accurate or appropriate.
Addional informaon on the book can be found at: www.plantgateway.com
An appropriate citaon for this eBook is:
Byng JW. 2014. The Flowering Plants Handbook: A praccal guide to families and genera of
the world. Plant Gateway Ltd., Herord, UK. eBook available from: www.plantgateway.com

From the war of nature, from famine and death, the most exalted object which we are capable of
conceiving, namely, the producon of the higher animals, directly follows. There is grandeur in this
view of life, with its several powers, having been originally breathed into a few forms or into one;
and that, whilst this planet has gone cycling on according to the xed law of gravity, from so simple
a beginning endless forms most beauful and most wonderful have been, and are being, evolved.
Charles Darwin On The Origin of Species (1859)
CONTENTS

The Flowering Plants Handbook


CONTENTS


DEDICATION
This work is a dwarf standing on the shoulders of giants and is dedicated to the many botanists, both
past and present, for the huge body of knowledge that exists today. Parcular reference is made to the
vision and works of the Angiosperm Phylogeny Group members (Kåre and Birgia Bremer, Mark Chase,
Mike Fay, James Reveal, Doug and Pam Sols and Peter Stevens); the inspiraonal morphological works
by Al Gentry, John Hutchinson, Armen Takhtajan, Cornelis Gysbert Gerrit Jan van Steenis and Dick
Brummi; and nally the great collectors whose many herbarium specimens I viewed at Kew, notably
Terry Pennington, Ghillean Prance, Marn Cheek, Mark Coode and John Dranseld, whose eldwork
makes everything else possible.
CONTENTS

PREFACE
This book was a complete accident and grew to become far more comprehensive than I ever intended.
It came about due to my desire to understand more about the plant diversity of the world, parcularly
during eldwork for my PhD and several surveys in Zambia. Another reason for compiling such a book is
there are no really praccal eld guides or handbooks on the market to both families and genera which
are comprehensive, not to menon comprehensive for teaching and learning. Notable excepons are
the Generic Tree Flora of Madagascar (Schatz, 2001) and A Field Guide to the Families and Genera of
Woody Plants of Northwest South America (Gentry, 1993) but both are geographically restricted and
slightly out-of-date. I persevered adding and dying notes I generated teaching botany at the University
of Aberdeen over several summers and this work is the result of many days teaching both specialists
and non-specialists how to idenfy plants, much of which I was teaching myself as I was going along.
This is the rst edion and comments are appreciated to improve future versions.
Idenfying plants is extremely important and probably understated at universies these days. Botany
and plant taxonomy is very much thought of as being done in the past and cells and biotechnology
is what we should be doing now. But plant names and understanding their classicaon is sll of
profound importance because taxonomy is the baseline data for all science. Once you have a name
of a species all the informaon associated with it (i.e. its uses, its distribuon, its conservaon status
and threats) becomes accessible. The classicaon system acts as a retrieval system for idencaon.
However, recent changes following molecular work mean that it is oen confusing for many users what
the classicaon system (parcularly at the generic level) actually looks like and how to idenfy plants
to it because the characters needed are scaered across hundreds of papers, books and websites.
In addion, names are constantly changing as new informaon comes to light and hundreds of new
species are described each year. We sll know we know so lile about most of the worlds species
and what is parcularly alarming is that many species severely threatened with exncon due to
deforestaon and other anthropogenic acvies. There is a strong possibility that many plant species
will disappear before we have even had a chance to nd out the most basic thing about them so
idenfying plants correctly is a necessity rather than a luxury.
This book aims to make possible the idencaon to genus and family of all owering plants of the
world or at least guide the user from everything with a ower and (usually) green to fewer potenal
groups. At the family level, this means following the currently accepted Angiosperm Phylogeny (APG)
III classicaon (strictly) which is now widely accepted. At the generic level, I have aempted to
synthesise currently accepted names from the literature, oen taking into account future changes.
No new combinaons nor aempts to reclassify any families are made in this rst edion. It is solely a
synthesis of the literature. There is no similar single-body of knowledge on the worlds ora available on
the web or in the bookshops which is praccal, aimed at both specialists and non-specialists, mobile,
easy to use, comprehensive, up-to-date, illustrated and with global coverage.
I must state that work of this kind is oen dicult to compile due to exisng errors in published
literature, the large quanty of exisng literature, somemes diering opinions on the taxonomy (so
called ‘lumping’ and spling’) and nally the simple fact that there are so many missing gaps in our
body of knowledge. Also, it must be stated that the preparaon and publicaon of this work has not
received any grants or nancial aid from any instuon or grant giving body for its compilaon but I
raised funds for it through my own extra-curricular plant idencaon courses run in various botanical
gardens in the UK, consultancy work and pre-sales of the book through Plant Gateway Ltd. I must
thank in parcular individuals who ordered pre-order books for showing faith in the work, as well as
numerous instuons who purchased copies for their libraries.
This book will be annually updated and this rst version contains descripons to all currently accepted
families and so far morphological notes for 6656 genera (complete for 398/413 families). I am sll not
CONTENTS

happy with several family treatments and generic synopses and these will be improved and added
to in future edions. The second version is ancipated to include a further completed nine families
(Acanthaceae, Amaranthaceae, Apocynaceae, Aracaceae, Brassicaceae, Euphorbiaceae, Lamiaceae,
Malvaceae and Melastomataceae), improving and correcng the current content and adding excing
new technological features. I would like this rst edion to encourage an open, collaborave process
where feedback and discussion shape future edions.
The hundreds of plant names and confusing botanical terminology are oen seen as factors that put
people o from idenfying plants or at least from looking at them and the subject more closely. But it’s
not about how many names you can remember, its just knowing where to look for that informaon.
This book aempts to use simple plain English where possible and if technical terms are unavoidable
they are included in the glossary at the back. I have avoided using too many keys (and abbreviaons)
as these are not always popular but I have included some to provide enough structure to allow
idencaon (and back track during idencaon where needed), as well as, to help learn about the
classicaon system at the same me.
By making things simple the intenon is to encourage more non-specialists to start looking and
idenfying plants without undermining their opportunity to develop advanced knowledge. This is
because I rmly believe that by enabling others to understand the diversity that exists can ulmately
help reduce its decline. I hope this is the beginning of my contribuon.
Happy botanising!
James W. Byng BSc (Hons) MSc PhD (j.byng@plantgateway.com)
London, October 2014
CONTENTS


This book aims to guide the user from many owering plant families and genera to one (or few) in four ‘easy’ steps.
The more botanical experience the user has the more likely some steps will be passed over, so non-specialists should
begin with steps 1 or 2 and specialists should begin with steps 3 or 4.
For disncve habits, such as parasic, carnivorous, grass-like or bulbous plants, and/or morphological characters,
such as the absence of the perianth or plants with exudates (sap, latex or resins), then rst see the list of useful
diagnosc characters.

Each major group is divided into orders. Ideally one order
matches the plant but somemes two or three do, so
explore each.... Again pracce and familiarity will help
over me. Please note that variable orders should always
be checked!
      

