Plants of the World: An Illustrated Encyclopedia of Vascular Plants 9780226536705

Citation preview

PLANTS OF THE

WORLD

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

Kew Publishing Royal Botanic Gardens, Kew

The University of Chicago Press www.press.uchicago.edu

Mark W. Chase

© 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 1 3 4 6 8

How to use this book Introduction Evolution of land plants Plants and human culture Naming plants 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

Order description with thumbnail illustration of phylogenetic tree and magnifying glass icon showing its position on the tree. See full-size tree on page 11 for more detail.

MAGNOLIALES Families 49 to 54 form the order Magnoliales. These woody plants can be recognised by their often two-ranked or spirally arranged leaves. Their petals are whorled (or spirally arranged) and their medium-sized seeds have an irregular ruminate endosperm (like nutmeg).

49. MYRISTICACEAE Nutmeg family

Family number and scientific name followed by common name. See page 14 for information on plant families and common names. These aromatic, often dioecious trees, sometimes shrubs, have red sap and red, long terminal buds. Leaves are simple, alternate, often oriented in a plane, short-petiolate and without stipules. Leaf margins are entire, and hairs on the leaf surfaces and stems are usually branched or stellate. Inflorescences are panicles or fascicled racemes. Flowers are small, actinomorphic and funnel-, bell- or urn-shaped. Tepals are usually three, basally fused and often fleshy. Male flowers have two to 40 stamens with fused filaments. Female flowers have a single carpel, superior ovary and bilobed stigma. The fruit is a fleshy to woody capsule, usually splitting in half, Myristica fragrans, Singapore (MC) [49]

Description of key characteristics of the family.

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exposing woody seeds that are usually covered in a lacy or entire, leathery or fleshy aril. Distribution: This is a pantropical family that are often canopy trees in rainforests.

Map showing approximate native range for the family (marked in orange). The accompanying text includes climatic and habitat information. See page 15 for further information.

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 of plants and page 15 for more information on genera.

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Phylogeny and evolution: Myristicaceae clearly belong to core Magnoliales, which probably evolved >100 million years ago. They are well supported as sister to Degeneriaceae. Fossils from upper Cretaceous deposits in the Sahara are known, and Eocene fossil seeds are known from Europe, e.g. from the London clay. Diversification of modern lineages happened fairly recently, c. 15–20 million years ago. Genera and species: Myristicaceae include 21 genera with c. 520 species: Bicuiba (1), Brochoneura (3), Cephalosphaera (1), Coelocaryon (4), Compsoneura (c. 19), Doyleanthus (1), Endocomia (4), Gymnacranthera (7), Haematodendron (1), Horsfieldia (c. 100), Iryanthera (20), Knema (c. 90), Mauloutchia (10), Myristica (c. 170), Osteophloeum (1), Otoba (8), Paramyristica (1), Pycnanthus (4), Scyphocephalium (4), Staudtia (1) and Virola (c. 65). Virola surinamensis, fruit, French Guiana [49]

Uses: Myristica fragrans is a tree with apricotlike fruits in which nutmeg (the seed) and mace (the aril) are formed. This native of the Banda Islands in the Maluku Archipelago (Moluccas) in Indonesia was important in the 17th century spice trade, giving the name “Spice Islands” to this region. Ground nutmeg is used as a culinary spice but with excessive use is addictive, toxic and potentially hallucinogenic. Nutmeg oil is used medicinally and for flavouring tobacco and toothpaste. Bark of Iryanthera, Virola elongata and Osteophloeum platyspermum is used locally as a hallocinogen. Gymnacranthera, Horsfieldia and Knema seeds have oils that are used to make candles. Fat from Otoba seeds is used to make soap, and Virola sebifera contains oils that are suitable for candle and soap making. Caihuba, Virola surinamensis, produces an edible oil that is similar to cocoa butter. Horsfieldia iryaghedhi, Pycnanthus angolensis, Staudtia stipitata and Virola koschnyi produce fine timbers. Etymology: Myristica is derived from the Greek μύρων (myron), a balm or ointment, probably derived from a Semitic root m’rr, meaning bitter, a cognate with myrrh. Compsoneura excelsa in fruit, Los Mogos, Osa, Puntarenas, Costa Rica (CD) [49]

HOW TO USE THIS BOOK

Informal higher category and order. See page 8 for information on taxonomic ranking.

MAGNOLIALES

MAGNOLIIDS

Tuliptree, Liriodendron tulipifera, Royal Botanic Gardens, Kew, UK [50]

Magnolia macrophylla, fruit, Nichols Arboretum, Ann Arbor, Michigan, USA [50]

Magnolia ×soulangeana (a hybrid of M. denudata and M. liliiflora), Royal Botanic Gardens, Kew, UK [50]

Magnolia stellata, private garden, Kingston upon Thames, Surrey, UK [50]

Magnolia doltsopa, Royal Botanic Gardens, Melbourne, Australia [50]

Magnolia campbellii, Royal Botanic Gardens, Kew, UK [50]

50. MAGNOLIACEAE

scent and attracting pollinators. The fruit is cone-like with free or fused follicles. In many species the carpels dehisce, and the pendent seeds exhibit a red aril.

flowers, Pachylarnax with few carpels but many ovules per carpel, and Talauma with fused carpels. These genera have, however, been found to be embedded in Magnolia sensu lato, expanding that genus to > 250 species, a number that is still growing. The two species of Liriodendron are well-supported as sister to Magnolia. Magnoliaceae are probably sister to the rest of Magnoliales.

Tuliptree family

Distribution: The family has a disjunct distribution in eastern North America, tropical America (Mexico to Brazil and Peru), southern India, Sri Lanka, the Himalayas and throughout temperate and tropical East Asia (Japan and Korea to New Guinea). These trees and shrubs have simple, entire and lobed, spirally arranged and petiolate leaves and stipules that enclose the bud and sheath the stem; these soon fall off leaving a scar. Stalked flowers are formed singly on the end of branches or short axillary shoots. Petals are free, six or more, spirally or whorled, sometimes differentiated into sepal-like outer petals and petal-like inner ones. Numerous stamens are free and spirally arranged, the filaments short or elongate, often flattened, and the anthers are elongate with the connective produced into a tip. Ovaries are superior, often stalked. Carpels are usually numerous, sometimes few, spirally arranged and free. Beetles are the most frequent pollinators, and some species create heat in their flowers, increasing the dissemination of

Phylogeny and evolution: The 98 million year old fossil flower Archaeanthus and fossil fruits of Lesqueria have been assigned to Magnoliaceae. Liriodendron in particular was widespread across the Northern Hemisphere during the late Cretaceous and Tertiary. Numerous now extinct lineages have been recorded from fossils. More modern representatives appeared in the late Miocene in Eurasia, especially when compared to North American extant taxa, which are considerably older. Several modern genera were described on the basis of deviating morphological characters not found in Magnolia sensu stricto; these include Elmerillia with sessile ovaries, Kmeria with unisexual flowers, Manglieta with four or more ovules per carpel, Michelia with axillary

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.

List of major economic uses including traditional and modern uses for food and construction, and religious, cultural and recreational practices, with notes on cultivation of ornamental species. See pages 4–5 and 16 for more information.

Origin and meaning of the scientific name on which the family name is based. For more information see page 7.

Genera and species: This is now a family consisting of just two genera with c. 267 species: Liriodendron (2) and Magnolia (c. 265). Uses: Essential oils from Magnolia champaca are used for perfumery; its leaves are used to feed silk worms. Timber of Magnolia is used for boxes, matches, engraving, flooring, broom handles, traditional Japanese shoes etc. Wood of Liriodendron (whitewood) is used for furniture, shingles, latches and formerly canoes. Many species are highly valued ornamentals. Etymology: Magnolia was named in honour of French botanist Pierre Magnol (1638–1715), who was the first to publish plant families in an intrinsic ‘natural’ classification. Plants of the World

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Discussion of the evolutionary history including important fossils, former and current ideas about relationships and groupings within the family (e.g. subfamilies, clades). See pages 3 and 13 for further information.

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INTRODUCTION When thinking of biodiversity, animals first come to mind for most people. Also, when people think of a tropical rainforest, they imagine walls of green vegetation with lots of animal life: monkeys, tapirs, panthers, snakes, colourful birds and butterflies. Perhaps there are some flowers in the rainforest too, but these are just bright blobs of colour without specific parts, and all too often people seem to forget the enormous diversity of plants that provide home, food and support for these emblematic animals. Without healthy and diverse forests of bamboo, there will be no pandas. Because of this ‘green blindness’, conservation efforts and the funding associated with these are often directed towards the protection and study of large mammals, birds and reptiles (roughly the same animals that dominate nature television documentaries), but conserving these animals is directly related to conserving their habitats and preserving the botanical diversity of which they are composed. No large mammals, including humans, can exist without the input from primary producers, the plants. Plants not only provide oxygen, food and shelter, but they also capture and purify water and provide numerous medicinal compounds needed to fight or ease diseases of animals including humans. Plant diversity parallels or exceeds most animal diversity, so there is no excuse for green blindness. With an estimated 321,000 species, plant diversity is much greater than the number of vertebrates (62,300), of which only 5,490 are mammals and some 10,000 are birds, small groups that receive lots of scientific attention in comparison. Only arthropods, including insects, spiders, scorpions, lobsters and crabs, have greater

diversity (c. 1,234,000 species), but these are also dependent on plants. The diversity and evolution of insects and land plants have proceeded hand-in-hand. Of course, there are organisms that happily thrive without plants; some detritivores, such as many fungi and bacteria, and some arthropods that have specialised on feeding on detritivores can exist without the input of land plants. These existed before plants evolved, but the organismic diversity of the world would be much diminished and different if plants did not exist, and of course human civilisation totally depends on our green friends. Plants absorb sunlight via chlorophyll in their leaves. Through photosynthesis, plants convert light energy into chemical energy in the form of carbohydrates, such as sugar and starch, that can later be released to fuel activities of the plants. This process is the most efficient energy conversion known in the living world, and this efficiency makes plants the powerhouses of our planet. Because sunlight is essentially limitless and photosynthesis so efficient, plants can produce more carbohydrates than they need for their own growth, and these can be employed to produce nectar to attract pollinators or to pack their fruits and seeds with extra nutrients to attract animals to disperse their seeds. Plants use animals to their benefit, and animals in turn, often unknowingly, help plants to achieve their reproductive goals: fertilisation of egg cells and dispersal of seeds. The flowering plants in particular are so efficient at reproducing themselves (sexually and asexually) that they are good sources of food for animals, but plants have in response developed a large diversity of chemical

compounds to combat being eaten. Therefore, many plants make poisons, but many of these, given at precise dosages, can also be employed as medicines or pesticides. Humans have thus benefited from plant defence mechanisms. In addition, humans avoided poisoning by domesticating plants to produce toxin-free resources to feed ever-burgeoning human populations, but these domesticated plants have in turn also benefited from this symbiosis and are now distributed in huge numbers worldwide under the protection of their human benefactors. We now know that the flowering plants are unique in that they have had multiple rounds of polyploidy, after which most of the extra copies of genes were stripped from their genomes, so that they retain only large numbers of controlling genes (transcription factors and gene regulators), which are used to provide more sophisticated patterns of gene expression, giving them advantages over other kinds of plants and allowing them to better utilise available resources. Higher chromosome numbers are the initial result of these polyploid events, but these episodes are followed in many plants by chromosome condensation/reorganisation, resulting in herbaceous species with low chromosome numbers, self-pollination and annual life histories, which are highly suited to domestication and production of large amounts of carbohydrates. Without the resources provided by annual herbaceous flowering plants (e.g. rice, maize, wheat, beans, tomato, lettuce, gourds etc.), it is difficult to imagine how human civilisations could ever have developed. Imagine trying 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.

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INTRODUCTION

Figure 1a. Plants on masonry, Gymnocarpium dryopteris (Aspleniaceae) on a wall in Uppsala, Sweden

by clubmosses, ferns and gymnosperms – it would have been impossible. Imagine an office without ink, coffee, tea, sugar or milk, let alone chocolate digestive biscuits, without which this book would not have been written. The diversity of habits, leaves, flowers and fruits of land plants is incredible. Plants grow in almost every habitat on the planet. They can be found in salt and fresh water, under icy glaciers and in hot deserts, completely underground or hanging in the air on exposed branches in the forest canopy. They even invade man-made structures, like masonry, concrete, asphalt, fences and electrical wires (Figure 1). Desirable or not, plants are practically everywhere. They drive our climate, provide drinking water, produce much of our food, medicine and construction

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Figure 1b. Plants on wires, Tillandsia recurvata (Bromeliaceae) in Tamaulipas, Mexico © Panoramio, wikimedia commons

material, beautify our world and make our planet a pleasant place in which to live. Of course several books have addressed the diversity of plants in the past, but DNA studies from the mid-1990s onward have revolutionised our ideas about classification and the evolution of the land flora. The diversity of plants is therefore here organised in an evolutionary sequence, placing closely related families near each other. Because a book is two-dimensional, we have to present the plant families in a linear sequence, which is to a degree artificial; in many cases the family placed next to another is equally closely related to the next two or more families that follow it, but there is a convention followed by most authors who use this sort of system, and we follow here the linear system as proposed

in the current version of the Angiosperm Phylogeny Group classification (APG IV 2016) and similar recent classifications for ferns and gymnosperms (Christenhusz & Chase 2014, Christenhusz et al. 2011). All families are represented here by one or more images, illustrating the diversity of the group and allowing comparisons. We aim to provide a useful overview of the botanical diversity of our green planet and hope the result is enjoyable. Our intention is that this book will provide an introduction to the wall of green that surrounds us, maybe even be a cure for ‘green blindness’, and we would be most unhappy if our readers do not go out and see for themselves what wonders the plant world has on display!

EVOLUTION OF LAND PLANTS As little as 30 years ago, the evolution of plants was portrayed as a linear, fairly straightforward increase in complexity. For example, Psilotum and Equisetum were considered perhaps to be related to the earliest vascular plants because they were at least superficially similar in appearance to fossils from the Devonian and Silurian. The complex algal charophytes were considered the closest relatives of land plants simply because they looked superficially like aquatic vascular plants. However, this comfortable sequence of plant evolution has in the past 20 years been challenged and shown to have been much more complex than previously assumed. Land plants (often called ‘Embryopsida’, but here Equisetopsida) form a lineage that is most closely related to just one genus of the green algae, Coleochaete. The early vascular plants are lycopods, whereas Psilotum and Equisetum are now known to be closely related to ferns, in which we now classify them. When plants emerged on land is not precisely known, but the estimated dates vary from 439 to 912 million years ago, although most estimates fall around 480–500 million years. Land plants can be distinguished by having sporopollenin in the spore wall, which probably facilitated their spread on land because it protects their spores from dehydration. The earliest land plants did not have roots, like their green algal

ancestors; in aquatic environments, roots are generally not required or are only used to secure a plant in position on the substrate. In the earliest stages of the invasion of land by plants, the functions of roots were performed by fungi in exchange for carbohydrates, and roots perhaps first evolved to make this process more efficient and less subject to the drier conditions that exist at the soil surface. Many plants today still maintain a relationship with fungi, called a mycorrhizal association, but some like the mustard family, Brassicaceae, appear to have foregone it. Among the earliest plants to colonise the land were relatives of liverworts, mosses and hornworts (Figure 2); also present were other groups of plants about which we know relatively little, other than their structure that is obvious from the study of their fossils; their life-history strategies are mostly in the realm of speculation. Mosses, hornworts and liverworts, collectively often termed bryophytes, exhibit the alternation of generations observed in the green algae, but they have a dominant gametophyte (the haploid generation, with one set of chromosomes). The sporophyte generation (diploid, with two sets of chromosomes) is dependent on the gametophyte. Most phylogenetic studies indicate that the hornworts alone are closer (sister) to the rest of the land plants, those with

a vascular system also called the “vascular plants”, which are the focus of this book. This term is a simplification and resulted in the bryophytes being termed “lower plants”; they do have water-conducting tissues, but these differ from those found in the remainder of the land plants, the large clade composed of the lycopods, ferns, gymnosperms and flowering plants. In all other groups of land plants, it is the sporophyte generation that is dominant; in ferns, the gametophytes are free-living and generally small, whereas in the seed plants, the gametophytes are entirely dependent on the sporophyte and not easily observed, unless one knows where to look in the developing pollen and ovary. In seed plants, pollen is released from the parent sporophyte. Pollen contains the male gametophytes, which release sperm; pollen is often considered the “male” part of a plant and described as the plant counterpart to animal sperm, but this is not strictly true. Pollen carries the sperm, and this grows out of the pollen when it lands on a stigma and makes its way via the pollen tube to the ovary where fertilisation happens. Only then is a seed formed, and seeds are inside a fruit, a structure that usually helps with the dispersal of the seed, either mechanically (explosion, wind, gravity, water) or by animals, in their fur or on their feet or inside their guts and deposited with their faeces.

