Plants of the World. An Illustrated Encyclopedia of Vascular Plants

800 Pages·2017·198.845 MB·English

by  Maarten J. M. Christenhusz, Michael F. Fay, Mark W. Chase

To our parents
Benno Christenhusz and Gerdi Leussink
Hugh and Doreen Fay
Wayne and Helen Chase
and Amy Morris
who all nurtured our interest in nature and encouraged our studies in botany
PLANTS OF THE
WORLD
AN ILLUSTRATED ENCYCLOPEDIA
OF VASCULAR PLANTS
Maarten J. M. Christenhusz Michael F. Fay Mark W. Chase
Kew Publishing
Royal Botanic Gardens, Kew
The University of Chicago Press
www.press.uchicago.edu
© The Board of Trustees of the Royal Botanic Gardens, Kew 2017
Text © the authors
Photographs © M. Christenhusz unless stated otherwise on page 672
The authors have asserted their rights to be identified as the authors of this work in accordance with the Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form, or by any means,
electronic, mechanical, photocopying, recording or otherwise, without written permission of the publisher unless in accordance with the
provisions of the Copyright Designs and Patents Act 1988.
Published in 2017 by
Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK
www.kew.org
and
The University of Chicago Press, Chicago 60637, USA
26 25 24 23 22 21 20 19 18 17 1 2 3 4 5
Kew Publishing
ISBN 978 1 84246 634 6 e-ISBN 978 1 84246 636 0
The University of Chicago Press
ISBN-13: 978-0-226-52292-0 (cloth) ISBN-13: 978-0-226-53670-5 (e-book) DOI: 10.7208/chicago/9780226536705.001.0001
Great care has been taken to maintain the accuracy of the information contained in this work. However, neither the publisher nor the authors can be
held responsible for any consequences arising from use of the information contained herein. The views expressed in this work are those of the authors
and do not necessarily reflect those of the publisher or of the Board of Trustees of the Royal Botanic Gardens, Kew.
British Library Cataloguing in Publication Data
A catalogue record for this book is available from the British Library.
Library of Congress Cataloging-in-Publication Data
Names: Christenhusz, Maarten J. M., 1976 – author. | Fay, Michael F., 1960 – author. | Chase, Mark W., 1951– author.
Title: Plants of the world : an illustrated encyclopedia of vascular plants / Maarten J. M. Christenhusz, Michael F. Fay, and Mark W. Chase.
Description: Chicago : The University of Chicago Press, 2017.
Identifiers: LCCN 2017013690 | ISBN 9780226522920 (cloth : alk. paper) | ISBN 9780226536705 (e-book)
Subjects: LCSH: Plants—Encyclopedias. | Plants—Pictorial works.
Classification: LCC QK7 .C47 2017 | DDC 580.3—dc23
LC record available at https://lccn.loc.gov/2017013690
∞ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
Copy editors: Ruth Linklater, Sharon Whitehead
Design, typesetting and page layout: Nicola Thompson, Culver Design
Production Management : Andrew Illes
Cover image: Freycinetia impavida (Pandanaceae), in fruit, Tahiti, French Polynesia (© M. Christenhusz).
Back cover images (from top to bottom): lycopod: Selaginella willdenowii (Selaginellaceae), Malaysia; fern: Angiopteris evecta (Marattiaceae), Moorea,
French Polynesia; gymnosperm: Picea likiangensis (Pinaceae), Yunnan, China; magnoliid: Magnolia hypoleuca (Magnoliaceae), North Carolina, USA;
monocot: Dracula verticulosa (Orchidaceae), Ecuador; eudicot: Cochlospermum fraseri (Bixaceae), Northern Territory, Australia
Printed and bound in China by C&C Offset Printing Co., Ltd
For information or to purchase all Kew titles please visit shop.kew.org/kewbooksonline or email [email protected]
Kew’s mission is to be the global resource in plant and fungal knowledge, and the world’s leading botanic garden.
Kew receives about half of its running costs from Government through the Department for Environment, Food and Rural Affairs (Defra). All other
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CONTENTS
vi How to use this book