Morphological notes and images for choosing which major
group are laid out on pages 4‒6. These are broad groups which
are aimed primarily for the beginner to have an idea where to
start. Over me familiarity and pracce will help choose the
correct one.
       


 

The keys try to include all species for each family but working
at such a high taxonomic level results in some extremely rare or
unusual species perhaps being dicult to key out. When using
long keys do check out a few families in the relevant group to
get a feel for them.
Read and compare generic notes, deliming along the
way. If two or more genera are delimited use local oras
and eld guides. Note deliming genera at a global scale is
oen dicult and idencaon oen only becomes easier
at a regional and/or local scale so geography can be very
useful.
CONTENTS


CONTENTS
................................................................................... 1
 ...................................................... 4
 ……………….........…....…………… 7
....................................................................... 10
Amborellales……………….............................................................. 11
Nymphaeales………………………..................................................... 12
Austrobaileyales………................................................................. 14
Chloranthales............................................................................. 16
Canellales……………………………………………………….........…....……… 17
Piperales………………………………………………………............…………….. 19
Magnoliales………………………………………………….........…….…………. 22
Laurales………………………………………………………….........………..……. 30
.......................................................................................... 38
Acorales………………………............................................................. 39
Alismatales……………………………................................................... 40
Petrosaviales……………................................................................. 53
Dioscoreales............................................................................... 54
Pandanales………………………………………………………...........………….. 57
Liliales……………………………………………………………..….........…………. 61
Asparagales…………………............................................................. 70
Arecales……….……………………….................................................... 94
Commelinales……………............................................................... 96
Zingiberales................................................................................ 101
Dasypogonaceae………………………………………..……….........…………. 109
Poales..……………………………………………………….……….........………… 110
 ............................................................................... 130
Ceratophyllales……………............................................................. 131
Ranunculales………………………..................................................... 132
Sabiaceae…………………................................................................ 143
Proteales.................................................................................... 144
Trochodendrales…………………………………………………...........………. 149
Buxales………………………………………………………………..........………… 150
Gunnerales………………….............................................................. 152
Dilleniaceae………………………….................................................... 153
ROSIDS ……….................……………………………........................................... 154
Saxifragales…………………............................................................. 156
Vitales…….…………………………….................................................... 167
Zygophyllales..…………................................................................. 168
Fabales....................................................................................... 170
Rosales………………………………………………………………...........………… 174

Fagales.……………………………………………………………….........…..…….. 189
Cucurbitales…………………............................................................ 193
Celastrales…………………………….................................................... 201
Oxalidales…………………................................................................ 206
Malpighiales…............................................................................ 212
Geraniales.………………………………………………………..…….........…….. 249
Myrtales…………………………………………………….………….........………. 251
Crossosomatales…………............................................................. 265
Picramniales………………………….................................................... 269
Sapindales……..…………................................................................ 270
Huerteales.................................................................................. 294
Malvales………………………………………………………..………….........…… 296
Brassicales…………………………………………………..…………….........…… 310
 ....................................................................... 324
Berberidopsidales……….............................................................. 325
Santalales.…………………………….................................................... 326
Caryophyllales…………................................................................. 337
ASTERIDS................................................................................................. 380
Cornales………………………............................................................ 382
Ericales…..…………………………….................................................... 387
Unplaced Basal Lamiid Families……………………………..........………. 414
Garryales……….…………................................................................ 417
Genanales................................................................................ 418
Vahliaceae…………………………………………………………...........………… 429
Boraginaceae………………………………………………………...........………. 429
Solanales..……………………............................................................ 437
Lamiales………………………………..................................................... 446
Aquifoliales………………................................................................ 489
Asterales..................................................................................... 492
Escalloniales……………………………………..………………….........………… 504
Bruniales…………………………………………….…………………..........……… 505
Paracryphiales…………….............................................................. 507
Dipsacales…………………………….................................................... 508
Apiales………….…………................................................................. 512
................................................................................. 520
 ......................................................................... 537
 ……….…..........……………...................................................... 539
............. 607
CONTENTS

1

For over 250 years owering plants were arranged in various classicaon systems according to their morphology
and biochemistry. Botanists and biologists from around the world supported and used dierent systems (e.g. de
Candolle, Lindley, Bentham & Hooker, Engler, Hutchinson, Takhtajan, Cronquist, Dahlgren) which caused some
confusion and even more debate. In the last twenty years the classicaon of plants has changed dramacally with
the accumulaon of molecular data that is unravelling the true evoluonary relaonships of plants.
Plants have adapted over millions of years to dierent pollinaon syndromes and/or challenging environments (e.g.
arid, saline) and this has resulted in some divergence of closely related groups and convergence of distantly related
ones. A classicaon system based on evoluonary relaonships has been the aim since Darwin and using DNA
sequences reects genealogy far more accurately than morphological characters alone.
The Angiosperm Phylogeny Group (APG) classicaon is a modern, mostly molecular-based system and is now
widely accepted. The rst version was published in 1998, this was superseded in 2003 by the APG II system and
the current APG III system was published in 2009. In the APG era this work is the rst and only to comprehensively
compile praccal morphological characters of all APG III families and their genera in the world based on the current
literature for idencaon.

The owering plant families are numerically arranged following the Linear Angiosperm Phylogeny Group (LAPG)
III (Haston et al., 2009). The generic synopsis of each family is arranged alphabecally, and if there are many
genera grouped, then also taxonomically (by subfamilies or tribes), geographically (by connent or region) and/or
morphologically. The formal family descripons contain important characters which are usually easily comparable
with each other and useful diagnosc characters are underlined. Morphological synapomorphies, as well as, general
characters to help compare similar genera are aempted for each genus listed. All characters used can be seen by
the naked eye or with a hand lens, though looking at placentaon and ovule number can somemes be challenging.

These formulas aid idencaon and can be useful for nong the number of parts to each whorl. There are several
ways of doing oral formulas with a very general one used in this work to allow its use by many users. The formulas
consist of four primary symbols (plus some addional secondary symbols) which go from le to right:
Primary: K number of sepals - with ‘K’ represenng “calyx”. K5 = calyx of 5 sepals.
C number of petals - with ‘C’ represenng “corolla”. C5 = corolla of 5 petals.
A number of stamens - with ‘A’ represenng “androecium”.
G number of carpels - with ‘G’ represenng “gynoecium”. G indicates a superior and  an inferior ovary.
Secondary: (....) brackets are used here to indicate less oen numerical values NOT fusion
4‒5 dashes between numbers are used here to show variaon in each whorl
P the perianth (i.e. calyx and corolla) and is used when the two whorls are indisnguishable from each other
+ symbol is used to indicate dierenaon within each whorl. For example, two large petals and
three small petals would be: C2+3
° symbol is used for sterile stamens (= staminodes) or carpels (= pisllodes)

Some families and genera have now been incorporated into other families and/or genera. A comprehensive list of
family synonyms is provided based primarily on the paper by Reveal & Chase (2011). Recent or relevant generic
synonymy, based on current literature, are included in each of the family treatments except for where extensive
changes have taken place and appropriate websites are cited for further informaon.