Figure 2a. Liverwort: Asterella drummondii (Asterellaceae), Western Australia

Figure 2b. Moss: Polytrichum commune (Polytrichaceae), Lake District, UK

Figure 2c. Hornwort: Anthoceros agrestis (Anthocerotaceae), Germany © BerndH, wikimedia commons

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PLANTS AND HUMAN CULTURE

Figure 3a. Stone carving of Adam and Eve with the Apple of Paradise. Facade of Notre Dame, Paris, France

Nearly all animals are totally dependent on plants. Plants fix carbon into energy-rich compounds with the help of sunlight, and as a waste product they produce oxygen, which animals use to metabolise this carbon-rich fuel. However, apart from providing nutrients, food and medicine, plants are important for the development of human culture, a particular trait that sets Homo sapiens apart from other animals. There was said to be a rotten Apple in Paradise (Figure 3), and, even though the apple did give humans ‘wisdom’ and a narrative, humans had to learn how to fend for themselves in the wide world. Luckily there were plants outside the Garden of Eden as well. As soon as humans learned how to make tools, plants became directly associated with hominid culture, perhaps a stick to poke a hole to retrieve honey from a bee hive was one of the first, followed by spears and arrows to hunt and wood to build a shelter, leaves to make a roof and fibres to bind it all together. Using plant material as fuel for fire is critical in survival strategies of the past and present. Even though leather, fur and bones were frequently used by early humans for clothing and tools, fibre from cotton and

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Figure 3b. Adam naming the plants and animals. Fresco in Gran Meteora monastery, Greece.

linen were much more abundant, renewable and much easier to obtain, and this certainly helped humans to expand into much hotter and colder areas than their original climatic range encompassed. From hunter-gatherer societies that were frequently on the move, humans started to settle and form villages. This was only possible because plants (and animals) could be domesticated, and cultivation at larger scales became possible, producing more than a single farmer’s family could eat. All major seed crops are annuals (or have been selected for an annual life history if they were perennial). In addition, plants with a clear dormant period were domesticated for crops like potatoes and onions. These crop plants differ from all longer-lived species in their inherent capacity to put all their resources into the production of seeds or perennating structures (such as the

stem tubers of potato and other root crops). Little or nothing is held back by the parent plant for the following season apart from seeds and tubers, which humans can use as food and to grow the crop the following year. This massive and easily harvested product allowed farmers to sell part of their harvest in exchange for other goods and in due course, after many independent and repeated attempts, this allowed humans to diversify, specialise and develop a variety of trades and industries. The basis of communities, towns and cities was set by the domestication of plants. For the success of agriculture, whether to produce animals or plants as the final product, plants are of enormous importance. They produce fodder, make and improve soil and recycle and purify water. Despite all our accomplishments, we owe our existence to a six-inch layer of topsoil, sunshine and rain.

PLANTS AND HUMAN CULTURE Most other industries are also dependent on plants, varying from the publishing and financial sectors (paper is made of plant fibres) to all manufacturing that uses coalgenerated electricity or carbon-based fuels in their production (coal and oil are fossil plant material). Human civilisation has exceeded the mitigating ability of plants to compensate for our activities, and our stripping of vegetation from large parts of the planet has now become the biggest threat to our existence. In addition, the climate is changing due to the burning of fossil fuels and the impact humans have on life through degradation of habitats, exploitation and depletion of natural resources. Carbon that was fixed by plants hundreds of millions of years ago and held in reserve below the Earth’s surface is burned and released into the atmosphere. To a certain extent, plants have benefited from the increased levels of CO2

and are growing more quickly and larger as a result of the increase in atmospheric carbon, but the amount being released increases so rapidly that plants cannot keep up. More carbon in the air results in a changing climate, rising sea levels, more and stronger storms, freak weather events, unusual cold periods, drought, heat waves and floods. Unfortunately it is easier to secure funding for the protection of cuddly animals like lemurs or impressive large cats (and even then the money available is relatively limited) than it is to obtain funds to secure the existence of a rare species of tree, even though a local economy may be based on the fruits of these trees. Financial aid to fund a goat for a small village in the Sahel is the worst development aid imaginable. Goats increase desertification by eating all vegetation and reduce the possibility for

people to grow food locally (Figure 4). In fact eradication of goats, especially feral ones, has in many arid regions improved the water balance and allowed more sustainable food production. Exclusion of people also allows ecosystems to reset, as seen in the area around Chernobyl in Ukraine, where wildlife is now rampant. With a growing human population, climate change and political instability in many parts of the world, the need for selfsustainable, renewable, environmentally friendly livelihoods increases, and plants will play a key role in helping to achieve this. This encyclopedia lists the usefulness of many plants, some well-known crops but also other useful plants that are poorly known. We hope that this will bring the economic importance of botany to the attention of people and perhaps spark the economic use of some ‘forgotten’ species.

Figure 4. Goats climbing an argan tree (Argania spinosa, Sapotaceae) in Morocco (HR)

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NAMING PLANTS Nomenclature deals with the internationally agreed rules for naming of plants, whereas taxonomy is how and to which organisms these names should be applied. Systematics studies how these organisms are related to each other and the characters by which they can be recognised or diagnosed. These three disciplines are inter-related and should not to be confused. Taxonomy always aims for the most stable solution when new classifications and name changes are proposed, and nomenclature should help with that through the principle of priority (the first published name is generally the one that should be used). There is an elaborate system of rules to decide on the correct name that should be chosen. Unfortunately, names will have to change when it is found that species were wrongly placed, when for instance some are found to belong to a different clade (a group of uniquely related organisms) than originally assumed. Earlier these changes were made on the basis of morphological characteristics alone, which sometimes resulted in the grouping of unrelated species based on similar characters that evolved independently (convergent evolution). With the help of DNA data, comparisons of which allow us to build phylogenetic trees or assess species limits, the taxonomist has gained a valuable, less subjective tool to help assess relationships and build classifications, while at the same time taking into account morphological, anatomical and chemical similarities; all are tools in the taxonomist’s toolbox. Systematics is increasingly using various techniques from genetics to understand relationships between plants and to revise classifications. Even though some major changes of classification occurred due to the use of DNA in classification, it should be noted that many plant families have retained their traditional circumscription. Why is it important that the correct scientific names of plants are used? The main function of names is communication. When you have an internationally accepted name of an organism, information about these organisms or group of organisms (such as genera, families or orders) can be communicated without confusion. Why not use the common names of species? Common names vary from region to region and country to country. For instance, a bluebell would in England be Hyacinthoides non-scripta (Asparagaceae), in Scotland Campanula rotundifolia (Campanulaceae), in Australia Billardiera heterophylla (Pittosporaceae), in New Zealand Wahlenbergia gracilis (Campanulaceae), in Texas Eustoma grandiflora (Gentianaceae), in eastern North America Mertensia virginica (Boraginaceae) and in California Phacelia campanularia (Boraginaceae) (Figure 5). On the other hand,

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Figure 5a. Bluebell Hyacinthoides non-scripta (Asparagaceae)

Figure 5b Bluebell Campanula rotundifolia (Campanulaceae)

Figure 5c. Bluebell Billardiera heterophylla (Pittosporaceae)

Figure 5d. Bluebell Phacelia campanularia (Boraginaceae)

NAMING PLANTS the same species can have many different names in different languages. Hyacinthoides non-scripta is restricted to Atlantic Europe, from northern Spain to England, but is known variously as almindelig klokkeskilla, bell bottle, bluebell, bokidi koukou, clychau’r gog, coinnle corra, fairy flower, Hasenglöckchen, jacinthe sauvage, jacinto de los bosques, muguet bleu, wild hyacinth, wilde hyacint and wood bell, among other names. This diversity of common names for the same plant makes it difficult to find and synthesise all information published about a species. However, a stable scientific name can be used by people across all languages, geographical borders and scientific disciplines. This makes it clear which of the bluebells is being discussed, although scientific names are often omitted from the popular press because they are deemed too complicated for the general public to comprehend, whereas in fact they make life easier when you want to look things up and cause less confusion when doing so. When carrying out any type of research on plants, it is always advisable to make a dried herbarium voucher accompanied by a label stating all information you have about the specimen: name, size, stature, colour, date of collection, precise locality of collection, geographical coordinates, elevation, what study the specimen was used in and the collector’s name with a unique collection number. This specimen should then be deposited in a public herbarium, where other researchers can use it to be certain about the plant species that was studied, in case they want to repeat this study in the future. For repeatability of scientific studies, this is of incredible importance. The value of herbaria and other natural history collections is sometimes downplayed as ‘stamp collecting’, but it is much more than that. New species are still regularly found among specimens in natural history collections, and if we want to know all about the organisms with which we share our planet, understanding their distributions, habitats and changes to these, we will have to start with the material already in our collections. Natural history collections house vast amounts of information about diversity, geography, phenology etc. that we are only now starting

to use on a broader scale, especially as a result of the mass digitisation of these collections, which makes these data available on the internet. These collections also document our cultural history, because so much of human history was founded on our uses of plants. By eliminating these collections, we will lose valuable evidence about our past and potential revivable uses of plants from our history. Using these georeferenced databased collections, we can now map where target or indicator species occurred in the past versus the present and use these data to predict future distributions and extinctions under a changing climate. We can study morphological and genetic diversity of species because we are able to extract DNA from type (the specimen upon which a name is based) and other historical specimens to place taxa correctly and with certainty in a phylogenetic analysis. Also we can always go back to the original material used in previous studies, making these repeatable and thus scientific. Why do scientific names have to change? The simple answer to this common question is that names should reflect evolutionary relationships and therefore a name may have to change when our understanding of relationships has changed, usually because more data are available. Admittedly, the English bluebell referred to above was formerly known as Scilla non-scripta, Hyacinthus non-scriptus and Endymion non-scriptus, but these changes have more to do with the historical delimitation of the genera Hyacinthus and Scilla than with actual nomenclatural change, as you can see the species epithet remains the same. There are also changes as a result of the nomenclatural rules, although one of the principles is to keep names as stable as possible. The rules of priority may result in the uptake of an older, forgotten name for a species, which takes precedence over a more recent one that has commonly been in use. It is also possible that a type specimen has been erroneously interpreted and that (part of) the species needs a different name. Then there are correctable errors, listed in the nomenclatural code, where endings and conjugations may need changing. In the 1750s, Carolus Linnaeus developed

the binomial system that we use today. This simplified naming, in which all species names are composed of two names (genus and species), replaced the short descriptive phrases common in earlier times. The longer phrase names were workable as long as the number of taxa was limited, but as more of the world was explored, this system became increasingly unworkable. Shortening the phrase to two words made it much easier to remember names and grouped similar species in genera. During the late 18th and the 19th century, there were many species discovered that could not be easily placed in one of the existing genera, and so these often were placed in genera of their own. These were also often tentatively placed in a family, although evidence for some of these placements was scant. Now that we have better understanding of relationships, many species have been reorganised into genera that make more sense on the basis of molecular and morphological data. To prevent such future nomenclatural change, it has been proposed that we use the phylogenetic tree as the sole evidence of their relationships. If, for example, an unknown animal was originally described as a ‘desert goat’ because it vaguely resembled a domestic goat, but later the animal was shown to be a kind of horse instead, it would continue to be called a goat, meaning that its name then tells us nothing about its relatedness. Similarly, scientific names of species would be stable and forever remain the same. Of course, stability of names should always be taken into account when changes of names are considered, and when there are several options, the choice resulting in the fewest name changes should be preferred, especially when economically important species are involved. If this is nomenclaturally impossible, because priority is involved, a case can be made for conservation or rejection of a name, which is then voted on by the Nomenclatural Committee. For this reason, we now use the name Amaryllidaceae instead of the older name Alliaceae. However, in principle, the rule of priority should be strictly followed. We prefer to retain the current system, in which a species name contains some information about what its general relationships are.

Plants of the World

7

NAMING PLANTS Taxonomic ranks in plants have standard endings so they can be easily recognised, as for instance an order or family. These endings are as follows: Division: -ophyta, e.g. Equisetophyta (this rank is rarely used) Subdivision: -ophytina, e.g. Equisetophytina (this rank is rarely used) Class: -opsida, e.g. Equisetopsida Subclass: -idae, e.g. Equisetidae Superorder: -anae, e.g. Equisetanae Order: -ales, e.g. Equisetales Suborder: -ineae, e.g. Equisetineae (this rank is rarely used) Family: -aceae, e.g. Equisetaceae Subfamily: -oideae, e.g. Equisetoideae Tribe: -ideae, e.g. Equisetideae Subtribe: -inae, e.g. Equisetinae

All these names are based on the stem of the genus that typifies these ranks. In this case the type genus is Equisetum (stem: Equiset-), which means ‘horse bristle’, the horsetail in English. Genus names and species epithets are always written in italics with the genus name capitalised. The species epithet (the second of the two words that make up the species name) is not capitalised. Below the rank of species there are several additional ranks, mainly subspecies (subsp.), variety (var.) and forma (fo.). It has to be noted that the rule of priority applies only within a rank, not across ranks, and that at and above the level of order the rule of priority does not apply. Some names have also been conserved or rejected, usually to prevent unwanted nomenclatural change.

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP In early classifications, plants were usually organised according to their presumed medicinal traits. The first comprehensive herbals of the 16th century provided some accurate circumscriptions but were organised by usefulness rather than similarity or putative relationships. In the mid-18th century, Carolus Linnaeus invented an artificial system for classification of plants based on numbers of flower parts (Figure 6). His ‘sexual system’ described how, for instance ‘six men were in bed with one woman’ (hexandria monogynia) in Allium, Hyacinthus and Hemerocallis, three genera we now know to belong to three families. These characters were useful for identification, but the classification was known to be artificial (unnatural) because other characters, for instance of leaves etc., were not taken into account. Many of these sexual characters evolved in parallel, and thus species were frequently placed together that we now know

8

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are distantly related and vice versa. Linnaeus himself knew that this system had these shortcomings, but he was more interested in an easily understood classification than in placing related species together. The prevailing species concept of the time was that species were unchangeable entities created by God. With the advent of Darwinian evolution and discovery of Mendelian genetics, species concepts changed. For classification, usually extrinsic morphological characters became used to secure a more ‘natural’ classification, but these were often subjectively interpreted characters. Workers such as Cronquist (1981), Takhtajan (1980, 1997) and Thorne (1992) based their classifications on all data available at the time (morphology, anatomy, ontogeny, chemistry etc.), but subjectively filtered the Figure 6. The sexual system of Linnaeus as illustrated by Ehret (1736)

All names are followed by the author name (often abbreviated), which is done to prevent confusion in the cases of similar names or homonyms by various authors. This book deals mostly with the levels above genus; at these higher levels, name confusion is not a big issue. Therefore in this encyclopedia we have omitted these author names because they are not commonly used outside taxonomic publications. We recommend that all scientific journals include author names as full references in their papers, as this links to descriptions and associated data on the species when it was first found.

NAMING PLANTS Taxonomic ranks in plants have standard endings so they can be easily recognised, as for instance an order or family. These endings are as follows: Division: -ophyta, e.g. Equisetophyta (this rank is rarely used) Subdivision: -ophytina, e.g. Equisetophytina (this rank is rarely used) Class: -opsida, e.g. Equisetopsida Subclass: -idae, e.g. Equisetidae Superorder: -anae, e.g. Equisetanae Order: -ales, e.g. Equisetales Suborder: -ineae, e.g. Equisetineae (this rank is rarely used) Family: -aceae, e.g. Equisetaceae Subfamily: -oideae, e.g. Equisetoideae Tribe: -ideae, e.g. Equisetideae Subtribe: -inae, e.g. Equisetinae

All these names are based on the stem of the genus that typifies these ranks. In this case the type genus is Equisetum (stem: Equiset-), which means ‘horse bristle’, the horsetail in English. Genus names and species epithets are always written in italics with the genus name capitalised. The species epithet (the second of the two words that make up the species name) is not capitalised. Below the rank of species there are several additional ranks, mainly subspecies (subsp.), variety (var.) and forma (fo.). It has to be noted that the rule of priority applies only within a rank, not across ranks, and that at and above the level of order the rule of priority does not apply. Some names have also been conserved or rejected, usually to prevent unwanted nomenclatural change.

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP In early classifications, plants were usually organised according to their presumed medicinal traits. The first comprehensive herbals of the 16th century provided some accurate circumscriptions but were organised by usefulness rather than similarity or putative relationships. In the mid-18th century, Carolus Linnaeus invented an artificial system for classification of plants based on numbers of flower parts (Figure 6). His ‘sexual system’ described how, for instance ‘six men were in bed with one woman’ (hexandria monogynia) in Allium, Hyacinthus and Hemerocallis, three genera we now know to belong to three families. These characters were useful for identification, but the classification was known to be artificial (unnatural) because other characters, for instance of leaves etc., were not taken into account. Many of these sexual characters evolved in parallel, and thus species were frequently placed together that we now know

8

Christenhusz, Fay & Chase

are distantly related and vice versa. Linnaeus himself knew that this system had these shortcomings, but he was more interested in an easily understood classification than in placing related species together. The prevailing species concept of the time was that species were unchangeable entities created by God. With the advent of Darwinian evolution and discovery of Mendelian genetics, species concepts changed. For classification, usually extrinsic morphological characters became used to secure a more ‘natural’ classification, but these were often subjectively interpreted characters. Workers such as Cronquist (1981), Takhtajan (1980, 1997) and Thorne (1992) based their classifications on all data available at the time (morphology, anatomy, ontogeny, chemistry etc.), but subjectively filtered the Figure 6. The sexual system of Linnaeus as illustrated by Ehret (1736)

All names are followed by the author name (often abbreviated), which is done to prevent confusion in the cases of similar names or homonyms by various authors. This book deals mostly with the levels above genus; at these higher levels, name confusion is not a big issue. Therefore in this encyclopedia we have omitted these author names because they are not commonly used outside taxonomic publications. We recommend that all scientific journals include author names as full references in their papers, as this links to descriptions and associated data on the species when it was first found.