1 Introduction

3 Evolution of land plants

4 Plants and human culture

6 Naming plants

8 Classification and the Angiosperm Phylogeny Group

13 Fossil plants

14 Families

14 Etymology and common names

15 Genera

15 Phytogeography

16 Economic botany

18 Lycopods

22 Ferns

71 Gymnosperms

88 The ANA grade families

95 Magnoliids

115 Monocots

213 Eudicots

638 Glossary

671 Acknowledgements

672 Photography credits

673 Further reading

753 General references

756 Index
HOW TO USE THIS BOOK
For an explanation of how plants are scientifically named and how their relationships are studied, see pages 6–8,
and for the types of plant groups, their relationships and how they are visualised, see pages 8–12.
MAGNOLIALES MAGNOLIIDS
Order description with thumbnail
illustration of phylogenetic
MAGNOLIALES
tree and magnifying glass icon
Families 49 to 54 form the order Magnoliales. These woody plants can be recognised by
showing its position on the tree. their often two-ranked or spirally arranged leaves. Their petals are whorled (or spirally
See full-size tree on page 11 for arranged) and their medium-sized seeds have an irregular ruminate endosperm (like nutmeg).
more detail.
49. MYRISTICACEAE exposing woody seeds that are usually covered Uses: Myristica fragrans is a tree with apricot-
Nutmeg family in a lacy or entire, leathery or fleshy aril. like fruits in which nutmeg (the seed) and mace
(the aril) are formed. This native of the Banda
Family number and scientific Distribution: This is a pantropical family Islands in the Maluku Archipelago (Moluccas)
name followed by common name. that are often canopy trees in rainforests. in Indonesia was important in the 17th century
spice trade, giving the name “Spice Islands”
See page 14 for information
Phylogeny and evolution: Myristicaceae to this region. Ground nutmeg is used as
on plant families and common clearly belong to core Magnoliales, which a culinary spice but with excessive use is
probably evolved >100 million years ago. They addictive, toxic and potentially hallucinogenic.
names.
are well supported as sister to Degeneriaceae. Nutmeg oil is used medicinally and for
Fossils from upper Cretaceous deposits in the flavouring tobacco and toothpaste. Bark of
Sahara are known, and Eocene fossil seeds are Iryanthera, Virola elongata and Osteophloeum
These aromatic, often dioecious trees, known from Europe, e.g. from the London clay. platyspermum is used locally as a hallocinogen.
sometimes shrubs, have red sap and red, long Diversification of modern lineages happened Gymnacranthera, Horsfieldia and Knema seeds
terminal buds. Leaves are simple, alternate, fairly recently, c. 15–20 million years ago. have oils that are used to make candles. Fat
Map showing approximate native often oriented in a plane, short-petiolate and from Otoba seeds is used to make soap, and
range for the family (marked in without stipules. Leaf margins are entire, Genera and species: Myristicaceae include Virola sebifera contains oils that are suitable
and hairs on the leaf surfaces and stems are 21 genera with c. 520 species: Bicuiba (1), for candle and soap making. Caihuba, Virola
orange). The accompanying text usually branched or stellate. Inflorescences Brochoneura (3), Cephalosphaera (1), Coelo- surinamensis, produces an edible oil that is
includes climatic and habitat are panicles or fascicled racemes. Flowers caryon (4), Compsoneura (c. 19), Doyleanthus similar to cocoa butter. Horsfieldia iryaghedhi,
are small, actinomorphic and funnel-, bell- or (1), Endocomia (4), Gymnacranthera (7), Pycnanthus angolensis, Staudtia stipitata and
information. See page 15 for urn-shaped. Tepals are usually three, basally Haematodendron (1), Horsfieldia (c. 100), Virola koschnyi produce fine timbers.
further information. fused and often fleshy. Male flowers have two Iryanthera (20), Knema (c. 90), Mauloutchia
to 40 stamens with fused filaments. Female (10), Myristica (c. 170), Osteophloeum (1), Etymology: Myristica is derived from the
flowers have a single carpel, superior ovary Otoba (8), Paramyristica (1), Pycnanthus Greek μύρων (myron), a balm or ointment,
and bilobed stigma. The fruit is a fleshy to (4), Scyphocephalium (4), Staudtia (1) and probably derived from a Semitic root m’rr,
woody capsule, usually splitting in half, Virola (c. 65). meaning bitter, a cognate with myrrh.
Compsoneura excelsa in fruit, Los Mogos, Osa,
Description of key characteristics Myristica fragrans, Singapore (MC) [49] Virola surinamensis, fruit, French Guiana [49] Puntarenas, Costa Rica (CD) [49]
of the family.
Number of genera and number
of species followed by a
list of genera, with species