The literature consulted for each family is given at the end of each treatment. General taxonomic works by Stevens
(2001-), Mabberley (2008) and Takhtajan (2009) were consulted for all or most family treatments and are not cited
for each treatment to avoid repeon but as sources of informaon they are giants and essenal reference works.
The 15 largest families are briey treated in this rst version so the literature cited is therefore oen minimal.
CONTENTS

2

Images and illustraons of families and genera are included to aid idencaon. Some images are of high quality
and some of poorer quality. In the laer case this is oen due to the scarcity of images for some rare groups and
it is hoped beer quality usable images will become available for future versions. Illustraons/images of technical
terms and/or ‘spot’ characters specic to certain families are also included for around a h of all families in grey
boxes. Herbarium specimens at RBG Kew and RBG Edinburgh are shown where images of living plants were not
available. The copyright of all images or illustraons in this work belong to the author unless otherwise stated.
Images obtained from wikimedia commons are somemes in the public domain (and variously noted with PD) or
have the following licenses associated with each image:
CC0 1.0 CC BY 2.0 CC BY-SA 2.1 ES CC BY-SA 2.5 CC BY 2.5 AU CC BY-SA 3.0 CC BY 4.0

Where possible botanical terminology is kept to a minimum and a basic glossary is included at the back of the
book for any terms used, as well as, common terms which are frequently used in other botanical literature. Also,
illustraons of essenal and oen complex terms are included to aid the non-specialists. For a more comprehensive
botanical glossary see The Kew Plant Glossary (Beentje, 2010).

Connental and country levels have generally been favoured in this work with provincial, state or regional (if
applicable) data in square brackets or vice versa. Maps with botanically interesng islands and/or island groups
are provided in the Geographical Maps secon along with maps of the tropical and subtropical regions. A standard
atlas should help with most distribuons but the following geographical terms are oen used and described below
to aid the reader:
Hispaniola ‒ island in the Caribbean comprised of Hai and the Dominican Republic.
 ‒ several island groups o the coast of Africa and Europe: Canary Islands [Spain], the Azores and Madeira [Portugal]
and Cape Verde Islands. See MAP D.
 ‒ region of southeast Asia comprised of Malaysia, Indonesia, Brunei, the Philippines and Papua New Guinea. See MAP C.
 ‒ group of islands in the Indian Ocean comprised of La Réunion [France], Maurius and Rodrigues [Maurius]. See
MAP D.
 ‒ region of Australasia comprised of Papua New Guinea, Irian Jaya [Indonesia], Solomon Islands, Vanuatu, Fiji and New
Caledonia [France]. See MAP D.
 ‒ region of Australasia comprised of the Federated States of Micronesia, Kiriba, Marshall Islands, Nauru, Palau, North-
ern Mariana Islands [USA], and Guam [USA].
Neotropics ‒ tropical America (i.e. tropical areas of North, Central and South America).
 ‒ the connents of North America and South America combined.
 ‒ the connents of Europe, Asia, Africa and [in this work] Australasia.
 ‒ tropical regions of the Old World.
Pantropical ‒ across tropical regions (i.e. in both the Neotropics and Old World tropics).
 ‒ triangular region of Australasia from Hawaii [USA] to New Zealand and Rapa Nui [Chile].
 ‒ region of the earth from the Tropics of Cancer/Capricorn to around the 38th parallel. See MAP B.
Tropics ‒ region of the Earth around the equator from the Tropic of Cancer to the Tropic of Capricorn. See MAP A.

Few abbreviaons are included in the text to make it easier for the user to read. Apart from abbreviaons associated
with oral formulas, the following abbreviaons and symbols are used:
ca. ‒ about/roughly p.p. ‒ pro parte (partly) UK ‒ United Kingdom
cm ‒ cenmetre sect. ‒ secon USA ‒ United States of America
e.g. ‒ for example s.l. ‒ sensu lato (broad sense) var. ‒ variety
i.e. ‒ meanng that is s.n. ‒ sin nombre (or name missing) ≤ ‒ less than or equal to
incl. ‒ including sp. ‒ species (singular) ≥ ‒ greater than or equal to
ined. ‒ unpublished spp. ‒ species (plural) < ‒ less than
m ‒ metre s.s. ‒ sensu stricto (narrow sense) > ‒ greater than
mm ‒ millimetre subg. ‒ subgenus ± ‒ more or less
mt ‒ mountain/mount subsp. ‒ subspecies
CONTENTS

3


Phylogenec tree showing relaonships of accepted orders and unplaced families of owering plants divided into
the six informal higher groups used in this work. Major clades are also highlighted for reference but not discussed
further.


 asterids
 Lamiales (-ales)
Gesneriaceae (-aceae)
Didymocarpoideae (-oideae)
 Trichosporeae (-eae)
Jerdoniinae (-inae)
Jerdonia
indica


 
 




Family names end in “-aceae”. However, some of
the largest families were tradionally known by
previous names and these names are sll widely
used. It was agreed amongst botanists to keep
these names in use, alongside more modern
names following the standard “-aceae” ending.
CONTENTS



























DILLENIACEAE

































ASTERIDS



ROSIDS










4
 earliest lineages
Perianth somemes spirally
arranged
Perianth somemes
3-merous
Oen many parts to
each whorl
(many carpels above)
Oen laments and anthers
poorly dierenated.
Plants usually woody, if herbaceous then plants aquac, perianth reduced to absent or (very rarely) grass-like

Usually parallel
venaon
Perianth usually 3-merous Plants usually
herbaceous

Prominent midrib
with parallel
secondary venaon
Perianth usually consisng of
similar whorls (= tepals)
© Denis Barthel /CC-BY-SA-3.0
© Marco Schmidt / CC-BY-SA-3.0
Asparagales

Bulbous plants


 Palm-like
plants

Flowers condensed
into spadix with bract-
like spathe
© Forest & Kim Starr / CC-BY-SA-3.0
 Grass-like
plants
© Sco Zona/ CC-BY-2.0

Flowers are needed for owering plant idencaon though when there are only fruits present it is sll possible but becomes more
dicult. In most cases a combinaon of vegetave and reproducve parts are needed and in some cases a combinaon of both oral
and fruit characters. Plants without a perianth (i.e. both sepals and petals absent) and parasic plants have evolved several mes and
are scaered across the following six higher groups. For these plants it is best to try and idenfy by a process of eliminaon from the
list of families with diagnosc characters pages.
rosidsasterids
© Chrisan Fischer/ CC-BY-SA-3.0

© Marnez-Azorín, Crespo, Dold & Barker/ CC-BY-SA-3.0
© Chris Davidson / oraoheworld.org

14

 Plants in the order have a woody habit, superior ovaries, oen a many parted and spiralling perianth, many stamens and
carpels free or 1. In addion, Schisandraceae have alternate, oen gland-doed and aromac leaves; Trimeniaceae have opposite,
oen gland-doed and aromac leaves, owers aggregated and not solitary and staminodes absent; and Austrobaileyaceae are
woody climbers endemic to Australia with opposite leaves and owers with a rong-sh smell.
 Leaves opposite....................................................................................................................................2
 Leaves alternate................................................................................................ 
 Flowers about 5‒6 cm in diameter...............................................................
2. Flowers less than 1 cm in diameter....................................................................... 