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP

Sister species Rapid radiation

Extinction

Speciation

Common ancestor

Figure 7a. Hypothetical visualisation of a phylogenetic tree on a real tree (Callitris glaucophyllum, Cupressaceae)

Sister species

tin on ct i

Speciation

Ex

information to determine which character was most important and should be the basis of classification. These classifications had a great following, but unfortunately ontogenetic data could not always be obtained, and thus parts of these classifications were sometimes still only a matter of speculation. Phylogenetic analysis of scored morphological and ontogenetic characters was sometimes carried out, sometimes also including fossil taxa. This made classifications less subjective, although they were dependent on the interpretation of the characters, and the manner in which they were scored occasionally influenced results. In the late 1980s, the molecular age began, and increasingly greater numbers of DNA sequences could be compared, analysed separately or together with morphological data, resulting in testable, much less subjective classifications. The use of DNA sequences in phylogenetic reconstructions almost entirely removed subjectivity and allowed a better understanding of relationships of morphologically difficult-toplace groups. Relationships of plants are usually visualised by means of a phylogenetic tree. In those based on DNA information, each terminal branch is based on the DNA sequence of an individual plant, which is often considered to represent its species, genus, family or (as in the case below) order. Species that are alive today may have had many extinct relatives, which in a phylogenetic tree will make such species look isolated (Figure 7). Sometimes, when a species-rich clade is sister to a species-poor clade, people tend to read it as if the small clade is “basal” or “primitive”, but this assumption is erroneous. It is a misconception that makes reading a phylogenetic tree problematic. For instance, most modern ferns have evolved as recently as most flowering plant clades, but the ferns are an old lineage and have retained many of the original fern characters. However, ferns are also competitive in modern ecosystems and by no means primitive in all their attributes. The original Angiosperm Phylogeny Group (APG 1998) ushered in a new way of constructing a classification: one based on an explicitly analysed set of data rather than on an intuitive interpretation of available data. Additionally, it was not authored by a single authority, but was based on an informal consensus among many researchers involved in various studies

Rapid radiation

Speciation

Speciation

Figure 7b. Abstract visualisation of evolution based on the same tree

Common ancestor

Plants of the World

9

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP of classification at the time. Data were still scant in the late 1990s, and the primary focus of APG was to place families in orders. Family circumscriptions became more important when the first update of the APG classification was proposed (APG II 2003). This classification included the so-called ‘bracketed system’, in which a researcher could either use a broader or narrower circumscription of several families but still follow the APG classification. This flexibility caused confusion and was therefore abolished in APG III (2009). The APG system is now widely established, and most libraries, herbaria, botanical gardens and online resources follow it. Even though most issues are now settled, the discussion about broader or narrower families still continues in some cases, and, as long as monophyletic groups are recognised, this choice is arbitrary. The fourth update of APG resolved some issues that could not be resolved with available data in 2009, and APG IV (2016) focused more on family delimitation than any of the APG systems did before. Due to an emphasis on stability, APG IV does not differ much from APG III. Apart from some minor additions of previously unplaced or misplaced genera, the last two updates of the APG classifications are practically the same. Only in the treatment of Gesneriaceae we have diverged from APG IV as that system left Peltanthera unplaced, and we propose that it is better placed with Calceolariaceae in an expanded Gesneriaceae. For lycopods, ferns and gymnosperms, similar molecular-based classifications have been proposed. Here, we follow those of Christenhusz & Chase (2014) for lycopods and ferns and Christenhusz et al. (2011) for gymnosperms. To the right, we present formal cladograms at the ordinal level, showing relationships between orders and larger groups. We realise that many people find these difficult to read, especially because they are a two-dimensional depiction of a multidimensional phenomenon. Therefore we also provide a less formal ‘tree of life’, in which the orders are placed on their respective branches and the approximate ages of these branches are given (Figures 8 and 9).

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Lycopods

Ferns

Gymnosperms

Angiosperms

Eudicots

Figure 8. Simplified phylogenetic tree showing relationships of the orders of vascular plants.

Algae Mosses Hornworts Liverworts Lycopodiales Isoëtales Selaginellales Ophioglossales Psilotales Equisetales Marattiales Osmundales Hymenophyllales Gleicheniales Schizaeales Salviniales Cyatheales Polypodiales Zamiales Cycadales Ginkgoales Welwitschiales Gnetales Ephedrales Araucariales Pinales Cupressales Amborellales Nymphaeales Austrobaileyales Chloranthales Piperales Canellales Magnoliales Laurales Acorales Alismatales Petrosaviales Dioscoreales Pandanales Liliales Asparagales Arecales Commelinales Zingiberales Poales Ceratophyllales Ranunculales Proteales Trochodendrales Buxales Gunnerales Dilleniales Saxifragales Vitales Zygophyllales Cucurbitales Fagales Fabales Rosales Oxalidales Celastrales Malpighiales Geraniales Myrtales Crossosomatales Picramniales Sapindales Huerteales Malvales Brassicales Santalales Berberidopsidales Caryophyllales Cornales Ericales Icacinales Metteniusales Garryales Vahliales Gentianales Solanales Lamiales Boraginales Aquifoliales Asterales Escalloniales Bruniales Dipsacales Paracryphiales Apiales

Magnoliids

Monocots

Rosids

Asterids

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP

EUDICOTS FLOWERING PLANTS

Malpighiales Oxalidales Celastrales

Santalales Ericales

Garryales Icacinales

Zygophyllales Vitales

Caryophyllales

Gentianales

100

Solanales

Fagales Cucurbitales Fagales Geraniales Myrtales Rosales Crossosomatales Huerteales 100 Malvales Brassicales Picramniales Sapindales

Vahliales Dilleniales Asterales Boraginales 115 115 100 Lamiales Saxifragales Escalloniales Aquifoliales Bruniales Gunnerales Cornales 120 Paracryphiales Apiales Berberidopsidales Petrosaviales Alismatales Trochodendrales Dipsacales Buxales 130 Proteales Ranunculales

120 Acorales

140

Ceratophyllales

Magnoliales

MAGNOLIIDS Polypodiales Cyatheales Salviniales Gleicheniales Osmundales

Austrobaileyales 150

145

Nymphaeales

Piperales

Canellales Amborellales Schizaeales Hymenophyllales Marattiales 200

336

Psilotales

155 255

245 370 Seed plants

Selaginellales

Lycopodiales

Equisetales 385 Hornworts Mosses Liverworts 484

GYMNOSPERMS Pinales Cupressales Araucariales Gnetales Welwitschiales Ephedrales Ginkgoales Cycadales

Isoëtales

380

Ophioglossales

FERNS

MONOCOTS

Chloranthales

Laurales

Asparagales Arecales Poales Commelinales Liliales Zingiberales Pandanales Dioscoreales

LYCOPODS

Vascular plants 446 290

379

Figure 9. Visualisation of simplified phylogenetic tree showing relationships of the orders of vascular plants. Numbers indicate the approximate ages of branches in millions of years.

Plants of the World

11

CLASSIFICATION AND THE ANGIOSPERM PHYLOGENY GROUP

Diorama of a forest during the Carboniferous with Cordaites (putative relative of Equisetales) Sigillaria (Isoëtales) and Psaronius (Marattiales). University of Michigan Museum of Natural History, Ann Arbor, USA.

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FOSSIL PLANTS When people think of fossils, dinosaur bones mostly come to mind. Of course skeletons and shells are the hardest part of an animal and are the most likely to become fossilised, but some plants also fossilise well even though they have no skeleton. Their cells have rigid walls of cellulose, so it is more likely that woody structures, seeds and leaves are preserved rather than soft tissues. Most of our knowledge of the history of life on earth is derived from fossils, and they are an important key in understanding evolution and relationships between organisms though so-called ‘missing links’, and fossils are essential for estimating ages of clades. Fossil plants are especially important in estimating past climate by comparing distributions of fossil taxa with the ranges of their extant relatives. Also leaf stomata are often well preserved and can be studied to estimate past CO2 concentrations, an invaluable tool in climate reconstruction and understanding past, present and future climate change. After plants become buried in sediment, the spaces left inside the cell wall are sometimes replaced by inorganic material, usually chert, a fine crystalline quartz. This process is called permineralisation, which most commonly occurs in woody tissues (forming petrified wood), but can also happen in herbaceous cells or porous animal skeletons. Sometimes the dead tissue is dissolved, leaving an imprint of the organic structures in the sediment. These impression fossils are often useful for understanding the shapes of soft tissues that do not permineralise well. Sometimes a residue of carbon from the organism remains in the impression fossil, which is called carbonisation. When this happens on a larger scale, as in the swamps of the Carboniferous, this can form coal or, under higher pressure, oil or diamonds. Rare fossils that preserved or show outlines of soft

parts such as flowers (Figure 10) are always preserved under special conditions, such as rapid burial in fine sediment, absence of oxygen or protection in an enclosed structure formed from chemicals emitted by the fossilising organism. Charcoalified plant parts (partially burned) are also an important type of fossilised material. Spores and pollen often preserve particularly well because they are covered in sporopollenin, a substance that does not easily decay and thus is readily preserved in the fossil record, especially under anaerobic conditions, such as lakebeds and swamps. The age of a certain fossil can be estimated from the particular layer, or stratum, in which it was found. Radioactive decay can also be measured to estimate the age of the sediment, and carbonised fossils can be carbon-dated, more precisely so when they are not too ancient. We do not attempt to provide a full overview of the fossil history of each family, as this is a different discipline and not relevant to the general reader interested in living plants, but where appropriate, we mention the most important fossils used for inferring the evolutionary age of the family or morphological evolution. The most important extinct clades are listed and placed tentatively in the classification above. In the Phylogeny and Evolution sections, we discuss previous and current ideas about the relationships of each family. The main changes in circumscription of these families and genera are also mentioned.

Figure 10 A fossil palm (Sabal sp.), Naturalis, Leiden, the Netherlands

Plants of the World

13

FAMILIES A plant family is composed of one or more genera that share some characters and are monophyletic (i.e. they form an exclusive group, with each being more closely related to others in the group than to plants belonging to another group). The definition of a plant family has varied in the past, and some such as daisies (Asteraceae), umbels (Apiaceae), crucifers (Brassicaceae), orchids (Orchidaceae), grasses (Poaceae) and palms (Arecaceae) have a long history of use with little change in concept over the years, whereas others, such as Scrophulariaceae, Liliaceae and Salicaceae have undergone dramatic changes in the number of genera included and their circumscription. In their current circumscriptions, there are 451 vascular plant families when following the classifications of APG IV (2016), Christenhusz & Chase (2014) and Christenhusz et al. (2014). These encompass an approximate total of c. 300,750 species of

which approximately 180,000 are eudicots, 109,000 are monocots, 10,550 are ferns, 1,290 are lycopods and 1,011 are gymnosperms. The ten largest vascular plant families comprise c. 121,180 species, and there is a big gap between the largest five families and the rest. Orchidaceae are probably the largest vascular family, comprising nearly 9% of botanical diversity, followed closely by Asteraceae. It has to be noted that there are still species being described, and this will result in all these numbers slowly increasing over the coming decades.

The 20 largest vascular plant families Family 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

Orchidaceae Asteraceae Fabaceae Rubiaceae Poaceae Lamiaceae Euphorbiaceae Myrtaceae Cyperaceae Melastomataceae Apocynaceae Ericaceae Malvaceae Polypodiaceae Acanthaceae Gesneriaceae Piperaceae Brassicaceae Apiaceae Bromeliaceae

Approximate number of species 26,470 24,700 16,020 13,620 11,000 6,800 6,243 5,828 5,500 5,000 4,300 4,250 4,225 4,070 4,000 3,810 3,700 3,628 3,575 3,475

ETYMOLOGY AND COMMON NAMES According to the International Code of Botanical Nomenclature, each family name is based on a genus, the type genus of that particular family (although due to priority, these names can be considered a later synonym of another genus and thus not in current use, e.g. Hydatella is a later name for Trithuria, and although it is the basis for Hydatellaceae, Hydatella is considered a synonym). For each family we have provided the likely origin of the name of the type genus. These scientific names are usually based on Greek or Latin, but they sometimes use another language or are chosen to honour a person or place. Etymology provides some insight into the

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botanical history of naming genera and families, and the construction/meaning of names may help the novice to get to grips with scientific names. For each family, we also provide a common name based either on the type genus or on a well-known representative of the family. As already stated earlier, common names can vary between areas, and therefore these are not stable. Usually the common name of a family is based on common names of representatives found in English language areas, mainly the UK, USA, South Africa or Australia. This of course causes issues with some common names; for instance for Platanaceae, we

decided not to call it the sycamore family because even though sycamore in America is Platanus occidentalis (Platanaceae), a sycamore in England is Acer pseudoplatanus (Sapindaceae) and in East Africa the name is reserved for Ficus sycomorus (Moraceae). So when names cause confusion, we opted for the least confusing option, in this case “plane tree family” for Platanaceae. In cases where no common names are available, we chose a common name derived from the most widely used vernacular name in another language, if reasonably pronounceable in English. In a few cases, we coined a new name, hoping that these will appeal to the user.

FAMILIES A plant family is composed of one or more genera that share some characters and are monophyletic (i.e. they form an exclusive group, with each being more closely related to others in the group than to plants belonging to another group). The definition of a plant family has varied in the past, and some such as daisies (Asteraceae), umbels (Apiaceae), crucifers (Brassicaceae), orchids (Orchidaceae), grasses (Poaceae) and palms (Arecaceae) have a long history of use with little change in concept over the years, whereas others, such as Scrophulariaceae, Liliaceae and Salicaceae have undergone dramatic changes in the number of genera included and their circumscription. In their current circumscriptions, there are 451 vascular plant families when following the classifications of APG IV (2016), Christenhusz & Chase (2014) and Christenhusz et al. (2014). These encompass an approximate total of c. 300,750 species of

which approximately 180,000 are eudicots, 109,000 are monocots, 10,550 are ferns, 1,290 are lycopods and 1,011 are gymnosperms. The ten largest vascular plant families comprise c. 121,180 species, and there is a big gap between the largest five families and the rest. Orchidaceae are probably the largest vascular family, comprising nearly 9% of botanical diversity, followed closely by Asteraceae. It has to be noted that there are still species being described, and this will result in all these numbers slowly increasing over the coming decades.

The 20 largest vascular plant families Family 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

Orchidaceae Asteraceae Fabaceae Rubiaceae Poaceae Lamiaceae Euphorbiaceae Myrtaceae Cyperaceae Melastomataceae Apocynaceae Ericaceae Malvaceae Polypodiaceae Acanthaceae Gesneriaceae Piperaceae Brassicaceae Apiaceae Bromeliaceae

Approximate number of species 26,470 24,700 16,020 13,620 11,000 6,800 6,243 5,828 5,500 5,000 4,300 4,250 4,225 4,070 4,000 3,810 3,700 3,628 3,575 3,475

ETYMOLOGY AND COMMON NAMES According to the International Code of Botanical Nomenclature, each family name is based on a genus, the type genus of that particular family (although due to priority, these names can be considered a later synonym of another genus and thus not in current use, e.g. Hydatella is a later name for Trithuria, and although it is the basis for Hydatellaceae, Hydatella is considered a synonym). For each family we have provided the likely origin of the name of the type genus. These scientific names are usually based on Greek or Latin, but they sometimes use another language or are chosen to honour a person or place. Etymology provides some insight into the

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botanical history of naming genera and families, and the construction/meaning of names may help the novice to get to grips with scientific names. For each family, we also provide a common name based either on the type genus or on a well-known representative of the family. As already stated earlier, common names can vary between areas, and therefore these are not stable. Usually the common name of a family is based on common names of representatives found in English language areas, mainly the UK, USA, South Africa or Australia. This of course causes issues with some common names; for instance for Platanaceae, we

decided not to call it the sycamore family because even though sycamore in America is Platanus occidentalis (Platanaceae), a sycamore in England is Acer pseudoplatanus (Sapindaceae) and in East Africa the name is reserved for Ficus sycomorus (Moraceae). So when names cause confusion, we opted for the least confusing option, in this case “plane tree family” for Platanaceae. In cases where no common names are available, we chose a common name derived from the most widely used vernacular name in another language, if reasonably pronounceable in English. In a few cases, we coined a new name, hoping that these will appeal to the user.

GENERA In this book we list the accepted genera with estimated numbers of species according to the latest treatments, following the Checklist of Selected Plant Families (http://apps.kew. org/wcsp/home.do) or, when a family is not treated there, other online or printed sources. Of course, as soon as this book is printed, these lists will be outdated: new genera and species will have been described, and new phylogenetic trees will show different configurations forcing reclassifications and inevitable nomenclatural changes. However, we believe that providing generic lists and species numbers will help people in placing genera in the correct family and give them an idea about size, even if it is only an informed estimate. Larger genera (with 100 or more species) are marked in boldface in the lists of genera under each family. The largest genera in the plant world are Astragalus (Fabaceae), Piper (Piperaceae) and Euphorbia (Euphorbiaceae). Segregate or smaller genera have not been widely accepted in these three genera, and therefore they remain large. Delimitation of genera is often based on arbitrarily selected morphological characters, but two options are possible if a genus is found not to be monophyletic: 1) split off one or more new genera to preserve a well-established

genus name for which the species were found to be embedded in another genus (splitting); or 2) enlarge a genus when it is found to be polyphyletic or paraphyletic and include genera found to be embedded in it (lumping), which is what has been done with most of the largest genera in the table (right). We advocate this for some groups in which genera are small and now finely split. This of course depends on the taxonomists working on the group and acceptance of new names by the users of these names. The species numbers for Sedum and Thelypteris given below are those if they are to be taken in their broad sense. Similarly, the large genus Alchemilla (Rosaceae) is embedded in Potentilla (along with several others, such as strawberry, Fragaria), and Potentilla can be divided. However, it may be better to expand it, which would then give rise to a genus that should be included in our list of large genera (right); for this to be formally accepted, many transfers of species from other genera to Potentilla would first be needed. Generic circumscription is a human construct and a matter of stability, historical concepts, morphological, genetic uniformity, and most of all a matter of opinion. Therefore the discussion between ‘lumpers’ and ‘splitters’ will likely be a never-ending story.