numbers for each genus given
in brackets. See pages 6–7 for
more information on the naming
102 Christenhusz, Fay & Chase
of plants and page 15 for more
information on genera.
vi Christenhusz, Fay & Chase
HOW TO USE THIS BOOK
Informal higher category and order. See
page 8 for information on taxonomic ranking.
MAGNOLIIDS MAGNOLIALES
All families are represented by
one or more images, showing
key characteristics as well as the
diversity of the group. Captions
include plant name and place
where photographed (if known).
Numbers in blue indicate the
family number.
Tuliptree, Liriodendron tulipifera, Royal Botanic Magnolia macrophylla, fruit, Nichols Arboretum, Magnolia ×soulangeana (a hybrid of M. denudata and
Gardens, Kew, UK [50] Ann Arbor, Michigan, USA [50] M. liliiflora), Royal Botanic Gardens, Kew, UK [50]
List of major economic uses
including traditional and modern
uses for food and construction,
and religious, cultural and
Magnolia stellata, private garden, Kingston upon Magnolia doltsopa, Royal Botanic Gardens, Magnolia campbellii, Royal Botanic Gardens,
Thames, Surrey, UK [50] Melbourne, Australia [50] Kew, UK [50] recreational practices, with notes
50. MAGNOLIACEAE scent and attracting pollinators. The fruit is flowers, Pachylarnax with few carpels but on cultivation of ornamental
Tuliptree family cone-like with free or fused follicles. In many many ovules per carpel, and Talauma with
species. See pages 4–5 and 16 for
species the carpels dehisce, and the pendent fused carpels. These genera have, however,
seeds exhibit a red aril. been found to be embedded in Magnolia sensu more information.
lato, expanding that genus to > 250 species, a
Distribution: The family has a disjunct number that is still growing. The two species
distribution in eastern North America, of Liriodendron are well-supported as sister to
tropical America (Mexico to Brazil and Peru), Magnolia. Magnoliaceae are probably sister to
southern India, Sri Lanka, the Himalayas and the rest of Magnoliales.
throughout temperate and tropical East Asia
(Japan and Korea to New Guinea). Genera and species: This is now a family Origin and meaning of the
consisting of just two genera with c. 267
scientific name on which the
These trees and shrubs have simple, entire and Phylogeny and evolution: The 98 million species: Liriodendron (2) and Magnolia
lobed, spirally arranged and petiolate leaves year old fossil flower Archaeanthus and fossil (c. 265). family name is based. For more
and stipules that enclose the bud and sheath fruits of Lesqueria have been assigned to
information see page 7.
the stem; these soon fall off leaving a scar. Magnoliaceae. Liriodendron in particular Uses: Essential oils from Magnolia champaca
Stalked flowers are formed singly on the end was widespread across the Northern Hemi- are used for perfumery; its leaves are used
of branches or short axillary shoots. Petals are sphere during the late Cretaceous and Tertiary. to feed silk worms. Timber of Magnolia is
free, six or more, spirally or whorled, sometimes Numerous now extinct lineages have been used for boxes, matches, engraving, flooring,
differentiated into sepal-like outer petals and recorded from fossils. More modern represent- broom handles, traditional Japanese shoes etc.
petal-like inner ones. Numerous stamens are atives appeared in the late Miocene in Eurasia, Wood of Liriodendron (whitewood) is used
free and spirally arranged, the filaments short especially when compared to North American for furniture, shingles, latches and formerly
or elongate, often flattened, and the anthers are extant taxa, which are considerably older. canoes. Many species are highly valued Discussion of the evolutionary
elongate with the connective produced into a Several modern genera were described on ornamentals. history including important
tip. Ovaries are superior, often stalked. Carpels the basis of deviating morphological charac-
are usually numerous, sometimes few, spirally ters not found in Magnolia sensu stricto; these Etymology: Magnolia was named in honour fossils, former and current
arranged and free. Beetles are the most frequent include Elmerillia with sessile ovaries, Kmeria of French botanist Pierre Magnol (1638–1715), ideas about relationships and
pollinators, and some species create heat in with unisexual flowers, Manglieta with four or who was the first to publish plant families in
their flowers, increasing the dissemination of more ovules per carpel, Michelia with axillary an intrinsic ‘natural’ classification. groupings within the family
(e.g. subfamilies, clades).
Plants of the World 103
See pages 3 and 13 for
further information.
Plants of the World vii