, up to 15 m tall.  simple, opposite (rarely sub-opposite);
margins enre; peoles present; spules present? deciduous and small
or absent.  axillary or rarely terminal solitary owers or 2‒3
aggregated; bracteate. Flowers 5‒6 cm in diameter, bisexual, acnomorphic;
bracteolate. Perianth tepal-like, imbricate; outer whorl oen sepaloid; inner
whorl oen petaloid, greenish-yellow, somemes with brown to purple
spots.  greenish and purple-blotched at the apex; anthers introrse;
staminodes smaller than stamens, ±warty purple-spots.  superior;
carpels free; locules (4‒)6‒9(‒14); ovules several per locule; placentaon
marginal; sgma 2-lobed.  berry-like and orange.
1/species 2; Austrobaileya.
 Australia [tropical Queensland, Atherton Tableland].
(9‒)12‒23(‒many) A6‒11+6°‒16° (4‒)6‒9(‒14)
 The owers are y pollinated and have an odour like rong sh and the gland-
like warts on the stamens and staminodes are thought to produce this unpleasant smell.
 Bailey & Swamy 1949; Endress 1980b, 1993a, 2001.

•A. maculata (stamens and staminodes have conspicuous embossed brown to purple
spots).
•A. scandens (stamen and staminodes lack spots; only found in Daintree Rainforest).

Trees,  or , up to 25 m tall.  oen gland-doed, somemes aromac, simple, opposite;
leaf apex acute to acuminate; margins enre to toothed; peoles present; spules absent; hairs reddish, woolly
or absent.  cymes, racemes or panicles. Flowers ca. 5‒7 mm in diameter, bisexual or unisexual,
acnomorphic. Perianth spirally arranged tepals, imbricate, merging into bracteoles, deciduous; white to cream.
 many, ca. 4 mm long, spirally arranged; anthers basixed.  superior; carpel 1, rarely 2, reduced to
absent in male owers; locule 1; ovule 1; placentaon apical; style absent.  a drupe or berry, red to purple or
black when mature.
1/species 8; Trimenia.
 Scaered from Malesia to Pacic Islands and eastern Australia.
2‒many A6‒many 1(‒2)
 Austrobaileyaceae ‒ carpels 3‒8; owers ca. 3‒5 mm in diameter.
 The generic name Piptocalyx is synonymous with Trimenia.
 Endress 2001; Endress & Sampson 1983; Philipson 1986, 1987a, 1993d; Qui et al. 1999; Rodenburg 1971; Wagner &
Lorence 1999; Whin 2007.
Austrobaileya scandens
© Dennis Stevenson/ plantsystemacs.org
Habit of Austrobaileya scandens

rosidsasterids

15

•T. bougainvilleensis (Solomon Islands).
•T. neocaledonica (leaves obovate; New Caledonia).
•T. nukuhivensis (Marquesas Island [Nuku Hiva]).
•T. macrura (woody climbers, tepals ≤ 10; New Guinea).
•T. marquesensis (Marquesas Islands [Hiva Oa and Tahuata]).
•T. moorei (woody climbers; tepals < 11; northeast New South
Wales to southern Queensland [Australia]).
•T. papuana (plants polygamous, stamen laments long and
thin; Sulawesi and Moluccas [Indonesia], New Guinea).
•T. weinmannifolia (stamen laments short and broad; Samoa,
Fiji).
 
 or small trees to .  usually gland-doed, aromac
(peppery in Illicium) simple, alternate (spiral); margins usually enre, somemes
toothed with chloranthoid teeth; peoles present; spules absent. 
usually axillary solitary owers or a few aggregated; somemes cauliorous.
Flowers bisexual (Illicium) or unisexual (plants monoecious or dioecious: Kadsura,
Schisandra), acnomorphic; oen bracteolate. Perianth spirally arranged tepals;
outer whorl sepaloid, bract-like; inner whorl petaloid, white, yellow to red.
laments free (Illicium) or fused basally (e.g. K. coccinea, Schisandra) or
laments ±fused (most Kadsura); anthers basixed; oen 3‒22 staminodes. 
superior; carpels free; ovule(s) 1 (Illicium), 2‒3 (Schisandra), 2‒5(‒11) (Kadsura);
placentaon marginal to basal.  an aggregaon of free carpels which are eshy
(red to yellow when mature) in Kadsura and Schisandra or star-shaped aggregated
follicles (Illicium).
3/species ca. 80.
Sri Lanka, east and southeast Asia to western Malesia, southeast USA, eastern
Mexico and large Caribbean Islands.
(5‒)9‒15(‒many) A4‒many 5‒many
 Illicium verum (star anise).
 The family have spirally arranged tepals, stamens and staminodes. Illicium is somemes
treated as a separate family. The owers of Illicium oridanum have a faint but unpleasant
odour.
 Endress 2001; Hao et al. 2000, 2001; Panero & Aranda 1998; Saunders 1998, 2000;
Xia & Saunders 2009; Xia et al. 2009.

•Illicium (trees or shrubs; fruit a follicle; southeast Asia, southeast USA and the Caribbean; ca.
40 spp.).
•Kadsura (woody climbers; fruits berry-like and widely separated in a spike; Sri Lanka to the
Philippines and from Java [Indonesia] to South Korea and Japan, parcularly species rich
in southern China; ca. 16 spp.).
•Schisandra (woody climbers; aggregated berry-like fruits are close together and oen glo-
bose; Russian Far East to Java and Bali [Indonesia] and Uar Pradesh [India] to the west,
S. glabra from southeast USA and Mexico; ca. 22 spp.).
Trimenia moorei
© Peter Weston
Schisandra chinensis
© Кордюков Александр/ CC0 1.0
Illicium anisatum
Illicium henryi
Aggregated fruits of
Kadsura japonica
© Alpsdake/CC-BY-SA-3.0
© KENPEI/CC-BY-SA-3.0
Fruits of Illicium verum Schisandra rubriora
rosidsasterids
© Sco Zona/ CC-BY-2.0
© Sco Zona / CC BY 2.0© Thor/ CC BY 2.0