The largest genera of vascular plants Number of species

Family

Genus

Fabaceae

Astragalus

c. 2,400

Piperaceae

Piper

c. 2,100

Euphorbiaceae

Euphorbia

1,933

Orchidaceae

Bulbophyllum

1,867

Rubiaceae

Psychotria

1,858

Cyperaceae

Carex

1,829

Begoniaceae

Begonia

1,803

Asteraceae

Senecio

Piperaceae

Peperomia

Orchidaceae

Dendrobium

Orchidaceae

Epidendrum

Solanaceae

Solanum

Euphorbiaceae

Croton

1,186

Myrtaceae

Syzygium

1,136

Aspleniaceae

Thelypteris s.l.

Orchidaceae

Lepanthes

1,085

Fabaceae

Acacia s.s.

1,075

Myrtaceae

Eugenia

Balsaminaceae

Impatiens

1,633 c. 1,600 1,509 1,413 c. 1,300

c. 1,100

1,044 c. 1,000

Ericaceae

Rhododendron

c. 1,000

Melastomataceae

Miconia

c. 1,000

Phyllanthaceae

Phyllanthus

c. 1,000

Rosaceae

Alchemilla

c. 1,000

Crassulaceae

Sedum s.l.

979

Amaryllidaceae

Allium

920

Lamiaceae

Salvia

c. 900

PHYTOGEOGRAPHY Phytogeography is the study of plant distributions, both historical and modern. When distributions and phylogenetic history are known, the current distribution can sometimes be explained. On the basis of georeferenced herbarium specimens, detailed maps can be made of where species have been found, and these data can be used to

make ecological niche models and to predict current, past and future distributions under changing climatic conditions. The maps provided here are not detailed, but for each family we provide their rough native range. Even though these ranges are rough estimates, they are based on known occurrences, and it is possible that occasional

finds of family members outside the given range occur. This may be due to occasional new finds, species naturalised from horticulture or agriculture or perhaps even range changes due to climate change. When possible, the area of largest diversity is provided in the text. These maps are just a general indication of where plants occur and are by no means precise. For

Plants of the World

15

GENERA In this book we list the accepted genera with estimated numbers of species according to the latest treatments, following the Checklist of Selected Plant Families (http://apps.kew. org/wcsp/home.do) or, when a family is not treated there, other online or printed sources. Of course, as soon as this book is printed, these lists will be outdated: new genera and species will have been described, and new phylogenetic trees will show different configurations forcing reclassifications and inevitable nomenclatural changes. However, we believe that providing generic lists and species numbers will help people in placing genera in the correct family and give them an idea about size, even if it is only an informed estimate. Larger genera (with 100 or more species) are marked in boldface in the lists of genera under each family. The largest genera in the plant world are Astragalus (Fabaceae), Piper (Piperaceae) and Euphorbia (Euphorbiaceae). Segregate or smaller genera have not been widely accepted in these three genera, and therefore they remain large. Delimitation of genera is often based on arbitrarily selected morphological characters, but two options are possible if a genus is found not to be monophyletic: 1) split off one or more new genera to preserve a well-established

genus name for which the species were found to be embedded in another genus (splitting); or 2) enlarge a genus when it is found to be polyphyletic or paraphyletic and include genera found to be embedded in it (lumping), which is what has been done with most of the largest genera in the table (right). We advocate this for some groups in which genera are small and now finely split. This of course depends on the taxonomists working on the group and acceptance of new names by the users of these names. The species numbers for Sedum and Thelypteris given below are those if they are to be taken in their broad sense. Similarly, the large genus Alchemilla (Rosaceae) is embedded in Potentilla (along with several others, such as strawberry, Fragaria), and Potentilla can be divided. However, it may be better to expand it, which would then give rise to a genus that should be included in our list of large genera (right); for this to be formally accepted, many transfers of species from other genera to Potentilla would first be needed. Generic circumscription is a human construct and a matter of stability, historical concepts, morphological, genetic uniformity, and most of all a matter of opinion. Therefore the discussion between ‘lumpers’ and ‘splitters’ will likely be a never-ending story.

The largest genera of vascular plants Number of species

Family

Genus

Fabaceae

Astragalus

c. 2,400

Piperaceae

Piper

c. 2,100

Euphorbiaceae

Euphorbia

1,933

Orchidaceae

Bulbophyllum

1,867

Rubiaceae

Psychotria

1,858

Cyperaceae

Carex

1,829

Begoniaceae

Begonia

1,803

Asteraceae

Senecio

Piperaceae

Peperomia

Orchidaceae

Dendrobium

Orchidaceae

Epidendrum

Solanaceae

Solanum

Euphorbiaceae

Croton

1,186

Myrtaceae

Syzygium

1,136

Aspleniaceae

Thelypteris s.l.

Orchidaceae

Lepanthes

1,085

Fabaceae

Acacia s.s.

1,075

Myrtaceae

Eugenia

Balsaminaceae

Impatiens

1,633 c. 1,600 1,509 1,413 c. 1,300

c. 1,100

1,044 c. 1,000

Ericaceae

Rhododendron

c. 1,000

Melastomataceae

Miconia

c. 1,000

Phyllanthaceae

Phyllanthus

c. 1,000

Rosaceae

Alchemilla

c. 1,000

Crassulaceae

Sedum s.l.

979

Amaryllidaceae

Allium

920

Lamiaceae

Salvia

c. 900

PHYTOGEOGRAPHY Phytogeography is the study of plant distributions, both historical and modern. When distributions and phylogenetic history are known, the current distribution can sometimes be explained. On the basis of georeferenced herbarium specimens, detailed maps can be made of where species have been found, and these data can be used to

make ecological niche models and to predict current, past and future distributions under changing climatic conditions. The maps provided here are not detailed, but for each family we provide their rough native range. Even though these ranges are rough estimates, they are based on known occurrences, and it is possible that occasional

finds of family members outside the given range occur. This may be due to occasional new finds, species naturalised from horticulture or agriculture or perhaps even range changes due to climate change. When possible, the area of largest diversity is provided in the text. These maps are just a general indication of where plants occur and are by no means precise. For

Plants of the World

15

PHYTOGEOGRAPHY Global Biodiversity: Species numbers of vascular plants © Barthlott 2014

example, Amborella (Amborellaceae) does not grow everywhere on New Caledonia. It is not uncommon in the upland forest, so the entire island is coloured, even though it is only found in a few places. If one overlays these distribution maps (Figure 11), we would get a map showing the hotspots of family diversity. The main areas are obviously in the tropics, but it should be pointed out that warm temperate areas like southwestern Australia, southern Africa and southern China also have remarkably diverse floras. This phylogenetic diversity agrees fairly well with the known hotspots of biodiversity (Myers et al. 2000), and these areas should be given priority for botanical conservation. Phylogenetic diversity is important, and thus when a family is restricted to a small area it is important that this area is considered for protection. Since such a family is unlikely to have close relatives, if it goes extinct, an entire lineage would be lost.

Diversity zones (DZ), number of vascular plant species per 10,000 km² DZ 1 ( 5000) Barthlott, W., Rafiqpoor, M.D., & Mutke, J. 2014 based on former publication starting with Barthlott, W., Lauer, W. & Placke, A. 1996

Figure 11. Overlay of family distributions, showing the areas of the world with the greatest diversity of vascular plant families.

ECONOMIC BOTANY Plants are the foundation of civilisation. We need the oxygen and clean air that plants produce, and we need fruits, vegetables, other foods, medicinal compounds and fibre, all of these are clear. What few people also realise is that most of our culture and economy is based on plants, the main exception being wind, wave and solar energy generation, the last imitating the capacity of plants to harvest energy from the sun. Plants underlie every part of our economy; plants form the basis of agriculture, banking, building, education, fashion, fishery, fossil fuels, health care, industry, medicine, trade, transportation etc. Botany should receive a much more central role in general research and research funding than it currently is given. Considering how much we still do not know about these organisms that make the world function, there is a great need to study the biodiversity of this planet, and plant diversity should be the highest priority.

16

Christenhusz, Fay & Chase

This book provides an overview of plant diversity on Earth, and for each family we provide a list of some economic uses, showing the diversity of products that different plant groups provide humans, including food (cereals, fruit, vegetables, nuts, tubers, grains), animal fodder (Figure 12), medicines, perfumes, oils, dyes, ornamental plants, timber, fibre, clothes and paper. We focus mainly on the common foods and crops, but also include products that are still harvested from the wild. Even though the enumeration is fairly exhaustive, we omit many medicinal plants simply because there are so many, particularly in Indian and Chinese traditional medicines, and there are other good books that focus on traditional medicinal usage of plants. We nevertheless hope to illustrate the great diversity of uses and incredible economic importance of botany, hopefully making plants better understood and more valued by the wider public. Plants should be studied, valued and loved.

Figure 12. Humans have altered their landscape for millennia.

Right: Fruit and vegetable market in Bogota, Colombia, showing a wide diversity of plant products available to humans

PHYTOGEOGRAPHY Global Biodiversity: Species numbers of vascular plants © Barthlott 2014

example, Amborella (Amborellaceae) does not grow everywhere on New Caledonia. It is not uncommon in the upland forest, so the entire island is coloured, even though it is only found in a few places. If one overlays these distribution maps (Figure 11), we would get a map showing the hotspots of family diversity. The main areas are obviously in the tropics, but it should be pointed out that warm temperate areas like southwestern Australia, southern Africa and southern China also have remarkably diverse floras. This phylogenetic diversity agrees fairly well with the known hotspots of biodiversity (Myers et al. 2000), and these areas should be given priority for botanical conservation. Phylogenetic diversity is important, and thus when a family is restricted to a small area it is important that this area is considered for protection. Since such a family is unlikely to have close relatives, if it goes extinct, an entire lineage would be lost.

Diversity zones (DZ), number of vascular plant species per 10,000 km² DZ 1 ( 5000) Barthlott, W., Rafiqpoor, M.D., & Mutke, J. 2014 based on former publication starting with Barthlott, W., Lauer, W. & Placke, A. 1996

Figure 11. Overlay of family distributions, showing the areas of the world with the greatest diversity of vascular plant families.

ECONOMIC BOTANY Plants are the foundation of civilisation. We need the oxygen and clean air that plants produce, and we need fruits, vegetables, other foods, medicinal compounds and fibre, all of these are clear. What few people also realise is that most of our culture and economy is based on plants, the main exception being wind, wave and solar energy generation, the last imitating the capacity of plants to harvest energy from the sun. Plants underlie every part of our economy; plants form the basis of agriculture, banking, building, education, fashion, fishery, fossil fuels, health care, industry, medicine, trade, transportation etc. Botany should receive a much more central role in general research and research funding than it currently is given. Considering how much we still do not know about these organisms that make the world function, there is a great need to study the biodiversity of this planet, and plant diversity should be the highest priority.

16

Christenhusz, Fay & Chase

This book provides an overview of plant diversity on Earth, and for each family we provide a list of some economic uses, showing the diversity of products that different plant groups provide humans, including food (cereals, fruit, vegetables, nuts, tubers, grains), animal fodder (Figure 12), medicines, perfumes, oils, dyes, ornamental plants, timber, fibre, clothes and paper. We focus mainly on the common foods and crops, but also include products that are still harvested from the wild. Even though the enumeration is fairly exhaustive, we omit many medicinal plants simply because there are so many, particularly in Indian and Chinese traditional medicines, and there are other good books that focus on traditional medicinal usage of plants. We nevertheless hope to illustrate the great diversity of uses and incredible economic importance of botany, hopefully making plants better understood and more valued by the wider public. Plants should be studied, valued and loved.

Figure 12. Humans have altered their landscape for millennia.

Right: Fruit and vegetable market in Bogota, Colombia, showing a wide diversity of plant products available to humans

LYCOPODS (LYCOPODIIDAE) Lycopods or clubmosses are the sister lineage to all other vascular plants (ferns plus seed plants) and are among the oldest lineages of living plants. Extant species are, however, highly divergent from the early members of this group. They were diverse in the Carboniferous, when some species formed large trees dominating the ancient forests. Modern lycopods are all low-growing herbaceous plants, with some scrambling Selaginella reaching the height of only a few metres. Isoëtes retains some of this anatomical diversity and has secondary (woody) growth in its short stem. Like ferns, lycopods have a clearly alternating life cycle, with sporophytes being the large plants we commonly observe. These make spores from which gametophytes grow upon germination and produce free-swimming sperm. This makes these plants dependent on water, although many sporophytes have evolved strategies to deal with drought. Sporophytes have roots and stems that

bear spirally arranged or whorled leaves called microphylls, served only by a single vein (except for a few Selaginella species). Lycopods differ from ferns and the seed plants in having microphylls instead of macrophylls (leaves supplied with several veins). Stems are usually protostelic or sometimes polystelic and sporangia are either born on a microphyll or are associated with one. Often placed among ‘fern allies’, the correct name for this lineage is lycopodiophytes or lycopods, not ‘lycophytes’ as often erroneously used. Due to their life-history strategy, which is like that of the ferns, and their relatively inconspicuous nature, most people think of them as ‘fern allies’, but they are equally closely related to the seed plants. General references: Ambrose BA. 2013. The morphology and development of lycophytes. Annual Plant Review 45: 91–114. Boyce CK. 2005. The evolutionary history of roots and leaves. pp. 479–499 in Holbrook

NM, Zwienieckl MA (eds). Vascular transport in plants. Elsevier, Amsterdam. Christenhusz MJM, Zhang X-C, Schneider H. 2011. A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. Horn K, Franke T, Unterseher M, Schnitter M, Beenken L. 2013. Morphological and molecular analysis of fungal endophytes of achlorophyllous gametophytes of Diphasiastrum alpinum (Lycopodiaceae). American Journal of Botany 100: 2158–2174. McLoughlin S, Jansson I-M, Vajda V. 2014. Megaspore and microfossil assemblages reveal diverse herbaceous lycophytes in the Australian Early Jurassic flora. Grana 53: 22–53. Ranker TA, Hauf ler CH. 2008. Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Renzaglia KS, Whittier DP. 2013. Microanatomy of the placenta of Lycopodium obscurum: novel design in an underground embryo. Annals of Botany 112: 1083–1088.

LYCOPODIALES The order is comprised of a single family, a group of herbaceous plants known collectively as the clubmosses and firmosses, all with dichotomously branching stems. They have exosporic gametophytes like those of ferns.

1. LYCOPODIACEAE Clubmoss family

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These plants are erect or pendulous, terrestrial or epiphytic, homosporous herbs. Stems are dichotomously branched and protostelic. Microphylls (simple leaves with a single unbranched vein) are spirally arranged or whorled, sometimes decussate. All leaves are similar or those bearing sporangia differ from the sterile ones; the latter can be aggregated in distinct strobili (‘cones’). Sporangia are kidney-shaped or globular, dehiscing by a slit, and are formed singularly in the leaf

axils. Spores are trilete and subglobose to tetrahedral. Gametophytes are tuberous, mycoheterotrophic when subterranean or photosynthetic when surface-living. Distribution: A cosmopolitan family, but absent from arid regions. Phylogeny and evolution: Lycopodiaceae have no close affinity with any other group of plants; even the relationship with

LYCOPODIALES

LYCOPODS

Selaginellaceae and Isoëtaceae is ancient. Fossil Lycopodiaceae cannot be referred with certainty to extant groups, generally due to their poor preservation. The group dates back to at least the Carboniferous when tree-like members of this lineage thrived and dominated the terrestrial sphere. The extant members of the family form three (to five) clades corresponding to the genera accepted below. Some authors prefer to divide these lineages further, resulting in up to 14 genera, but many of these are difficult to distinguish morphologically. The peculiar Huperzia drummondii (formerly Phylloglossum drummondii) differs in having a subterranean tuber with a rosette of leaves and a single strobilus on a naked peduncle. This morphology is an adaptation to the seasonally dry region in its native Australia, but otherwise it fits in the genus Huperzia where it is now placed.

Huperzia drummondii, Western Australia [1]

Genera and species: Lycopodiaceae consist of three genera with c. 400 species: Huperzia (c. 300), Lycopodiella (c. 40) and Lycopodium (c. 40). Uses: Due to their high flammability, the oil-rich spores of stag’s-horn clubmoss (Lycopodium clavatum) and related species are harvested and sold as ‘lycopodium powder’. When mixed with air it creates an explosive mixture, which is widely used in special effects in the film industry. It was also previously used as a fine dusting on latex products such as condoms and surgical gloves, as a coating of metal moulds to prevent metal from sticking to them when casting the iron, in pharmaceuticals as a pill coating and as a cosmetic powder in make-up. Exposure to Lycopodium spores may cause allergic

Huperzia crassa, Ecuador [1]

Lycopodiella torta, Guadeloupe [1]

Huperzia ribourtii, Tahiti [1]

reactions ranging from mild dermatitis to severe asthma, so its uses involving human contact such as cosmetics have largely disappeared. Spores of Alpine clubmoss (Lycopodium alpinum) have been used in Scandinavia to produce a pale yellow dye in combination with twigs of bog whortleberry (Vaccinium uliginosum, Ericaceae). Several Lycopodium species have been used to weave mats, hats, bags and fish traps. Etymology: Lycopodium is composed of the Greek λύκος (lykos), wolf and ποδιών ( podion), feet, referring to the fuzzy, leafclad branches. Hence, the common name, lycopods (‘wolf claws’ is the translation of the common name in many other languages), for the whole group is appropriate, whereas lycophytes (wolf plants) is not.