INTRODUCTION
When thinking of biodiversity, animals first diversity (c. 1,234,000 species), but these compounds to combat being eaten. Therefore,
come to mind for most people. Also, when are also dependent on plants. The diversity many plants make poisons, but many of these,
people think of a tropical rainforest, they and evolution of insects and land plants have given at precise dosages, can also be employed
imagine walls of green vegetation with lots of proceeded hand-in-hand. Of course, there as medicines or pesticides. Humans have thus
animal life: monkeys, tapirs, panthers, snakes, are organisms that happily thrive without benefited from plant defence mechanisms.
colourful birds and butterflies. Perhaps there plants; some detritivores, such as many In addition, humans avoided poisoning by
are some flowers in the rainforest too, but these fungi and bacteria, and some arthropods that domesticating plants to produce toxin-free
are just bright blobs of colour without specific have specialised on feeding on detritivores resources to feed ever-burgeoning human
parts, and all too often people seem to forget can exist without the input of land plants. populations, but these domesticated plants
the enormous diversity of plants that provide These existed before plants evolved, but the have in turn also benefited from this symbiosis
home, food and support for these emblematic organismic diversity of the world would be and are now distributed in huge numbers
animals. Without healthy and diverse forests much diminished and different if plants did worldwide under the protection of their human
of bamboo, there will be no pandas. Because not exist, and of course human civilisation benefactors.
of this ‘green blindness’, conservation efforts totally depends on our green friends. We now know that the flowering plants
and the funding associated with these are often Plants absorb sunlight via chlorophyll in are unique in that they have had multiple
directed towards the protection and study of their leaves. Through photosynthesis, plants rounds of polyploidy, after which most of
large mammals, birds and reptiles (roughly the convert light energy into chemical energy the extra copies of genes were stripped from
same animals that dominate nature television in the form of carbohydrates, such as sugar their genomes, so that they retain only large
documentaries), but conserving these animals and starch, that can later be released to fuel numbers of controlling genes (transcription
is directly related to conserving their habitats activities of the plants. This process is the factors and gene regulators), which are used
and preserving the botanical diversity of most efficient energy conversion known in to provide more sophisticated patterns of
which they are composed. No large mammals, the living world, and this efficiency makes gene expression, giving them advantages
including humans, can exist without the input plants the powerhouses of our planet. over other kinds of plants and allowing
from primary producers, the plants. Plants Because sunlight is essentially limitless and them to better utilise available resources.
not only provide oxygen, food and shelter, photosynthesis so efficient, plants can produce Higher chromosome numbers are the initial
but they also capture and purify water and more carbohydrates than they need for their result of these polyploid events, but these
provide numerous medicinal compounds own growth, and these can be employed to episodes are followed in many plants by
needed to fight or ease diseases of animals produce nectar to attract pollinators or to pack chromosome condensation/reorganisation,
including humans. their fruits and seeds with extra nutrients to resulting in herbaceous species with low
Plant diversity parallels or exceeds most attract animals to disperse their seeds. Plants chromosome numbers, self-pollination
animal diversity, so there is no excuse for use animals to their benefit, and animals and annual life histories, which are highly
green blindness. With an estimated 321,000 in turn, often unknowingly, help plants to suited to domestication and production of