154
ROSIDS
Rosids are a diverse group usually with  , somemes a    present and
stamens usually . They comprise about 25% of all angiosperms diversity with 17
orders and 176 families. Rosid orders are oen morphologically diverse but most large families usually have
disncve characters. A few families have a fused perianth which could be confused with many asterids, notably
Cucurbitaceae and Thymelaeaceae.
 - Trees, shrubs, woody climb-
ers or herbs, somemes succulent; owers bisexual
or unisexual; perianth conspicuous or inconspicuous
to absent; hypanthium somemes present; carpels
usually 2; fruits oen dry. 
VITALES - Woody climbers, shrubs or herbs, oen
tendrils present; bark oen with lencels; leaf vena-
on palmate venaon; spules usually present; ovary
superior; fruit a berry.
 - Trees, shrubs, or herbs,
oen thorny in saline habitats or hemiparasites; sp-
ules present, if absent owers zygomorphic; ovary
superior; fruits dry or a drupe (then plants spiny).
 - Trees, shrubs, woody climbers or
herbs; leaves simple or compound with usually carpel
1 and legume fruits; spules present or rarely absent;
owers oen zygomorphic; stamens usually 8-many;
ovary superior; fruits dry or eshy.
ROSALES - Trees, shrubs, woody climbers or
herbs; spules present or absent; usually owers
bisexual with few to many stamens and conspicuous
perianth OR unisexual with reduced to absent peri-
anth, oen wind-pollinated, usually few stamens and
somemes exudates present, snging hairs or cysto-
liths present. 
 - Usually trees or shrubs; spules pres-
ent; owers usually unisexual, oen in catkins or
spikes, usually wind-pollinated; perianth oen re-
duced to absent; ovary oen inferior; fruits dry.
 - Erect or climbing herbs
or shrubs, rarely trees; spules present or absent;
owers usually unisexual; petals fused, free or ab-
sent, oen valvate; leaf venaon oen palmate or
3‒9-veined from base; ovary usually inferior (if supe-
rior then owers oen bisexual).
CELASTRALES - Trees, shrubs or woody climb-
ers, rarely herbs; spules present or absent; leaves
usually simple; stamens 1-whorled (if 2-whorled then
1-foliate leaves); usually conspicuous nectar disk
present; ovary superior.
 - Trees, shrubs, woody climbers
or herbs; leaves compound (if simple then plants
carnivorous OR with garlic-odour OR petals valvate
with oen fringed apices); spules present or absent;
stamens 1‒2-whorled; ovary superior.
 - Trees, shrubs, woody
climbers or herbs, somemes succulents; owers
acnomorphic or rarely zygomorphic; spules oen
present in most families; petals present or absent;
stamens 1‒2-whorled; ovary usually superior. Vari-

rosidsasterids
© pabloendemico / CC BY 2.0
© Frank Vincentz / CC-BY-SA-3.0 © brewbooks/ CC-BY-2.0
© 4028mdk09/ CC-BY-SA-3.0 © Stan Shebs/ CC-BY-SA-3.0
© Joao Medeiros / CC BY 2.0 © Ji-Elle/ CC-BY-SA-3.0 © H. Zell/ CC-BY-SA-3.0
©Jason Hollinger / CC BY 2.0 © Forest & Kim Starr/ CC-BY-SA-3.0 © Phyzome / CC-BY-SA-3.0
© Michael Clarke/ CC-BY-2.0 © Jacopo Werther/ CC-BY-2.0 © Noblevmy / CC-BY-SA-3.0
©Böhringer Friedrich/ CC BY-SA 3.0 AT
© prenn/ CC-BY-SA-3.0 © H. Zell/ CC-BY-SA-3.0
© Randi Hausken /CC BY 2.0 © Berichard/ CC-BY-2.0
© Chrisan Ordenes/ CC-BY-2.0 © Vojtech Zavadil/ CC-BY-SA-3.0
© Sten Porse/ CC-BY-SA-3.0© J.M. Garg/ CC-BY-SA-3.0


212

Plants in the order are morphologically diverse ranging from trees, shrubs, woody climbers to herbs, as well as, submerged
aquacs (Podostemonaceae), parasites (Raesiaceae) and cactus-like succulents (some Euphorbia in Euphorbiaceae).
There are few constant macromorphological characters which unite the order, though Malpighiales plants typically have 4‒5-merous
owers with free petals, stamens 4‒5 (oen) to 2-whorled or many, and usually fused carpels with superior ovaries (inferior in
Raesiaceae and some Rhizophoraceae; part-inferior in some Salicaceae and superior to inferior in Dichapetalaceae). Flowers are
usually acnomorphic, though zygomorphic owers are somemes present, notably in Viola (Violaceae) and Trigoniaceae. Spules
are usually present in most families.
Some families have: petals absent (Balanopaceae, some Euphorbiaceae, Lacistemataceae, Picrodendraceae, Podostemaceae,
Putranjivaceae, Raesiaceae, some Salicaceae);   (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae,
Goupiaceae, Passioraceae, Rhizophoraceae); or are  (some Euphorbiaceae, some Phyllanthaceae).
Molecular data found the Euphorbiaceae sensu lato and Flacouraceae were among the few tradionally recognised families not
to be found monophylec. This has resulted in spling the Euphorbiaceae into several families and the complete breakup of the
Flacouraceae (see Salicaceae and Achariaceae).
Root-holoparasic herbs without chlorophyll [parasic on Tetrasgma, Vitaceae]
......................................................................................................................
Non-parasic plants with chlorophyll present………………............................................……………………… 2
Aquac plants with lile dierenaon between roots, stems and leaves.
Plants terrestrial, if aquac then roots, stems and leaves dierenated............................................. 3
Woody climbers with leafy tendrils..........................................................
Trees, shrubs or herbs, if woody climbers then leafy tendrils absent.................................................. 4
4. Stamens many, laments basally fused in a ring with small warty structures on laments;
leaves 3-foliate or (rarely) bi-pinnate; peoles long [Costa Rica to Paraguay]
...................................................................................................................
Stamens 1‒many, laments not as above; leaves various, if 3-foliate then peoles not long............ 5
5. Ovule 1 per locule; spules present; ovary ≥1-locular; owers unisexual; perianth 1‒2-whorled
[usually inconspicuous, if conspicuous then exudates oen present]............................................. 
Ovules ≥2 per locule, if 1 ovule then owers bisexual (if unisexual then spules absent
with exudates present or perianth conspicuous and exudates absent).......................................... 
. Plants dioecious; perianth 2-whorled; exudates absent; plants non-succulent; fruits eshy
..........................................................................................................................
. Plants monoecious or dioecious; perianth 1‒2-whorled; exudates present or absent; plants
succulent or non-succulent; fruits non-eshy or (rarely) eshy................
Style (usually) gynobasic; spules present...........................................................................................8
Style terminal or lateral; spules present or absent............................................................................
Anthers dorsixed.................................................................................. 
Anthers basixed...................................................................................................

rosidsasterids


Branch of Bonnea
Herbarium specimen Moreira et al. 7503 (K)

Stout-stemmed trees or .  crowded at branch ends, simple, alternate; margins usually minutely toothed
to enre; peoles short to absent; spules absent; resinous exudates present.  cymes, panicles or
solitary owers (Ploiarium, some Bonnea); bracts 2. Flowers bisexual, acnomorphic; bracteoles 2. Sepals free,
imbricate, unequal, persistent. Petals free, contorted, free; red to pink. many, laments are free (Bonnea)
or basally fused into bundles; anthers basixed.  superior; carpels fused, 3 (Bonnea) or 5; locules 3‒5; ovules
many per locule; placentaon axile; sgmac branches 3‒5.  a sepcidal capsule.
3/species 35.
 Cuba, South America, and Indochina to Malesia.
5 C5 Amany 3‒5
 The family oen occurs in sandy savannas, swampy peaty soils, open rocky areas or
lowland, or mid to high elevaon cerrado or tepuis forests.
 Gustafsson et al. 2002; Weitzman 2005; Weitzman & Stevens 1997; Weitzman
et al. 2007.