Huperzia sieberiana, Guadeloupe [1]

Lycopodiella inundata, Twickel, the Netherlands [1]

Plants of the World

19

ISOËTALES

LYCOPODS

ISOËTALES This order with secondary growth is considered to be sister to the fossil Lepidodendrales, which were giant trees during the Carboniferous. These together are sister to the strictly herbaceous Selaginellales, with which they share heterospory (two sizes of spores) and endosporic gametophytes.

2. ISOËTACEAE Quillwort family

These aquatic or terrestrial heterosporous plants are evergreen or deciduous. Their woody stems (rhizomes) are covered with irregular ‘bark’ from which bulbils often develop. Leaves are spirally arranged, needle-like, filiform, flat or terete, forming a rosette or a tuft. The vascular bundle in the leaf is surrounded by four air canals. Ligules arise from a sac at the leaf base, above the sporangium. Sporangia are globose to ovoid

and are formed above the leaf base and have no dehiscence mechanism. Spores are of two kinds (heterosporous) and are usually formed in separate sporangia. Megaspores are trilete and variously ornamented, whereas microspores are monolete and patterned. Gametophytes are formed within the megaspores (endosporic), which are released and thus considered to be free-living. Microspores produce sperm only. Distribution: Isoëtaceae have a global distribution. They are usually found in clean, low-nutrient (oligotrophic) lakes, streams and ephemeral pools or are terrestrial in humid alpine vegetation. Phylogeny and evolution: The genus Isoëtes forms two clades, one including only two Andean terrestrial species (subgenus Stylites), which differ in their erect stems. Even though most species are aquatic, many

Isoëtes lacustris from C. Johnson (1856) The Fern Allies: a supplement to the Ferns of Great Britain, London, plate 13.

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Christenhusz, Fay & Chase

occur in temporary bodies of water, surviving the dry periods as dormant corms. Fossils attributable to Isoëtes have been found in the Triassic (as Isoëtites), many of which are described from spores only. Genera and species: Isoëtaceae include the single genus Isoëtes with c. 140 species. Uses: The corms of some species are consumed by birds, fish, muskrats and pigs. They contain oils and starch and are not poisonous, but do not have a flavour pleasant enough to be worthy of human consumption. Etymology: Isoëtes is derived from the Greek ίσος (isos), equal, and ετήσ (etes), yearly, meaning equal at all seasons of the year, referring to some evergreen species. The diacritical sign is used to indicate that the o and e are to be pronounced as separate vowels.

Isoëtes drummondii, Perth, Western Australia [2]

SELAGINELLALES

LYCOPODS

SELAGINELLALES This order is comprised of a single family, which previously had been thought to be more closely related to Lycopodiaceae (based on their shared primary growth), but they are instead more closely related to Isoëtales, with which they share heterospory and endosporic gametophytes.

3. SELAGINELLACEAE Spikemoss family

These terrestrial, rarely epiphytic, perennial, heterosporous plants have forked roots (rhizophores). Their main stems are erect or prostrate and not distinguished into rhizomes, but some species with erect branches (pseudofronds) spread by creeping stems. The microphylls (leaves) are spirally arranged, minute, with a ligule, and are all similar or of two kinds (then arranged into four ranks). Fertile leaves are leaf-like and aggregated Selaginella selaginoides, Lake District, UK [3]

in terminal strobili (‘cones’). Sporangia are stalked, placed on the upper surface of fertile leaves, just above the ligule. Spores are of two kinds (heterosporous): megaspores are variously shaped and ornamented, trilete, about four per sporangium, whereas microspores are globose-ellipsoid, variously ornamented, trilete and over 100 per sporangium. Distribution: Spikemosses are distributed worldwide, but with their highest diversity in the tropics. A few species occur in the Arctic, and some can even be found in desert areas, where they are “resurrection plants”: plants that dry to within a small percentage of their fresh weight, but then revive and grow again once provided with water. Phylogeny and evolution: Among arborescent lycopodiophytes of the Carboniferous, some heterosporous herbaceous plants gave rise to Selaginellaceae. The morphogenus

Selaginella lepidophylla, Mexico [3]

Selaginella flabellata, Guadeloupe [3]

Selaginella willdenowii with iridescent leaves, Singapore [3]

Pseudodendron, a common fossil in the Carboniferous that shares some characters with Sigillaria, is now regarded as a species of Selaginellaceae, but the majority of fossils from this family have been described from Cretaceous and Tertiary megaspores. The genus has been divided into five subgenera, but molecular studies have not corroborated these. The DNA markers (plastid genes) that have been used thus far have unusually high substitution rates in this genus, which may impede the reliability of the phylogenetic results. Selaginella moellendorfii has the smallest genome measured so far in vascular plants. A few species have more complex venation but are derived from a group with simple-veined microphylls. Therefore the macrophyll condition evolved independently several (at least two) times. Genera and species: Selaginellaceae consist of the single genus, Selaginella, with c. 750 species. Uses: Some Selaginella species are grown as ornamentals, most notably S. uncinata and S. willdenowii, both with remarkable iridescent leaves. Selaginella lepidophylla is sold bare-root in its dry state as ‘Rose of Jericho’ (although being native to Mexico). This species can be resuscitated when placed in water (similar to Anastatica, Brassicaceae, from the Middle East). Young shoots of S. tamariscina are cooked and eaten in East Asia. Etymology: Selaginella is the diminutive form of selago, Latin for a plant resembling savin (Juniperus sabina). The genus Selago is currently in use for a genus of Plantaginaceae; it also is the epithet of the fir clubmoss (Huperzia selago, Lycopodiaceae), which has some general resemblance to the type species of Selaginella, club spikemoss (Selaginella selaginoides), which is probably how the generic name originated. Plants of the World

21

FERNS Ferns are vascular plants that lack f lowers and are remarkable among plants in having two alternating freeliving generations, called gametophytes and sporophytes. Mosses, gymnosperms and angiosperms (seed plants) also have alternating generations, but these are not both free-living. In mosses and liverworts, sporophytes often lack chlorophyll and are then parasitic on free-living gametophytes. By contrast, the reduced gametophytes of seed plants are dependent (parasitic) on the sporophytes; they are present but never released into the environment. Lycopods have a life cycle that is similar to ferns and were therefore often included in the ‘fern-allies’ together with a number of other groups that are now known to be ferns (e.g. Equisetaceae, Psilotaceae and Ophioglossaceae). Lycopods differ from ferns in having simple one-veined leaves called microphylls, instead of the more complex types (macrophylls) found in ferns. In ferns and lycopods, gametophytes are free-living, although they may be subterrean and provided with

Gametophytes (arrow) and young sporophytes of Danaea geniculata (Marattiaceae) on a damp forest bank near Tapirai, Brazil

22

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carbohydrates through a mycorrhizal (fungal) association. They grow from a spore and are usually reduced thalloid structures, dependent on a humid environment due to the lack of true roots and their thin, flat body. They usually resemble a liverwort (a thallus) on which the gametangia, or gameteproducing organs, are produced (female archegonia and male antheridia). On the surface of the gametophyte, the archegonia have a female cell that produces an egg, and the antheridia produce free-swimming f lagellate sperms; most gametophytes are bisexual and others unisexual. When sufficient water is available, the sperm swims into the archegonia, and after fertilisation a sporophyte develops, leaving the gametophyte to wither away. When suitable conditions for fertilisation are not present, gametophytes may live for a long time and even reproduce asexually, as in some species of Hymenophyllaceae and Pteridaceae that have persisted for centuries or even millennia outside their ideal climatic region. Some species have tuberous gametophytes that lack chlorophyll, live underground and take all their nutrients from fungi. Sporophytes are the plants everyone generally knows as ‘ferns’. These are usually large complex vascular plants that produce spores. Spores are formed in groups called sporangia. In earlybranching ferns (families 4 to 7) sporangia develop from a group of cells. These are often called ‘eusporangiate ferns’, a term that is better avoided because it is simply a descriptive term referring to all groups that are not leptosporangiate; the latter group of ferns are sister to one family of eusporangiate ferns, Osmundaceae. Leptosporangiate ferns are all classified in subclass Polypodiidae and have sporangia that are formed from a single

cell. Sporangia are normally grouped in a sorus, which in most leptosporangiate ferns is covered by a scale-like structure called the indusium. Often sporangia are surrounded by an annulus, a structure that aids in ejecting the spores due to it drying unevenly and rupturing he sporangium wall. Leptosporangiate ferns comprise the vast majority of extant fern species. The oldest fern fossils are found in Carboniferous deposits, when the lineage soon became dominant. They, with lycopods, fixed carbon in fossil oiland coal-rich deposits, which are now mined for energy, causing an increase of carbon in the modern atmosphere, resulting in a greenhouse effect and a changing climate, similarly to how these plants changed the climate when they fixed the atmospheric carbon into the sediment several hundred million years ago. The rise of angiosperms during the Cretaceous is often portrayed as causing a dramatic decline of fern and gymnosperm lineages. However, most modern lineages of ferns show an increase in diversity in response to the increase in habitat diversity created by the diversification and formation of angiosperm-dominated rainforests. For other groups of ferns and fernlike plants, those that produced large forest trees, a decline associated with the rise of the seed plants, especially the angiosperms, may in fact be the case. Because ferns have neither flowers nor seeds, numerous legends surrounded the reproduction of ferns. In the Middle Ages, people believed that fern seeds existed, but that they were invisible and were supposed to give the bearer magical powers, especially invisibility. Seeds only develop on the longest day of the year when they would be difficult to harvest because the devil

FERNS

would, of course, want these powers of invisibility. For the same reason, ferns were considered sacred by early Celtic and Germanic peoples. How to collect fern seeds? You could probably try to shoot an arrow into the sun and if hit, drops of blood would fall on a fern and turn into fern seed. A dozen pewter plates could also serve to catch fern seed, but if pewter plates were not available, a white sheet or even a white napkin might do the trick as well, as long as you were in the right frame of mind. Of course, no one ever saw ‘fern seeds’. These and many other legends arose around the gathering of fern seeds, but it was not until the 18th and early 19th century that germination of fern spores, development and fertilisation of gametophytes and development of sporophytes were discovered. This late discovery of the sexual system of ferns resulted in a poor understanding of the relationships among ferns. In the early days they were generally classified on the basis of their soral structures, but it was soon found that this was artificial and other characters such as stem and leaf anatomy had to be employed. With the dawning of the molecular age in the 1990s, DNA sequences could be compared and ferns were reclassified. The classification of ferns has long suffered due to a lack of consensus as to what constitutes a family comparable to those of the seed plants, and family concepts in ferns have long been unstable and subject to great debate. Here we follow a recent classification of ferns (Christenhusz & Chase 2014). The two major lineages in the ‘eupolypods’ are treated as single large families to prevent recognition of numerous monogeneric families. The earlier classifications of Smith et al. (2006) and Christenhusz et al. (2011) treated eupolypods in a number of small families, which have found some acceptance and are preferred by some, but there has been resistance in following the smaller families segregated

from Woodsiaceae and Dryopteridaceae due to polyphyly of these lineages. However, these families are treated here fully as subfamilies, so that both systems can be applied if preferred. One issue of the system used here is that Aspleniaceae are difficult to define with the inclusion of Thelypteridoideae. They can only be defined now by having two vascular bundles in the petiole, even though the indusium placement also may be a synapomorphy. Expanded Polypodiaceae are problematic due to the inclusion of Hypodematioideae that also have two vascular bundles (and are thus similar to Aspleniaceae), but molecular studies place Hypodematioideae near the basal node of Polypodiaceae s.l., and could possibly be maintained separate if synapomorphies are required. The larger families are, however, in line with the currently widely accepted broad concept of Pteridaceae, which is similarly difficult to define morphologically, but is now widely accepted. Where family limits are drawn is a matter of convention and taste, and time will tell which system users will find easiest to apply. We hope that this encyclopedia will shed light on how ferns are interrelated and showcase the fantastic diversity of this unusual group of plants. With an estimated 11,000 species in 20 families, ferns make up only 2% of vascular plant species. Occurring worldwide, they are most diverse in the wet subtropics and tropical mountains at middle elevations. General references: Bower FO. 1923–1928. The ferns, vols. 1–3. Cambridge University Press, Cambridge. Boyce CK. 2005. Patterns of segregation and convergence in the evolution of fern and seed plant leaf morphologies. Paleobiology 31: 117–140. Camus GM, Gibby M, Johns RJ (eds). 1995. Pteridology in perspective. Kew Publishing, Richmond. Christenhusz MJM, Chase MW. 2014. Trends and concepts in fern classification. Annals of Botany 113: 571–594.

Lehtonen S. 2011. Towards resolving the complete fern tree of life. PLoS One 6: e24851. Manton I. 1958. Chromosomes and fern phylogeny with special reference to Pteridaceae. Journal of the Linnean Society, Botany 56: 73–92. Ranker TA, Haufler CH. (eds). 2008. Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Schneider H. 2013. Evolutionary morphology of ferns (monilophytes). Annual Plant Reviews 45: 115–140. Schneider H, Schuettpelz E, Pryer KM, Cranfill R, Magallón S, Lupia R. 2004. Ferns diversified in the shadow of angiosperms. Nature 428: 553–557. Tryon RM, Tryon AF. Ferns and allied plants with special reference to tropical America. Springer, New York. Verdoorn F (ed). 1938. Manual of pteridology. Nijhof, The Hague.

Thelypteris hydrophila crozier showing pneumatophores, Guadeloupe

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EQUISETALES

FERNS

EQUISETALES This order formerly included a number of early fossil taxa like the giant Calamites trees, but it is now believed that modern Equisetales are more closely related to ferns, whereas most ancient fossils probably formed a lineage of their own called the Sphenophytes, although equisetalean fossils were also dominant in the Carboniferous and Devonian. Equisetales are easily recognised by their jointed, leafless stems and terminal strobili.

4. EQUISETACEAE Horsetail family

(elaters), which aid dispersal. The gametophytes are green and live on the soil surface, and male gametes are free-swimming with several flagellae, whereas female gametes are sessile on the gametophyte. Distribution: Equisetaceae mostly occur in the temperate Northern Hemisphere and South America, but one species extends into the Old World tropics. Some species are naturalised as agricultural weeds, especially in the temperate areas.

Horsetails are herbaceous terrestrial plants with hollow stems and underground hollow rhizomes called stolons. The leaves are reduced to sheaths that surround the nodes. Typically in most species, stems branch regularly in whorls around the base of the sheaths, but irregular branching and no branching also frequently occur. The spores are formed in sporangia that form on umbrella-like stalked structures that are organised in cone-shaped strobili (‘cones’) on the ends of otherwise vegetative branches, or, in some species, on separate strictly strobilus-bearing non-green shoots. The spores are green and numerous (> 1,000 per sporangium) and have several ‘legs’

Equisetum myriochaetum, Guatemala [4]

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Phylogeny and evolution: The ancestors of horsetails were dominant trees in the vast swamp-forests during the Carboniferous and Devonian periods. Only a single genus remains of this formerly diverse group of plants, and it has been suggested that it is the oldest genus of extant vascular plants. Because of its ancient origin and the lack of characters comparable with other vascular spore-bearing plants, it has often been treated as a class on its own or grouped with Psilotaceae and lycopods as ‘fern allies’. Molecular evidence places the family on a long branch sister to Ophioglos-

Equisetum arvense, Helsinki Botanical Garden, Finland [4]

saceae+Psilotaceae with poor support. It is nevertheless part of the fern clade. Genera and species: A family composed of a single isolated genus, Equisetum, with c. 20 species, hybridisation between many species blurs their delimitation. Uses: Because of the high concentration of silicates, the plants are poisonous in large quantities, but young shoots are consumed in some traditional societies, usually after a complicated preparation process. Marestail (Equisetum arvense) has been used as a diuretic in Europe, but there is little scientific evidence for its effectiveness. It may, however, be a good source of antioxidants. The young stalks have been eaten as a vegetable since Roman times, and some species (e.g. E. arvense, E. laevigatum) were used as food by native Americans. Strobili of E. braunii are reported to be juicy and sweet. Field horsetail (Equisetum arvense) can be a nuisance as a garden weed in Europe and North America and, as an introduction, in South America, Australia and New Zealand. American scouring rush (E. praealtum;

Equisetum bogotense, Ecuador [4]

OPHIOGLOSSALES

FERNS

Botrychium lunaria, Scotland, UK [5]

Botrychium virginianum, Michigan, USA [5]

marketed as E. ‘ japonicum’) and E. scirpoides are sometimes cultivated as pond-margin ornamentals. Like most horsetails, they prefer wet sandy soils. The stems of European scouring rush, Equisetum hyemale, are loaded with silicates

Ophioglossum vulgatum, private garden, A new species of Ophioglossum Kingston upon Thames, Surrey, UK [5] from New Caledonia [5]

and were used historically as an abrasive for scrubbing pots and polishing wood. Equisetum arvense can also be used as a source of green dye, whereas wood horsetail (E. sylvaticum) is used in Scandinavia to produce a grey-yellow dye.

