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 funding needed to support Kew’s vital work comes from members, foundations, donors and commercial activities including book sales. 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