•Archytaea (inorescences 3‒many-owered; northeast South America; 2 spp.).
•Bonnea (stamen laments free and not in fascicles; northeast South America [Guiana
highlands], B. paniculata from the Andes, B. stricta from Atlanc coast of Brazil
south to Rio de Janiero or B. cubensis from Cuba; ca. 30 spp.).
•Ploiarium (inorescences solitary owers; Cambodia to Malesia; 3 spp.).
 
Annual or perennial aquac herbs, oen submerged; vegetavely variable: thalloid lichen-like, moss-like to
seaweed-like; roots usually aened, thalloid or liform.  well-developed, reduced or rarely absent, simple
or compound; margins enre, lobed or pinnasect; peoles present or absent; somemes spule-like appendages.
 solitary owers, cymes, spikes or racemes. Flower bisexual, acnomorphic or zygomorphic, buds
naked (some Trischoideae, Weddellinoideae), surrounded by a collar-like cup (some Trischoideae) or enclosed
by a spathella (= tube or sac-like envelope; Podostemoideae); bracteolate. Perianth 1-whorled, free or basally
fused. oen 2 basally fused, 1‒2-whorled or conned to one side, laments when whorled free or fused
(Tulasneantha), oen alternang with tepals.  superior; carpels fused; locules (1‒)2‒3; ovules 2‒many per
locule; placentaon free central or axile.  a capsule, smooth or ribbed.
54/species ca. 300.
 Tropical to temperate regions, plants oen found aached to rocks in fast-owing water.
(0‒)2‒5(‒20) A1‒2(‒many) (1‒)2(‒3)
When the plant is sterile the family could be mistaken for lichen, moss or algae.
 Plants are submerged when water levels are high but during dry seasons they have a terrestrial existence and this is when
they ower. The following generic names are synonymous: Hydrobryopsis = Zeylanidium; Lonchostephus = Mourera; and Maferria
= Farmeria.
 Cook & Rushauser 2006; Cusset 1992; Jäger-Zürn 1997; Kato et al. 2004; Kita 2002; Kita & Kato 2001, 2004; Koi & Kato
2012; Koi et al. 2009, 2012; Les et al. 1997; Moline et al. 2007; Philbrick et al. 2011; Sols et al. 1999a; Tippery et al. 2011; Thiv et
al. 2009.




•Angolaea (leaves repeatedly forked; stamens 3(‒4); sgma 1; capsule valves equal; Angola; 1 sp., A. uitans).
•Dicraeanthus (roots star-shaped; leaves linear, fan-like or forked; stamens 2; capsules ribbed longitudinally, valves equal,
3-ribbed; West and Central Africa).
•Djinga (leaves enre or forked; stamen 1; capsules valves unequal, 3-ribbed; Cameroon; 1 sp., D. felicis).
•Endocaulos (leaves simple; stamens 2; capsules asymmetric, unequal and valves 7-ribbed; Madagascar; 1 sp., E. mangorense).
•Inversodicraea (leaves repeatedly forked; stamen 1; Ghana to Democrac Republic of the Congo).
•Ledermanniella (leaves simple, lobed or forked; stamens 1‒2(‒3); capsule valves usually unequal, 3-ribbed; tropical Africa; ca. 45
spp.).

rosidsasterids

312
 
Annual or perennial, climbing herbs.  aromac (mustard odour
in sap), oen peltate, simple (palmately lobed) or compound (palmate),
alternate or rarely opposite; venaon palmate; margins toothed to enre;
peoles long, usually twining; spules small to absent. 
axillary solitary owers or rarely few-owered and umbel-like; bracteate.
Flowers bisexual, zygomorphic. Sepals fused, petaloid, 1 modied
into a conspicuous long to short spur. Petals free, imbricate, clawed,
upper 2 smaller, oen brightly coloured.  2-whorled, laments
free; anthers basixed.  superior; carpels fused; locules 3; ovule
1 per locule; placentaon axile to rarely apical; style 1.  a 3-lobed
schizocarp or rarely a 3-winged samara.
1/species ca. 95; Tropaeolum.
 Mexico to southern South America, oen culvated.
5 C5 A4+4 3
The generic names Magallana
and Tropaeastrum are synonymous
with Tropaeolum.
 Andersson & Andersson
2000; Bayer & Appel 2002e; Sparre
& Andersson 1991; Watson & Flores
2010.
Fruit of Tropaeolum majus
© Jamain/ CC-BY-SA-3.0
Frontal view of Tropaeolum majus
© Amada44 / CC-BY-SA-3.0
 
 to trees, rarely subshrubs (M. pygmaea), somemes trunk swollen (see
notes).  aromac (mustard or foed odour), deciduous, compound (bi-
pinnate to odd-pinnate), alternate, leaets mainly opposite; margins enre;
peoles present; spules and spels present, somemes represented by
conspicicous glands.  axillary panicles to thyrses; bracteate. Flowers
bisexual, zygomorphic; bracteoles 2. Sepals free, imbricate, petaloid. Petals free,
imbricate, outer petals larger than smaller inner 2; yellow, white or red (M. arborea,
M. longituba).  opposite petals; anthers dorsixed; staminodes present.
 superior, on a stalk; carpels fused; locule 1; ovules many; placentaon
parietal; style 1; sgma 1.  a large capsule, woody, explosively dehiscing and
3-valved.
1/species 13; Moringa.
 Africa, Madagascar, Middle East, India and Pakistan.
5 C5 A5+3‒5° (2‒)3(‒4)
 The genus is variable in habit ranging from
small herbs to subshrubs about 5 cm tall (M.
pygmaea) to trees with swollen trunks. Species
oen ower before the leaves appear and both
are usually needed for idencaon. Species with
swollen trunks include the African M. drouhardii, M.
ovalifolia and M. stenopetala, and Madagascan M.
hildebrandi.
 Hall et al. 2004; Kubitzki 2002h; Olson
2002, 2012; Thulin 1993; Verdcourt 1985.
Moringa ovalifolia
© Muhammad Mahdi Karim / CC-BY-SA-3.0
© Muhammad Mahdi Karim / CC-BY-SA-3.0
©Hans Hillewaert / CC-BY-SA-3.0

rosidsasterids
Tropaeolaceae character
Conspicuous calyx spur

Flowers are transversally zygomorphic (vs. median) which is unusual in angiosperms
Moringa oleifera

332
Thesiaoideae   

     