Etymology: Equisetum is derived from the Latin equus, horse, and seta, bristle, in reference to the coarse black roots of the type species, Equisetum fluviatile, that resemble the tail of a horse.

OPHIOGLOSSALES Even though there is practically no fossil record for this order, molecular clock studies suggest this order diverged c. 160 million years ago. The order is composed of the single family Ophioglossaceae, which some authors in the past have not considered to be closely related to ferns, thanks largely to their non-circinnate vernation.

5. OPHIOGLOSSACEAE Adder’s-tongue family

These terrestrial or epiphytic ferns are usually small (rarely larger than 30 cm) and fleshy with a nodding vernation (not rolled as usual in ferns). Rhizomes and petioles are succulent and have secondary (woody) growth, and roots lack root hairs. Leaves emerge singly (or a few) per growing season and are usually divided in a sterile leaf-like part and a basally or centrally attached non-leafy fertile portion (called the sporophore). Venation of the leaves is free in species with divided

blades and reticulate in simple-leaved species. Sporangia are composed of many cells (eusporangiate), globose and marginal or terminal. They contain thousands of spores and because they lack an annulus, the sporangia open along a lateral or vertical line. The spores are globose-tetrahedral and trilete. Gametophytes are tuberous, subterranean, fleshy and achlorophyllous (they are mycoheterotrophic). Distribution: Ophioglossaceae have a nearly worldwide distribution but are excluded from some of the driest habitats. Phylogeny and evolution: Ophioglossaceae stand alone morphologically, but molecular results indicate a relationship with the equally peculiar Psilotaceae. Both families lack root hairs and have similar mycoheterotrophic gametophytes. Morphological studies have suggested a relationship to the extinct lineage of progymnosperms, but there is no fossil evidence to support this. In fact, despite the

presumed ancient origin of the family, few fossils of Ophioglossaceae are known. Genera and species: Ophioglossaceae consist of four genera and c. 80 species: Botrychium (c. 50), Helminthostachys (1), Mankyua (1) and Ophioglossum (c. 30). Uses: Moonwort (Botrychium lunaria) has a long tradition as a herb to treat wounds, and several other Botrychium and Ophioglossum species are also used for this purpose. Young shoots and leaves of Botrychium lunaria, Helminthostachys zeylanica and Ophioglossum vulgatum are sometimes eaten as vegetables in Asia. The young unexpanded shoots of Botrychium australe was previously consumed by Maoris in New Zealand. Etymology: Ophioglossum is derived from the Greek, όφις (ofis), a snake and γλώσσα (glossa), tongue, in reference to the tongueshaped fertile parts of the leaf. Plants of the World

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PSILOTALES

FERNS

PSILOTALES In the past, early land plants with similar bifurcating branches were included here, but these (e.g. Rhynia, Psilophyton) are now known not to be related, a case of extreme convergence. Psilotales were formerly placed among lycopods as they have leaves that are simple, resembling microphylls. Nevertheless they are an ancient plant lineage based on a molecular clock, but the order has scanty fossil evidence and estimates are therefore fairly uncertain. They include the single extant family Psilotaceae and are distantly related to Ophioglossales, with which they share some morphological characters. Distribution: The family is pantropical to warm-temperate, but absent from dry areas. They extend northwards into the southern USA, southwestern Europe and Japan and southwards to New Zealand (Psilotum nudum). The greatest diversity of Tmesipteris is in Australasia and Pacific islands, where it occurs almost exclusively on treefern trunks. Psilotum can be terrestrial, epiphytic or epilithic.

6. PSILOTACEAE Whisk-fern family

These are epiphytic or terrestrial plants without roots. Their rootless rhizomes are fleshy and forked. The stems are usually also forked in twos (dichotomously branched), but are poorly or unbranched in some taxa. Their scale-like leaves have (or lack) a single central vein and therefore resemble microphylls. The sporangia are large and lack an annulus. Sporangia are often fused with two or three together to form a synangium, which appears to be borne on the adaxial side of a forked leaf. The spores are reniform and monolete, and there are > 1,000 per sporangium. The gametophytes are tuberous, subterranean, achlorophyllous and mycoheterotrophic.

Psilotum nudum, Australian National Botanic Gardens, Canberra, Australia [6]

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Phylogeny and evolution: Because of the reduced leaves, the simple dichotomous branching and the protostelic rhizomes that lack roots, Psilotum was often considered to be a “living fossil”, a descendant of the earliest land plants such as the fossil Rhynia (or other Psilotophyta). Because of their microphyll-like leaves they have in the past been associated with lycopods. Due to molecular studies, it is now known with certainty that this resemblance is superficial and that extant Psilotaceae and fossil Psilotophytaceae are not related. Psilotaceae belong to the fern lineage, where they are generally placed as the sister to Ophioglossaceae. The resemblance with that family is maybe not obvious at first, but the similarity in rhizome

Psilotum nudum, Réunion [6]

structure, absence of roots and mycoheterotrophic gametophytes with multiflagellate sperm cells (antherozoids) make Psilotaceae/Ophioglossaceae a good morphological entity. There is little known of these groups in the fossil record. Devonian fossils previously attributed to this lineage are not related to this family. Some species have large genomes, among the largest of any organism (but see also Melanthiaceae). Genera and species: Psilotaceae consist of two genera with a dozen species: Psilotum (2) and Tmesipteris (c. 10). Uses: Spores of Psilotum nudum were collected and used by Hawaiians and Polynesians as talcum powder applied under the loincloth to prevent chafing. It was a sacred plant in Japan during the 17th century, resulting in a great horticultural interest and the development of over 100 cultivars, many of which are still grown in bonsai collections. Etymology: Psilotum is derived from Greek ψηλός (psilos), bare, in reference to the naked stems of Psilotum nudum.

Tmesipteris ovata, New South Wales, Australia [6]

Tmesipteris vieillardii, New Caledonia [6]

MARATTIALES

FERNS

MARATTIALES This is an ancient lineage of ferns and includes the bulk of early fern fossils, such as Psaronius and Asterotheca (fossil Asterothecaceae), which were tall tree-like ferns. Modern Marattiales are easily recognised by their fleshy leaves that arise from succulent or papery outgrowths along the rhizome and their sporangia that are fused into complex synangia.

Angiopteris madagascariensis, Seychelles, with Michael Fay for scale [7]

7. MARATTIACEAE King-fern family

These sometimes enormous, mostly terrestrial ferns have mucilage canals in all their parts. They have large fleshy roots and fleshy rhizomes that are short and upright or creeping and elongate, sometimes globose. The rhizomes bear f leshy or papery, starchy stipule-like outgrowths at the base of leaf insertions that often bear adventitious buds from which new plants can grow. Leaves are usually large, fleshy or leathery and one- to four-times pinnate or palmate, rarely simple. The petioles and rachises have swollen nodes (pulvini) and stems, blades and rhizomes are covered with peltate scales. Lenticels are scattered along petioles and rachises. Their sporangia are fused but nearly free in some species, usually united in round or elongate

Angiopteris angustifolia, Hortus botanicus, Leiden, the Netherlands [7]

Angiopteris evecta, rhizome, Tahiti [7]

synangia (fused sporangia) that lack an annulus, but open by slits or pores, enclosing 1,000–7,000 spores. The spores are usually bilateral or ellipsoid and monolete. The gametophytes are superficial, heart-shaped, large and green. Distribution: Marattiaceae are distributed throughout tropical and subtropical regions of the world. Danaea, Eupodium and Marattia are found in tropical America from Mexico to Argentina, with Marattia also extending to Hawaii. Christensenia is found from the Himalayas through Southeast Asia and Malesia to the Solomon Islands. Angiopteris occurs from Madagascar, the Seychelles and southern India throughout tropical Asia to northeastern Australia and numerous Pacific islands as far east as Pitcairn. Ptisana (formerly Marattia of the Old World) occurs in tropical Africa, Madagascar, the Mascarenes, tropical Asia and Australasia, south to New Zealand and north to Japan. Phylogeny and evolution: Fossils of the distinctive stem anatomy and synangia place the origin of marattioid ferns in the Upper Carboniferous, making them one of the oldest extant lineages of land plants. Modern genera

are known with certainty from the Triassic and Jurassic and were widespread on continents of the Northern Hemisphere. The centre of diversity now lies in the northern Andes and southern China. The lineage is isolated and has no close extant relatives. The family is divided in two subfamilies: Danaeoideae, including only Danaea, and Marattioideae, in which the other genera are placed. Genera and species: Marattiaceae include six genera with c. 135 species: Angiopteris (c. 30), Christensenia (1), Danaea (c. 55), Eupodium (3), Marattia (c. 10) and Ptisana (c. 35). Uses: Starch from the rhizomes of some Angiopteris species is sometimes eaten in southern Asia, and Angiopteris rhizomes are used to flavour rice in Polynesia. In India, Angiopteris stems are used to make ruchshi, an intoxicating drink. The rhizome of Marattia douglasii was steamed and eaten by Hawaiians, and similarly Ptisana salicina was cultivated for consumption by Maoris in New Zealand. Etymology: Marattia was named for Italian botanist Giovanni Francesco Maratti (died 1777).

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MARATTIALES

FERNS

Danaea geniculata, Paraná, Brazil, detail of sori [7]

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Ptisana oreades, detail of sori, Australian National Botanic Gardens, Canberra [7]

Danaea polymorpha, Guadeloupe [7]

Eupodium kaulfusii, detail of sori, São Paulo, Brazil [7]

Eupodium kaulfusii, São Paulo, Brazil [7]

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OSMUNDALES

FERNS

LEPTOSPORANGIATE FERNS Families 8 to 33 are called leptosporangiate ferns. This is the crown-group of ferns. In all species the sporangia develop from a single epidermis cell. All other fern lineages (and seed plants and lycopods) are eusporangiate, meaning that the sporangia are developed from a group of cells. In leptosporangiate ferns, the sori are often covered in a scale-like structure called an indusium, a character frequently used in identification. The vast majority of extant ferns are leptosporangiate.

OSMUNDALES A widespread order with a fossil history going back to the Permian, they now comprise the single family Osmundaceae with sporangia that are intermediate between those of the Ophioglossales and the typical leptosporangiate ferns.

8. OSMUNDACEAE Royal-fern family

Todea barbara, National Botanic Gardens of Ireland, Glasnevin [8]

These terrestrial ferns have trunk-like rhizomes that are clothed in roots and leaf bases. Their petioles have stipules at the base. Leaves are monomorphic, fully dimorphic or with fertile portions dissimilar to the sterile parts on the same leaf, and are once to twice pinnate and hairy when young. The venation is free, usually forked. Their sporangia are free and not assembled in sori. They have a short stout stalk and a large capsule that can contain >100 to >1,000 spores. The capsule has an annulus that consists of a group of cells thickened on one side that makes the

Osmundastrum cinnamomeum, Hortus botanicus, Leiden, the Netherlands [8]

capsule open by an apical slit. Spores are green, subglobose, trilete, smooth and shortlived. The germination of the spores is bipolar. Gametophytes are large, superficial, green, heart- or ribbon-shaped and long-lived, and have a strong midrib, apical rhizoids and basal meristems. Distribution: Osmundaceae are cosmopolitan, except for hot, dry areas. Leptopteris is confined to Australasia, and Todea is found in three widely disjunct areas: South Africa, New Guinea and eastern Australia.

Leptopteris fraseri, New South Wales, Australia [8]

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OSMUNDALES

FERNS

Phylogeny and evolution: The stelar (root) structure of Osmundaceae is unique among ferns and allows assignment of fossils to this family, with certainty dating back to the Permian. No close relatives are known, which is not unusual for a lineage of such antiquity. It has been suggested that Osmundaceae are an intermediate stage towards leptosporangiate ferns. Molecular phylogenetic analyses have placed them as sister to all other leptosporangiate ferns, so it is possible that they represent a group of ferns that emerged around the time of the development of the leptosporangium. There are affinities to other early diverging fern lineages, including the presence of stipules, free venation, large spore output and sporangia that are sometimes formed by more than a single cell.

The cinnamon fern, often included in Osmunda, is sister to the rest of the extant taxa. To keep Todea and Leptopteris separate from Osmunda and to make the last monophyletic, the cinnamon fern has been segregated as Osmundastrum cinnamomeum. Genera and species: Osmundaceae consist of four genera with 23 species: Leptopteris (10), Osmundastrum (1), Osmunda (c. 10) and Todea (2). Uses: Young shoots (fiddleheads) and winter buds of cinnamon fern, Osmundastrum cinnamomeum, can be cooked and eaten. ‘Fern butter’ is the edible pith of the rhizome of the interrupted fern, Osmunda claytoniana. An edible starch is extracted from ‘zendai’, Osmunda japonica, in Japan.

The fibrous trunks of Osmunda regalis are sometimes used for orchid cultivation as a substitute for tree fern fibre, resulting in a threat to local populations and legal protection of the species in some countries. Etymology: The name Osmunda is of uncertain origin and was in use long before Linnaeus applied it. It may have originated from Old French osmonde, a kind of fern. It alternatively could have been derived from Old Saxon ås mund, god’s strength, referring maybe to the stately and rapid growth of the fern in spring. Alternatively it is possible that it was named for Saint Osmund, Bishop of Salisbury (died 1099), or another historical person with a similar name. The origin of the name remains shrouded in the mists of time.

HYMENOPHYLLALES This unusual group of ferns has thin leaves that are one cell thick. They diverged c. 243 million years ago. The order includes the single family Hymenophyllaceae with two genera that diverged during the Jurassic. They largely occur in the tropics where they are epiphytic.

Trichomanes elegans, São Paulo, Brazil [9]

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Trichomanes cuspidatum, Réunion [9]

Trichomanes endlicherianum, Tahiti [9]

HYMENOPHYLLALES

FERNS

Hymenophyllum sibthorpioides, Réunion [9]

9. HYMENOPHYLLACEAE Filmy-fern family

These are moss-like terrestrial or epiphytic ferns. They have slender, wiry and creeping or climbing, or erect and stout rhizomes, with or without roots. Leaves are simple to variously pinnately compound, or variously regularly or irregularly divided and can be small (< 2 mm) to intermediate in size (c. 50 cm) when mature. The blades are usually one or only a few cell layers thick and transparent and lack stomata. Venation is free or somewhat anastomosing, simple, forked or flabellately arranged. Sori terminate veinlets and are formed at the tips of ultimate segments or on lobe margins. The involucres

Hymenophyllum elegans, Paraná, Brazil [9]

are cup-shaped, usually two-lipped or bilobed, with a central receptacle that is often elongate and exserted, along which the sporangia are formed. The sporangia are short-stalked with a well-defined annulus. Spores are spheroidal, variously ornamented, green and short-lived, usually germinating inside the sporangium. Gametophytes are filamentous or ribbon-shaped and long-lived, reproducing vegetatively by proliferating buds in some species. Distribution: These moisture-dependent ferns occur predominantly in mossy forests in tropical montane regions and in temperate rain forests. They have a nearly worldwide distribution but are absent in areas with dry cold winters or hot dry summers. They can be found north to British Columbia and Norway and south to Tierra del Fuego and New Zealand. Vegetatively reproducing gametophytes have been reported from Germany, Luxemburg and eastern North America, in areas far colder than those where sporophytes of the same species occur. These gametophyte

populations have been presumed to be relics of warmer eras. Phylogeny and evolution: The fossil record of Hymenophyllaceae is sparse, but it has been suggested that the filmy leaf morphology is a derived character that evolved with the evolution of seed-plant-dominated tropical rainforests. They are an isolated lineage placed near Gleicheniales. Within Hymenophyllaceae, the two clades found correspond to the genera accepted here. Genera and species: Hymenophyllaceae consist of two genera with c. 650 species: Hymenophyllum (c. 350) and Trichomanes (c. 275). Trichomanes is sometimes divided into several smaller genera, but the genus is monophyletic when maintained in the broad sense, so the division into several genera serves no particular purpose. Etymology: Hymenophyllum is derived from the Greek υμένας (ymenas), a membrane or skin and φύλλων ( fyllon), a leaf, referring to the thin leaf texture of most species. Plants of the World

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GLEICHENIALES

FERNS

GLEICHENIALES This is a relatively isolated assembly of three similar families with a long fossil record. They are good indicators for present and past distributions of tropical vegetation.

Gleichenia microphylla, sori, New Caledonia [10]

Gleichenella pectinata, Guadeloupe [10]

Dicranopteris flexuosa, sori, Brazil [10]

Stromatopteris moniliformis, New Caledonia [10]

10. GLEICHENIACEAE

in regular clusters. Sporangia are short-stalked with a large capsule and numerous spores. The capsule has an oblique, nearly complete annulus. Spores are achlorophyllous. Gametophytes are elongate to subcordate, green in all genera except Stromatopteris, which has tuberous, achlorophyllous, mycoheterotrophic gametophytes.

Antarctica. With Dipteridaceae and Matoniaceae, the family forms an isolated lineage. Stromatopteris, endemic to New Caledonia, is sister to all other Gleicheniaceae and sometimes placed in its own family or subfamily.

Forking-fern family

These terrestrial ferns have creeping rhizomes that bear scales and/or hairs. Leaves are once-pinnate, the midribs often forking dichotomously and with axillary buds from which leaves can continue to grow indeterminately. Petioles have a single vascular bundle that is c-shaped in cross-section. Pinna segments are adnate and usually firm in texture, and venation is free. Sori are round, formed on the dorsal side of the leaf and exindusiate, with few sporangia, usually arranged

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Genera and species: Gleicheniaceae consist of six genera with c. 165 species: Dicranopteris (c. 20), Diplopterygium (c. 25), Gleichenella (1), Gleichenia (18), Sticherus (c. 100) and Stromatopteris (1).