species, plant diversity is much greater achieve their reproductive goals: fertilisation large amounts of carbohydrates. Without the
than the number of vertebrates (62,300), of of egg cells and dispersal of seeds. resources provided by annual herbaceous
which only 5,490 are mammals and some The flowering plants in particular are so flowering plants (e.g. rice, maize, wheat,
10,000 are birds, small groups that receive efficient at reproducing themselves (sexually beans, tomato, lettuce, gourds etc.), it is
lots of scientific attention in comparison. and asexually) that they are good sources of difficult to imagine how human civilisations
Only arthropods, including insects, spiders, food for animals, but plants have in response could ever have developed. Imagine trying
scorpions, lobsters and crabs, have greater developed a large diversity of chemical to feed large cities on the resources provided
Orchis Bank, near Downe (Kent, UK) was one of Charles Darwin’s favourite spots. The diversity of
plants and animals on this herb-rich hillside on chalk provided Darwin with an ideal place to observe
orchids (e.g. Anacamptis pyramidalis, Orchidaceae, foreground) being pollinated by insects, climbing
plants including Bryonia dioica (Cucurbitaceae) and Dioscorea communis (Dioscoreaceae), plant
movement (Oxalis acetosa; Oxalidaceae) and heterostylous species of Primula (Primulaceae). The
fundamental studies he carried out there featured prominently in his books and seminal works on
orchid pollination, climbing plants, the power of movement in plants and heterostyly.
Plants of the World 1
INTRODUCTION
Figure 1a. Plants on masonry, Gymnocarpium dryopteris Figure 1b. Plants on wires, Tillandsia recurvata (Bromeliaceae) in
(Aspleniaceae) on a wall in Uppsala, Sweden Tamaulipas, Mexico © Panoramio, wikimedia commons
by clubmosses, ferns and gymnosperms – it material, beautify our world and make our in the current version of the Angiosperm
would have been impossible. Imagine an planet a pleasant place in which to live. Phylogeny Group classification (APG IV
office without ink, coffee, tea, sugar or milk, Of course several books have addressed 2016) and similar recent classifications for
let alone chocolate digestive biscuits, without the diversity of plants in the past, but DNA ferns and gymnosperms (Christenhusz &
which this book would not have been written. studies from the mid-1990s onward have Chase 2014, Christenhusz et al. 2011). All
The diversity of habits, leaves, flowers revolutionised our ideas about classification families are represented here by one or more
and fruits of land plants is incredible. and the evolution of the land flora. The images, illustrating the diversity of the group
Plants grow in almost every habitat on the diversity of plants is therefore here organised and allowing comparisons. We aim to provide
planet. They can be found in salt and fresh in an evolutionary sequence, placing closely a useful overview of the botanical diversity
water, under icy glaciers and in hot deserts, related families near each other. Because a of our green planet and hope the result is
completely underground or hanging in the book is two-dimensional, we have to present enjoyable. Our intention is that this book will
air on exposed branches in the forest canopy. the plant families in a linear sequence, which provide an introduction to the wall of green
They even invade man-made structures, is to a degree artificial; in many cases the that surrounds us, maybe even be a cure for
like masonry, concrete, asphalt, fences and family placed next to another is equally closely ‘green blindness’, and we would be most
electrical wires (Figure 1). Desirable or not, related to the next two or more families that unhappy if our readers do not go out and see
plants are practically everywhere. They drive follow it, but there is a convention followed by for themselves what wonders the plant world
our climate, provide drinking water, produce most authors who use this sort of system, and has on display!
much of our food, medicine and construction we follow here the linear system as proposed
2 Christenhusz, Fay & Chase