•Buckleya (shrubs; leaves well-developed; owers unisexual,
4(‒5)-merous; fruit a drupe; eastern USA, East Asia).
•Osyridicarpos (climbing shrubs; leaves well-developed;
owers bisexual, 5-merous; fruit a drupe; Africa; 1 sp., O.
schimperianus).
•Thesium (herbs or subshrubs; leaves small to scale-like;
owers bisexual, usually 5-merous; fruit a nut or drupe;
worldwide; ca. 330 spp.).
Viscaoideae    
     
     


•Arceuthobium (dioecious; leaves reduced to scales; para-
sises Pinaceae or Cuppressaceae species; North and
Central America, Asia, Africa; ca. 45 spp.).
•Dendrophthora (dioecious or monoecious; leaves developed
or scale-like; anthers 1-locular; Caribbean, Mexico, South
America; ca. 70 spp.).
•Ginalloa (leaves well-developed and reduced on same
branch; inorescences spike-like or 1‒3-owered; an-
thers opening by longitudinal slits; India to Malesia).
•Korthalsella (leaves reduced; owers oen < 0.5 mm long;
stamens 3; Old World tropics and subtropics).
•Notothixos (leaves well-developed, 3‒5-nerved; white,
golden or brown stellate hairs especially on young
growth; owers usually 4-merous; Sri Lanka to Malesia,
Australia).
•Phoradendron (monoecious or dioecious; leaves usually
developed; anthers 2-locular; Neotropics; ca. 230 spp.).
•Viscum (monoecious or dioecious; leaves developed or re-
duced; inorescences 3‒5-owered; owers 4-merous;
anthers opening by pores; temperate and tropical Old
World; ca. 150 spp.).Fruits of Phoradendron californicum
© Stan Shebs/ CC-BY-SA-3.0
Thesium alpinum
© Liuthalas / CC-BY-SA-3.0
Viscum album
© Schnobby / CC-BY-SA-3.0
Habit of Arceuthobium cyanocarpum
© Stan Shebs/ CC-BY-SA-3.0
Close-up of
Arceuthobium
divaricatum
© Stan Shebs/ CC-BY-SA-3.0

rosidsasterids
LORANTHACEAE
 or rarely small trees, usually stem-hemiparasites or rarely root-hemiparasites
(±Nuytsia, Atkinsonia, Gaiadendron); stems oen with swollen nodes.  somemes
reduced to scales, oen leathery to eshy, simple, opposite or rarely alternate to whorled;
margins enre; venaon oen inconspicuous; spules absent.  cymes,
racemes, spikes, fascicles, umbels or solitary owers. Flowers bisexual or rarely unisexual,
acnomorphic or somemes zygomorphic. Perianth present or very rarely absent. Sepals
fused, reduced to a calyculus. Petals well-developed, fused or free, valvate, oen tubular;
oen red to yellow.  alternang with the sepals, equal in number to petals,
laments aached to petals; anthers basixed.  inferior; carpels fused; locules
usually 1; ovules 4‒12 per locule; placentaon basal.  a 1-seeded berry or nut to
samara-like (Nuytsia).
76/species ca. 950.
 Tropics to southern hemisphere
temperate regions and Italy, the Balkans to
central Europe.
0 C(0‒3‒)4‒6(‒9) A(3‒)4‒6(‒8)
3‒4(‒12)
  Olacaceae ‒ leaves alternate.
Viscaoideae (Santalaceae) ‒ owers unisexual,
ny tepals.
 Balle 1956; Barlow 1966, 1974, 1984a, 1984b, 1997a, 1997b; Barlow & Wiens 1973; Der & Nickrent 2008; Hopper 2010;
Kuijt 1986, 1988, 1991, 2011, 2012, 2014; Nickrent 1997-; Nickrent et al. 1998, 2010; Polhill & Wiens 1998, 1999a, 1999b; Roldán &
Kuijt 2005; Vidal-Russell & Nickrent 2008a, 2008b.
Loranthaceae characters
Stem-parasic plants (A);
well-developed petals, oen
tubular (B: opened ower of
Agelanthus)
A


450
OLEACEAE 
Trees, ,   (Jasminum, Myxopyrum) or woody herbs (some
Jasminum, Dimetra); stems somemes 4-angled.  simple or compound,
opposite or rarely alternate (Chrysojasminum); margins enre to toothed; peoles
present; hairs lepidote, stellate or glandular; spules absent.  solitary
owers, thyrses, cymes or racemes; somemes bracts. Flowers bisexual or rarely
unisexual (plants polygamomonoecious or dioecious), acnomorphic; nectar disk
present or absent. Sepals fused or absent (some Fraxinus), usually valvate. Petals
fused, rarely free (e.g. Fontanesia) or absent (most Foresera, Fraxinus, Nestegis);
usually yellow or white.  usually 2, opposite the sepals, laments aached
to petals when petals fused, somemes heterostylous, rarely 4 stamens (see notes).
 superior; carpels fused; locules 2; ovules 1‒2(‒many) per locule; placentaon
axile.  a capsule, samara, berry or drupe.
26/species ca. 600.
 ± Worldwide, especially East Asia.
(0‒)4(‒15) C(0‒)4(‒12) A2(‒4) 2
 Olea europaea (olives).
 Apocynaceae ‒ exudates (latex) present. Loganiaceae and Rubiaceae ‒ usually
spules present.
 Swollen nodes have no line across like in most other Lamiales families. Several genera are
non-monophylec, including Olea, Osmanthus and Chionanthus, and further work is needed. The
generic name Menodora is synonymous with Jasminum. Four stamens oen occur in Nestegis and
Osmanthus scortecinii from Malesia (which is ca. 5 species in total).
 Besnard et al. 2009; Filonenko et al. 2009; Green 2004; Guo et al. 2011; Hong-Wa &
Besnard 2013; Kiew & Baas 1984; Kim & Kim 2011; Wallander & Albert 2000; WCSP 2014.

Fontanesieae         

•Fontanesia (owers bisexual or polygamous; stamens longer than corolla tube; Sicily [Italy],
southwest Asia, China).


•Abeliophyllum (petals white to pinkish; stamens inside the corolla tube; fruit a samara; Korea;
1 sp., A. dischum).
•Forsythia (petals yellow; fruit a capsule; southeast Europe, China, Korea, Japan; ca. 12 spp.).
     
   

•Chrysojasminum (leaves alternate; corolla yellow; petals valvate; fruit a eshy berry; Canary
Islands [Spain], Madeira [Portugal], Mediterranean region, Central Asia to Himalayas and
Myanmar, China, southern India to Sri Lanka; ca. 15 spp.).
•Jasminum (leaves opposite; corolla white, yellow or reddish; fruit usually a eshy berry; tropics
to temperate regions; ca. 200 spp.).
•Menodora (leaves opposite or alternate; fruit a dehiscent or indehiscent dry capsule; Central
America, South America, South Africa).