Distribution: Gleicheniaceae are a pantropical family with their greatest diversity in the Neotropics and tropical Asia, extending occasionally into the subtropics. Many species are found in pioneer habitats or disturbed areas, roadsides, landslides etc. Gleicheniaceae prefer nutrient-poor soils and are often found in full sun, but then usually in montane areas where ample water is available.

Uses: Petioles of Dicranopteris linearis were woven into hats and cigar boxes on Java. In Malaysia they were used as pens and woven into mats, fishing traps and chair seats; they have even been used to make the walls of huts.

Phylogeny and evolution: This ancient family dates back at least to the early Mesozoic and possibly the Carboniferous. Fossils are known from all continents, including

Etymology: Gleichenia was named for German naturalist and plant physiologist Baron Wilhelm Friedrich von Gleichen (1717–1783).

GLEICHENIALES

FERNS

11. DIPTERIDACEAE Double-fern family

These are terrestrial or epilithic ferns. Their rhizomes are long-creeping and clothed with scale-like dark bristles or hairs. Petioles are placed centrally or basally on the blade and have one to several vascular bundles that are organised in a U-shape in cross-section. Leaves are monomorphic or strongly dimorphic. The blade is entire or divided deeply into two more or less equal parts, usually

glabrous (at least when mature), the margins deeply lobed, cleft or entire. Larger veins are palmately arranged at the base and forking several times, the smaller veins reticulate, and areoles with many included veinlets. The sori are small, round and exindusiate and cover the lower surface of the fertile leaves or are irregularly seriate. Sporangia are stalked, the stalks are four cells wide and the annulus is longitudinal, somewhat oblique. Sporangia are surrounded by hairs with a thickened tip. Spores are monolete or trilete, ellipsoid or tetrahedral, achlorophyllous, smooth or rugulose. Gametophytes are naked and heart-shaped, with a thick midrib and usually gametangia on both surfaces.

Phylogeny and evolution: This family consists of two isolated genera that, with Matoniaceae and Gleicheniaceae, form an isolated ancient lineage (Gleicheniales). In the past the family was associated with Polypodium, but this was only based on the shared lack of indusia. The family dates back to the Upper Triassic, with some fossils probably assignable to the extant genus Dipteris; the genus is thus of considerable antiquity and can be considered a “living fossil”.

Distribution: Dipteridaceae occur in tropical Asia and Australasia, from Assam to southern China and Okinawa, south to Queensland, New Caledonia and Fiji.

Etymology: Dipteris is derived from the Greek δυο (dyo), two, and πτέρης (pteris), fern, in reference to the biparted leaf.

Genera and species: Dipteridaceae consist of nine species in two genera: Cheiropleuria (1) and Dipteris (8).

Cheiropleuria integrifolia, Taiwan (RK) [11]

Dipteris conjugata, sori, New Caledonia [11]

Dipteris conjugata, New Caledonia [11]

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GLEICHENIALES

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Matonia pectinata, Malaysia (HW) [12]

12. MATONIACEAE Umbrella-fern family

These terrestrial or epilithic ferns have longcreeping, densely hairy rhizomes. Leaves are monomorphic, and petioles are dark and glossy, with a single simple vascular bundle. Blades have either pinnatifid pinnae that are pedately arranged or pinnae that are once forked and have axillary bulbils that cause indeterminate growth. Venation is free or

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somewhat reticulate. Sori are formed on the lower side of the leaves and are round and indusiate. The indusium is centrally attached and umbrella-shaped. Sporangia are few, large and placed in a circle around the indusium. The sporangium stalk is short, the capsule is subglobose and the annulus is oblique and incomplete (not surrounding the capsule entirely). Spores are trilete. Gametophytes are superficial, heart-shaped and naked, with antheridia on both sides and archegonia only on the lower side. Distribution: All extant members are restricted to the Malesian region and exhibit a relictual distribution. Matonia is restricted to the Malay Peninsula and Borneo, and Phanerosorus occurs only in the Moluccas. Usually they are found locally in tropical mountain areas on soils with low mineral content.

Phylogeny and evolution: Matoniaceae are of great antiquity, dating back to the Lower Mesozoic, and extant Matonia pectinata has been even considered to have been present since the Cretaceous. The family had a much wider distribution in the past based on their much more extensive fossil distribution. They have no close relatives, but the family is placed with Dipteridaceae and Gleicheniaceae in Gleicheniales based on DNA studies. Genera and species: Matoniaceae consist of two genera with four species: Matonia (2) and Phanerosorus (2). Etymology: Matonia is named for William George Maton M.D. (1774–1835), a fellow of the Royal College of Physicians, who had a keen interest in botany and was a friend of Robert Brown, who named this genus for him.

SCHIZAEALES

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SCHIZAEALES This order previously consisted of three families that are now united under the single family Schizaeaceae. They are remarkable in having sporangia that are borne on a marginal, pinnately arranged sporangiophore with each sporangium covered with a flap of leaf tissue.

These are terrestrial or epilithic ferns with creeping or suberect, articulately hairy rhizomes. Petioles have a single vascular bundle and are erect, sometimes twining. Blades are once or twice (to many-fold in

Lygodium) pinnate-pinnatifid, pinnatepalmatifid, or simple and pinnately lobed or dichotomously branched, sometimes absent or reduced to wings along a midrib. Veins are free, (dichotomously or flabellately) forking or rarely casually anastomosing; areoles, when present, are without included veinlets. Sporangia usually borne on sporangiophores or in two rows along a vein in a leaf tooth, usually pinnately or digitately arranged, usually each lateral veinlet bearing a sorus with a single sporangium and a flap of leaf tissue covering the sorus. Sporangia are asymmetrical, ovoid or pear-shaped, the annulus just below or at the apex. Spores are trilete or monolete, ellipsoidal, (sub-)globose or angular and are variously ornate. Gametophytes are

Actinostachys confusa, Seychelles [13]

Schizaea elegans, Peru [13]

13. SCHIZAEACEAE Fan-fern family

heart-shaped or filamentous, (partly) chlorophyllous and often with rhizoid cells, or (in Actinostachys) mycoheterotrophic, fleshy and tuber-like. Distribution: The diverse Schizaeaceae are mainly distributed in the tropics but extend in the Americas south to temperate South America and north to Newfoundland. They also occur in southern Africa, Madagascar and the Mascarenes, throughout tropical Asia, north to southern Japan and south to New Zealand. They usually grow on in mineralpoor soils often in open habitats, sometimes on decaying wood or on ultrabasic soils, but unlike other Schizaeaceae, Lygodium is usually found on fertile soils. Lygodium japonicum, Copenhagen Botanical Garden, Denmark [13]

Schizaea dichotoma (WA) [13]

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SCHIZAEALES

Lygodium volubile, sori, Brazil [13]

Phylogeny and evolution: Schizaeaceae are the single extant family in the order Schizaeales, which form the sister lineage to core leptoporangiate ferns (including Polypodiales, Cyatheales and Salviniales), a lineage of considerable age originating in the Jurassic. The three lineages, represented here at the subfamily level, diverged early in their evolutionary history and have therefore been separated as independent families in some classifications, even though they share major characters and were in historical classifications usually placed in a single family (as done here). Lygodioideae are sister to the rest and stand apart morphologically from the other two subfamilies in having oblique sporangia and climbing habits. The two genera in Schizaeoideae differ

FERNS

Anemia sp., near Ubatuba, Brazil [13]

Anemia vestita, Kenya [13]

mainly in gametophyte and sporangium structure, Actinostachys having mycoheterotrophic gametophytes and pseudodigitate sporangiophores. The oldest fossil assigned to Schizaeoideae dates from the Cretaceous. In Anemioideae two lineages were recognised on the basis of morphological differences, but the African genus Mohria was found to be embedded in Anemia s.s., Mohria being clearly derived from Anemia. The resemblance of Mohria to cheilanthoid ferns rests mostly upon a superficial similarity, convergence probably due to similarities in habitat preference. Genera and species: Schizaeaceae, with three subfamilies, comprise four genera and c. 190 species: Lygodioideae – Lygodium (c. 35); Anemioideae – Anemia (c. 115);

Schizaeoideae – Actinostachys (16) and Schizaea (21). Uses: Anemia caffrorum and related species give off an aroma of balsam when bruised. The essential oils are antibacterial, can be used to treat burns and are used in South Africa to prevent children’s nightmares. The twining rachises of Lygodium salicifolium are used in Thailand as a material for weaving. Occasionally used as an ornamental in the tropics, some species especially Lygodium japonicum and L. microphyllum have locally become serious weeds outside their native range. Etymology: Schizaea is derived from Greek σκίζει (skizei), to split.

SALVINIALES Families 14 and 15 are placed in the order Salviniales, also sometimes referred to as the heterosporous ferns. They are distinguished from other ferns in having two sizes of spores, megaspores and microspores, that are formed in a sporocarp, a hard structure in which the sporangia are formed; it is hypothesised to be an adaptation to their aquatic habitat.

14. MARSILEACEAE Pillwort family

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These are small terrestrial ferns of swampy, periodically wet or permanently flooded places. Their rhizomes are slender, creeping and much branched. They have air canals and bear hairs, but become glabrous later. Leaves are formed in two rows along the rhizome and are hairy when young. Petioles have a single V-shaped vascular bundle. Blades are lacking (Pilularia) or the lamina has two (Regnellidium) or four (Marsilea) segments, these pinnately

arranged, but the rachis nearly lacking, making the lamina appear palmate and clover-like. Veins are flabellately arranged, forking, and usually anastomosing on both ends, without included veinlets. Sori are formed inside firm, globose or ellipsoidal stalked sporocarps (a modified globose indusium), inserted on the petiole or on the rhizome near the petiole base. Sporocarps are long-lived and dehisce into two valves upon water uptake by the gelatinous interior

SALVINIALES

FERNS

Marsilea drummondii, South Australia [14]

Marsilea mutica, private pond, Kingston upon Thames, Surrey, UK[14]

that contains two or more sori carried out by the protruding gelatinous mass, containing the mega- and microsporangia that lack an annulus. Spores are trilete and of two different sizes: a single large one is formed in each megasporangium, and many small ones are formed in each microsporangium; the microspores are free. Spores germinate rapidly, the microspores producing spermatozoids within a day of germination, fertilisation occurring soon after germination of spores. Gametophytes are greatly reduced, consisting of a few cells only; they remain attached to the spores and are often developed and fertilised within the spore.

relationships of heterosporous water ferns (Marsileaceae and Salviniaceae) have long been disputed. They were previously placed in Hydropteridales, with some fossil taxa, usually among ferns, but sometimes among ‘fern allies’. Molecular phylogenetic studies have placed the heterosporous ferns as sister to the rest of the families in core leptosporangiates, heterospory (among vascular plants otherwise only known in Selaginellaceae and Isoëtaceae) being a derived character and probably an adaptation to their aquatic habitat. Macrofossils of Marsileaceae are known from the Upper Jurassic and later.

Distribution: A family of nearly worldwide distribution, but absent from cold or extremely dry regions and most oceanic islands. Some Australian species of Marsilea are drought resistant and can be found in desert environments, but usually in periodically wet areas. Sporocarps are usually produced in areas where the water level fluctuates and the plant dries out. Plants thrive in permanently flooded areas, but sporocarps are not always produced when the plant is submerged.

Genera and species: Marsileaceae consist of three genera with c. 65 species: Marsilea (c. 60), Pilularia (5) and Regnellidium (1).

Phylogeny and evolution: Placement and

Uses: Sporocarps of nardoo (Marsilea drummondii) were ground into flour and baked into bread by aboriginal Australians. The spores are, however, indigestible, provide no nutrition and are astringent. A diet of just nardoo bread results in death from starvation with a full stomach. Etymology: Marsilea was named for Italian mycologist Count Luigi Marsigli (1656–1730).

Pilularia globulifera illustration from O. W. Thomé (1885) Flora von Deutschland, Österreich und der Schweiz, Gera, Germany. [14]

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SALVINIALES

15. SALVINIACEAE Water-fern family

These are free-floating aquatic plants. Stems are horizontal, green, branched and floating. The two genera differ substantially (and were previously placed in separate families) and therefore are described here separately. Azolla has simple, unbranched roots on the lower side of the stems. Leaves are sessile, alternate, and formed in two rows on the upper side of the stem, each leaf bilobed, with one lobe upright, chlorophyllous, with a cavity harbouring colonies of the blue-green alga Anabaena, the other lobe floating, mostly without chlorophyll. Sori are formed inside sporocarps (modified indusia), the sporocarps formed on the first leaf of a side branch, replacing the lower lobe and the upper lobe covering the sporocarp. Sporocarps contain either micro- or megasporangia. The megasporocarps (containing a single sporangium) are smaller than the microsporocarps (containing numerous sporangia). Sporangia lack an annulus and spores are filamentous. Microspores

FERNS

are trilete and embedded in hardened mucilage. Megaspores have hair-like filaments and floats. Gametophytes of microspores reduced with a single antheridium. Gametophytes of the megaspores develop within the spore. Salvinia lacks roots. Leaves are formed in whorls of three along the length of the stems; the two upper leaves are floating and green, the lower one is submerged, much branched and root-like. The floating leaves are sessile or slightly stalked, ovate to cordate, with specialised water-repelling hairs on the upper surface. The veins are pinnate and lateral veins anastomose, the areoles without included veinlets. The submerged leaves are petiolate, finely divided and hairy, and the branches bear sporocarps. Sori develop inside sporocarps (globose indusia), one or a few of these sporocarps producing megasporangia and the other sporocarps producing microsporangia. Sporangia lack an annulus. Spores are spherical, plain to rugose. Microspores are enclosed in a hardened mass. Megaspores are trilete. Gametophytes of microspores are reduced, formed of only a few cells, and bear reduced antheridia. Gametophytes of megaspores are free-floating, heart-shaped and connected to the megasporangium. Distribution: The genera occur in southern North America, South America, Europe and western Asia and throughout the Palaeotropics, except Australasia. The plants are free-floating on fresh water surfaces such as

Salvinia natans, Helsinki Botanical Garden, Finland [15]

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Salvinia molesta, submerged root-like leaf, Tahiti [15]

ponds, rice paddies, river margins, ditches, drainage and irrigation canals, in swamps, bogs etc. Phylogeny and evolution: The two genera (previously often placed in separate families) are closely related and the pair are sister to Marsileaceae. Together these families form a lineage of heterosporous ferns, which is sister to the rest of the core leptosporangiates. Their fossil history dates back to the Cretaceous. The genera are morphologically divergent and for that reason each has been placed in its own subfamily. Genera and species: Salviniaceae consist of two genera with c. 20 species in two subfamilies: Azolloideae – Azolla (7); Salvinioideae – Salvinia (12). Uses: Because of its nitrogen-fixing cyanobacteria, Azolla pinnata is released in rice paddies in East Asia to fertilise the soil. A paddy overgrown with Azolla is drained, and the plants are ploughed into the soil before young rice is planted. Azolla filiculoides and Salvinia molesta are invasive species outside their native range and, when cultivated, release of these plants into nature needs to be prevented. Both genera are employed as biofilters to remove toxic compounds from waste water. Etymology: Salvinia was named for Italian botanist A.W. Salvini (1633–1729).

Azolla filiculoides, France [15]

CYATHEALES

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CYATHEALES The order Cyatheales was previously divided into eight families that vary greatly in general appearance, but these all share a helicogyroid annulus that bypasses the stalk of the sporangium. There are members that look like filmy ferns (some Cyathea), some that are tree-like (many Cyatheoideae and Dicksonioideae) and some (Plagiogyria) that more closely resemble hard ferns (Blechnum, Aspleniaceae) with creeping rhizomes and pinnate leaves, but all are now united in a single family: Cyatheaceae. Because some of the former family names are still commonly in use and are equivalent to current subfamilies, we opted to treat all subfamilies separately here.

16. CYATHEACEAE Tree-fern family

Cyatheaceae are a variable group of terrestrial or epilithic ferns, including some of the smallest and some of the largest ferns known. Rhizomes are creeping or erect, often trunklike, and covered in scales or hairs. Leaves are minute to large, 5–500 cm. Petioles have a single vascular bundle that is usually U- or V-shaped in cross section. Blades are rarely simple, usually once or up to five times pinnate. Veins are usually free or somewhat anastomosing near the margin, rarely fully anastomosing, the free veins often not reaching the margin. Sori usually develop on veins, dorsally or marginally, round, the receptacle raised or cup-shaped, with the indusia usually surrounding the receptacle, or the indusia fixed on one side of the sorus and scale-like; indusia are sometimes completely absent. Sporangia are stalked with a nearly vertical, oblique annulus that bypasses the sporangium stalk. Spores are trilete and globose, sometimes angular and variously ornamented. Gametophytes are green and heart-shaped and often scaly. Distribution: This is a pantropical family, extending into the subtropics and wet temperate regions of the Southern Hemisphere. Some temperate, mainly Australian

species are naturalised in suitable habitats in the Northern Hemisphere, including coastal British Columbia, Hawaii, the Macaronesian Islands and western Ireland and Britain. Phylogeny and evolution: This family is the single extant representative of the tree fern lineage, order Cyatheales. These include treelike species with tall trunks and large leaves, but also include minute species with trunks and leaves only a few centimetres long (former genus Hymenophyllopsis, now included in Cyathea). The family also includes some species of intermediate size with creeping rhizomes similar to members of Polypodiales, and species of this group have often been confused with members of Dennstaedtiaceae or Davallia (Polypodiaceae). The group is characterised by oblique annuli on the sporangium capsules. Molecular divergence of tree ferns is slow, which could be due to their long generation times, as has been hypothesised for palms. The current diversity of tree ferns is thus possibly relatively recent and the subfamilies discussed below probably date from the Jurassic with some lineages being of a relictual nature. Others, especially the species-rich Cyatheoideae, have diversified in more recent times, with many species in montane rainforests of the tropics. Genera and species: Cyatheaceae include eight subfamilies with 12 genera totalling about 700 species: Alsophila (c. 260), Calochlaena (6), Cibotium (11), Culcita (2), Cyathea (c. 275), Dicksonia (22), Lophosoria (1), Loxsoma (2), Metaxya (2, probably more), Plagiogyria (c. 15), Sphaeropteris (c. 100) and Thyrsopteris (1). The subfamilies are described separately below because they are sometimes treated at the family level.