•Dimetra (subshrubs; leaves hairy; petals imbricate; fruit a capsule; northern Thailand; 1 sp., D.
craibiana).
•Myxopyrum (woody climbers; stems 4-angled; leaves 3-nerved; fruit a drupe; India to southern
China, Philippines and New Guinea; ca. 4 spp.).
•Nyctanthes (shrubs or small trees; fruit a schizocarp; India, Thailand, Sumatra, Java [Indonesia]).
Oleeae

•Chionanthus (trees or shrubs; leaves simple; owers bisexual or unisexual; petals valvate; stamens 2 or 4; fruit a drupe; world-
wide; ca. 60 spp.).
•Comoranthus (trees or shrubs; leaves simple; owers bisexual; corolla hairy; fruit a capsule; Madagascar, Comoros).
Forsythia
Nyctanthes arbor-triss
Fruit of Nyctanthes arbor-triss
Jasminum ocinale
© Adityamadhav83 / CC-BY-SA-3.0
© J.M. Garg / CC-BY-SA-3.0
Chionanthus virginicus
© Cephas / CC-BY-SA-3.0
© Hedwig Storch / CC-BY-SA-3.0

rosidsasterids
Oleaceae characters
Flowers acnomorphic with usually
2 stamens (arrowed)
Phillyrea lafolia
© C T Johansson / CC-BY-SA-3.0


 
 to trees or perennial herbs, oen xerophyc, rarely annual or aquac (Trapella); stems somemes swollen.
 somemes deciduous, simple, opposite or somemes alternate (e.g. Sesamum); margins enre to lobed
or pinnad; peoles usually present; spules absent; conspicuous scky (mucilage) glandular hairs (give a slimy
clammy feel).  axillary solitary owers or cymes, rarely raceme-like (Sesamothamnus); pedicels
with 2(‒many) extra-oral nectaries at base (absent from Uncarina and Trapella); bracteate. Flowers bisexual,
±zygomorphic. Sepals fused, blunt-lobed, imbricate, unequal. Petals fused, imbricate, spurred (e.g. Holubia, Rogeria
longiora, some Sesamothamnus); white to yellow or rarely red.  4 and unequal or rarely 2 (Trapella),
opposite the sepals, laments aached to petals; oen staminode 1.  superior or inferior (Trapella), somemes
a nectar disk below; carpels fused, 2 or rarely 3‒4 carpels; locules usually 2; ovules 1‒many per locule; placentaon
axile.  a capsule, drupe or nut; oen winged, spiny or horned.
14/species 70.
 Subtropical to tropical regions, mostly in coastal
or arid habitats in Old World.
5 C5 A(2‒)4+(0‒)1° 2(‒3‒4)
 Sesamum indicum (sesame seeds).
 Martyniaceae ‒ inorescences terminal.
 There are rarely 3‒4 carpels in Josephinia imperatricis, J.
grandiora and some Sesamum orientale.
 Manning 1991; Olmstead et al. 2001; Ihlenfeldt
2004b.

•Ceratotheca (deciduous shrubs or perennial to annual herbs;
corolla white, pink or purple; fruits with 2 lateral horns;
East to southern Africa).
•Dicerocaryum (perennial herbs; owers longly pedicellate;
corolla white, pink, purple or yellow; fruits disc-like with
2 erect conical spines; Madagascar, East to South Africa).
•Harpagophytum (perennial herbs; inorescences 1-ow-
ered; corolla purple, pink or yellow; fruits 4-ridged;
southern Africa).
•Holubia (annual herbs; inorescences 1-owered; corolla
yellow-green, with basal sac-like spur; fruits 4-winged,
indehiscent; southern Africa; 1 sp., H. saccata).
•Josephinia (shrubs, subshrubs or annuals; corolla mauve,
pink or whish; carpels 2(3‒4); fruits globular with
spines; Kenya, Somalia, Malesia, northern Australia).
•Linariopsis (perennial herbs or subshrubs; corolla mauve;
fruits with ny warts; tropical Africa).
•Pedaliodiscus (perennial herbs; inorescences 1-owered;
corolla cream; fruits with 4-longitudinal wings; East
Africa; 1 sp., P. macrocarpus).
•Pedalium (annual herbs; inorescences 1-owered; corolla
yellow; fruits 4-angled; northeast Africa, widespread
pantropical weed; 1 sp., P. murex).
•Pterodiscus (perennial herbs; inorescences 1-owered;
corolla yellow, red, orange or purple; fruits with 4-longi-
tudinal wings; Africa).
•Rogeria (perennial or annual herbs; inorescences 2-ow-
ered; corolla white, red or purple; fruits smooth, winged
or spiny; southern edge of Sahara, southwest Africa).
•Sesamothamnus (trees or shrubs, trunk swollen at base,
spines present; corolla white, pink or yellow; fruits
smooth; northeast and southern Africa).
•Sesamum (deciduous shrubs or perennial to annual herbs;
corolla pink, white or purple; fruits smooth; sub-Saharan
Africa, India, Sri Lanka; ca. 20 spp.).
•Trapella (aquac perennial herbs; corolla white to pink;
stamens 2; staminodes 2; Japan, Russian Far East, Korea,
southeast China; 1‒2 spp.).
•Uncarina (trees or shrubs; inorescences 1‒9-owered;
corolla white, yellow or purple; fruits spiny; Madagascar;
ca. 13 spp.).
Ceratotheca triloba
Habit of Harpagophytum
procumbens
© Henri Pidoux / PD-self
Pedalium murex
© Marco Schmidt/ CC-BY-SA-3.0
Pterodiscus ngamicus
© Michael Wolf/ CC-BY-SA-3.0
Sesamothamnus rivae
© Caroline Waltenberg Wolf/ CC-BY-SA-3.0
Habit of Trapella sinensis
Fruit of Uncarina sp.
© Marco Schmidt/ CC-BY-SA-3.0

rosidsasterids
© Neelix / PD-self
© Show_ryu / PD-self
© Ton Rulkens / CC BY 2.0
© JM Garg / CC BY 2.0
Pedaliaceae character
Extra-oral nectaries (arrowed) are present at the base of the pedicel


Flora Malesiana
1‒  (incl. Singapore]; 2‒ ; 3‒ ; 4‒  [Brunei, Sabah, Sarawak, Kalimantan]; 5‒ ; 6‒
the Philippines; 7‒ ; 8‒  [Bali, Lombok, Sumbawa, Flores, Wetar, Sumba, Timor and Babar]; 9‒ New
[Irian Jaya, Papua New Guinea incl. Bismarck Archipelago].

© Maphobbyist / CC-BY-3.0

Tropic of Cancer

Tropic of Capricorn

CONTENTS
1
2
3
4
5


8




Aagesen L, Sanso AM. 2003. The phylogeny of the Alstroemeriaceae, based on morphology, rps16 intron, and rbcL sequence data.
Syst. Bot. 28: 47-69.
Aagesen L, Medan D, Kellermann J, Hilger HH. 2005. Phylogeny of the tribe Colleeae (Rhamnaceae) – a sensivity analysis of the
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