Uses: The growing tips of tree ferns are rich in starch and sometimes roasted and eaten, although this usage is not sustainable because it kills these slow-growing plants. This usage has, however, been reported from native peoples in Madagascar, Australia, New Zealand and New Caledonia, and Cibotium billardieri was eaten in Hawaii. The young leaves (fiddle-heads) are also eaten in some cultures and supposedly taste like bitter celery or palm heart. Tree fern trunks are widely used in construction and carving, and the peaty root-masses make good flower pots or growing medium for orchids; due to this last utilisation, international trade in Cyathea and Dicksonia is currently regulated by CITES. The fluffy scales of Cibotium, Culcita and other tree ferns were harvested on a large scale as stuffing for pillows and were frequently used in the seat cushions of early cars. Many species are popular ornamental plants in the tropics and subtropics. Etymology: Cyathea is derived from the Greek κυάθιος (kyathios), a cup or beaker, in reference to the shape of the indusium in some species.

16a. CYATHEACEAE SUBFAMILY THYRSOPTERIDOIDEAE Robinson-Crusoe ferns These terrestrial ferns have erect or curved upright rhizomes that are covered with stiff hairs. Leaves are large and long-petiolate, the petioles hairy at the base, and have a single Ω-shaped vascular bundle in cross-section. Blades are three- to five-pinnate, firm and glabrous when mature. Veins are free and do not reach the leaf margins. Fertile portions are

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CYATHEALES

FERNS

Thyrsopteris elegans, Glasgow Botanic Gardens, Scotland, UK [16a]

found on the basal part of the blade, where the pinnae are much dissected and almost without blade tissue. The sori are formed terminally on stalk-like veins, and the indusia are fused to form a cup in which the sporangia are placed on a column-shaped receptacle. Sporangia have a stalk with a complete annulus. Spores are globose-tetrahedral, prominently angular, verrucate. Gametophytes are green and superficial. Distribution: Thyrsopteridoideae are endemic to the Juan Fernández Islands, an archipelago off the coast of Chile in the southeastern Pacific. Phylogeny and evolution: Fossils of Thyrsopteridoideae are widespread, and the current restricted distribution is therefore certainly relictual. The lineage is of greater age than the age of the volcanic Juan Fernández Islands, and the genus was once widespread across South America, but died out there and only remained on these isolated islands. Genera: Thyrsopteridoideae include only a single, relictual, extant species: Thyrsopteris elegans. Etymology: Thyrsopteris is derived from the Greek θύρσος (thyrsos), a stalk or stem of a

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Loxsoma cunninghamii, collected by Colenso in New Zealand (Herbarium Kew, ex coll. Hooker) [16b]

plant, and πτέρης (pteris), a fern, in reference to the stalked sori.

northern Andes. They occur in open places in moist lowland forests.

16b. CYATHEACEAE SUBFAMILY LOXSOMATOIDEAE Cross ferns

Phylogeny and evolution: The disjunct distribution is an indication that the distribution of this subfamily may be relictual, although long-distance dispersal cannot be excluded. The widespread fossil genus Stachypteris from the Jurassic has been placed in this subfamily with certainty. Previously the subfamily was associated with Hymenophyllaceae and Dipteridaceae, based on superficial similarity (the urn-shaped sori and the bristle-like rhizome hairs, respectively). The gametophytes with scale-like hairs are similar to some Cyatheoideae, and an association, albeit distant, with that subfamily has been demonstrated in molecular studies.

These terrestrial ferns have long-creeping, branched rhizomes with stiff bristles. Leaves are placed distantly along the rhizome and are two- or three-pinnate-pinnatifid. Petioles are bristly-hairy at base, naked elsewhere and long, with a single, sickle-shaped vascular bundle in cross-section. Blades are firm with deltate-lanceolate pinnules. Veins are free, forked or somewhat pinnate. Sori terminate a vein and have an urn-shaped indusium; they are marginal and attached to the lower side of the blade, with a columnar central receptacle. Sporangia are formed among multicellular hairs and are stalked, pear-shaped, with a subvertical, somewhat oblique annulus. Spores are trilete, tetrahedral-globose, tuberculose or rugose, pitted and ridged. Gametophytes are a superficial, green, elongate, heart-shaped prothallus with multicellular scale-like hairs, especially above. Distribution: Loxsomatoideae are disjunctly distributed in northern New Zealand and in the Neotropics from Costa Rica to the

Genera and species: Loxsomatoideae consist of two genera with two species: Loxsoma cunninghamii in New Zealand and Loxsomopsis pearcei in the Neotropics. The species are closely related, morphologically similar and could be treated in a single genus. Etymology: Loxsoma is derived from the Greek λοξός (loxos), crosswise or oblique, and σώματος (somatos), body. The name was originally published erroneously as ‘Loxoma’, and later ‘corrected’ by the authors as Loxsoma.

CYATHEALES

FERNS

16c. CYATHEACEAE SUBFAMILY CULCITOIDEAE Cushion ferns

Caribbean, and in the Azores, Madeira, Tenerife and northwestern Spain. They are usually found in wet open forests.

These large terrestrial ferns have thick creeping or ascending rhizomes. The rhizomes are covered in persistent petiole bases and long hairs. Leaves are closely placed at the apices of the thick trunks. Petioles are covered with yellowish or brownish long hairs and have a single U-shaped vascular bundle in cross-section. Blades are four- to five-pinnatepinnatifid and are slightly hairy to glabrous. Veins are free, usually forking and reaching the leaf margins. Fertile portions of the leaf are similar to sterile portions or the lamina of fertile portions are somewhat reduced in size. Sori are formed marginally, terminating a vein, and are surrounded by two indusia on either side and have a central transversely elongate receptacle on which the sporangia are formed. The outer indusium is similar to the leaf blade and is fused with the inner indusium. Spores are globose-tetrahedral, depressed between the angles and variously ornate. Gametophytes are superficial, green and heart-shaped.

Phylogeny and evolution: Culcitoideae are sister to Plagiogyrioideae and not closely related to Calochlaena, a genus previously included in Culcita, but now placed in Dicksonioideae. The disjunct distribution of Culcitoideae may be relictual, but is more probably due to long-distance dispersal along the prevailing winds following the Gulf Stream.

Distribution: Culcitoideae occur disjunctly in tropical America from southern Mexico to Bolivia, Venezuela and throughout the

These are terrestrial ferns with erect or rarely creeping, naked rhizomes. Leaves are spirally arranged and closely placed on the rhizome

Genera and species: Culcitoideae consist of the single genus Culcita with only two species: C. coniifolia in the Neotropics and C. macrocarpa in Europe. Etymology: Culcita is Latin for cushion, in reference to the densely woolly rhizomes, which were harvested to fill pillows and car seat-cushions.

16d. CYATHEACEAE SUBFAMILY PLAGIOGYRIOIDEAE Pheasant-tail ferns

Culcita macrocarpa, private garden, England, UK [16c]

forming a rosette. Petioles are distinct, swollen at base, with knob-like aerophores along the side and with a single V- or U-shaped vascular bundle, quadrangular or triangular, sometimes round in cross-section. Leaves are hairy when young, the hairs excreting mucilage (hence the aerophores), becoming glabrous when mature. The blades are simple and pinnatifid or once pinnate and strongly dimorphic (sterile leaves different from fertile leaves). Sterile pinnae are sessile or adnate to the rachis and are usually serrate. Veins are free, simple or forking and the tips are enlarged near the margins, ending in margin teeth. Fertile leaves are much contracted, nearly lacking blade tissue and are held upright, the pinnae are narrowly linear, covered below in the sporangia (acrostichoid). Sporangia are borne on veins on the lower side of the lamina; indusia are absent, but the sorus is often (partly) covered by the recurved leaf margins. Sporangia are longstalked and the capsule is asymmetrical with a complete, oblique, thickened annulus. Spores are trilete, tetrahedral, depressed between angles, papillate and tuberculate. The gametophytes are green and heart-shaped. Distribution: Plagiogyrioideae usually occur in mountain areas in the tropics, where they can occur at high elevations. They are found in the Neotropics from Mexico to Bolivia,

Plagiogyria falcata, Taiwan (RK) [16d]

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CYATHEALES southeastern Brazil and the Greater Antilles and in Asia from the eastern Himalayas to New Guinea and Japan (Hokkaido). Plagiogyrioideae are most species-rich in East Asia. Phylogeny and evolution: Previously Plagiogyria was included in Blechnum (Aspleniaceae) due to a striking superficial resemblance, but it was already then known that the oblique annulus was a character more closely associated with the tree ferns than with Blechnum. Molecular phylogenetic analyses confirm this and place Plagiogyria as sister to Culcita in the family Cyatheaceae. Genera and species: Plagiogyrioideae consist of the single genus Plagiogyria, with c. 15 species.

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persistent roots and petiole bases, and are densely clothed in long yellowish-brown multicellular hairs. Leaves are clustered at the rhizome apex. Petioles have pneumatophores forming a line on each side and three vascular bundles. Blades are two or three-pinnatepinnatifid, firm and frequently whitish below. Veins are free, simple, forked or pinnate, reaching the leaf margin. Fertile and sterile segments are similar. Sori are formed at a vein ending and are marginal with two indusia, the outer indusium differentiated from the lamina, the inner tongue-shaped. The two indusia are fused at the base. Sporangia are usually formed among hairs. Spores are tetrahedralglobose, often with a prominent equatorial flange and a coarse ridge. The gametophytes are green and heart-shaped.

Legend: Cibotium barometz is the “vegetable lamb of Tartary”, which was harvested for symbolic and ritual purposes. The legendary vegetable lamb was originally described as “being both animal and plant and was a fruit of a tree that sprang from a seed like a melon or gourd that, when fully ripe, burst open and contained a perfectly formed little lamb”. This original description may have actually referred to ‘vegetable wool’ or cotton (Gossypium, Malvaceae), but in later descriptions fantasy took over and the vegetable lamb was described as “a lamb from flesh, bone and blood that was attached by its navel on a stem growing from the ground, sufficiently flexible to allow the lamb to graze on the vegetation around it”. When turned upside down, a rhizome of Cibotium with four cut petioles somewhat resembles a sheep and these creations, popular in curiosity cabinets of the Enlightenment, probably originated from toy-like figures formerly made in China from the rhizomes in that fashion.

Etymology: Plagiogyria is derived from the Greek πλάγιος (plagios), sideways or oblique and γύρος (gyros), round, referring to the oblique annulus.

Distribution: The subfamily occurs in southern Mexico and Mesoamerica (two species), in tropical East Asia from northeastern India to Japan and Borneo (three species) and on the Hawaiian Islands (six species).

16e. CYATHEACEAE SUBFAMILY CIBOTIOIDEAE Vegetable lambs

Phylogeny and evolution: There is some morphological similarity with Dicksonioideae, to which this subfamily is related.

These huge terrestrial tree-like ferns have massive, creeping to erect rhizomes, with

Genera and species : Cibotioidae consist of a single genus Cibotium with 11 species.

Etymology: Cibotium is derived from the Greek κιβωτίων (kibotion), a chest, container or box. Barometz is the Tartar name for the Scythian lamb, referring to the woolly rhizome that resembles a lamb with the legs being formed by the petiole bases.

Cibotium schiedei, Royal Botanic Gardens, Kew, UK [16e]

Cibotium barometz, rhizome, Singapore Botanic Garden [16e]

The barometz or vegetable lamb, from H. Lee (1887), The vegetable lamb of Tartary, London [16e]

Christenhusz, Fay & Chase

CYATHEALES

FERNS

Sphaeropteris intermedia, Jardin des Plantes, Paris, France [16f]

Cyathea arborea, Guadeloupe [16f]

Cyathea grandifolia, Guadeloupe [16f]

Alsophila imrayana, Guadeloupe [16f]

16f. CYATHEACEAE SUBFAMILY CYATHEOIDEAE Scaly tree-ferns

hairy, the pinnae or pinnules are variously shaped, depending on species and placement on the leaf. Veins are pinnate, free and simple or forked or anastomosing, forming arches along the midrib, usually the veins not reaching the margin. Sori are formed dorsally on the veins (rarely terminally), round, the receptacle is strongly raised or not. Indusia are attached around the receptacle bases and sometimes rupture upon opening, attached at one side only and scale-like or completely absent. Sometimes (as in Sphaeropteris) the indusium is replaced by a group of scales around the receptacle. Sporangia are stalked and usually surrounded by multicellular hairs, with a nearly vertical (slightly or strongly oblique) complete annulus that bypasses the sporangium stalk. Spores are trilete and globose. Gametophytes are green and heart-shaped.

some of the smallest. They are more common and more diverse in montane areas in the tropics, but a suite of species prefers lowland rainforests.

These are often large, occasionally tiny (Cyathea subgenus Hymenophyllopsis), terrestrial or epilithic ferns. The rhizomes are usually long and erect, slender or robust, usually tree-like and erect, rarely scandent, sometimes short and ascending or long creeping, often covered in a mantle of adventitious roots and bearing prominent leaf scars. The rhizomes are covered in non-clathrate scales, which distinguish Cyatheoideae from Dicksonioideae (that have only hairs on the tips of the trunks). Leaves are spirally arranged at the tips of the rhizomes. Petioles bear scales that are often inserted on wart- or spine-like outgrowths and are often also hairy. Petioles have discontinuous lines of pneumathodes, and vascular bundles are grouped in two arches, but often with additional series within this, or the vascular bundles are reduced to two simple ones only. Blades are pinnate or bipinnate-pinnatifid, rarely simple or tripinnate-pinnatifid, sometimes with the lowest pinnae strongly reduced (aphlebia), monomorphic. The lamina is scaly and usually

Distribution: This is a pantropical subfamily, extending into warm temperate zones in Australia, New Zealand, southern South America, southern China and southern Japan. Cyatheoideae are widely distributed and reaches many tropical islands. The subfamily includes the tallest extant ferns, but also

Phylogeny and evolution: Cyatheoideae are related to the other tree ferns, especially closely to Dicksonioideae. The oldest fossils assigned to the subfamily date from the late Jurassic. Being the most diverse and dominant tree fern lineage, Cyatheoideae are often associated with the Carboniferous forests, but the dominant tree ferns in that era were Marattiales. Molecular studies have shown that the genera Cnemidaria, Trichipteris and Hymenophyllopsis fall within Cyathea, all sharing morphological characters. Genera and species: This subfamily includes one to five genera (depending on the taxonomist; we accept three here), with c. 635 species: Alsophila (c. 260), Cyathea (c. 275) and Sphaeropteris (c. 100). Etymology: Cyathea is derived from the Greek κυάθιος (kyathios), a cup or beaker, in reference to the cup-shaped indusia of some species.

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CYATHEALES

Dicksonia antarctica, private garden, Kingston upon Thames, Surrey, UK [16g]

Calochlaena dubia, Helsinki Botanical Garden, Finland [16g]

Lophosoria quadripinnata, University of California Botanical Garden, Berkeley, USA [16g]

16g. CYATHEACEAE SUBFAMILY DICKSONIOIDEAE Hairy tree-ferns

are similar to the lamina. They surround the sorus forming a cup or the sori are exindusiate (Lophosoria). The central receptacle is hairy and raised or not. Sporangia are stalked with a complete, oblique or subvertical annulus. Spores are globose, sometimes angular and variously ornamented. Gametophytes are green, heart-shaped and somewhat irregular.

Culcita, but it differs from Culcitoideae in numerous characters. The two subfamilies are not closest relatives. The relationship with Metaxyoideae is also distant.

Distribution: This subfamily occurs in mountain areas of the Neotropics from Mexico to Venezuela and southeastern Brazil and temperate Chile, the Greater Antilles and on the Juan Fernández Islands, on St Helena in the mid Atlantic, and in tropical Asia from the Philippines, through Indonesia into eastern Australia, Tasmania, New Zealand and Samoa, with the greatest diversity in the mountains of New Guinea.

Etymology: Dicksonia was named for Scottish botanist James Dickson (1738–1822), who studied cryptogamic plants under the guidance of Joseph Banks.

These terrestrial ferns have roots that often form a thick mantle around the trunk-like rhizomes that can be erect and up to several metres tall in most Dicksonia species, but are creeping-ascending and sometimes branching in the other genera of the subfamily. The trunk apices are covered with multicellular hairs, distinguishing them from Cyatheoideae (which bear scales at the trunk apex). Petioles have three vascular bundles or have the bundles converging into a single U-shaped bundle in cross-section. The petiole base is clothed in hairs, and pneumathodes form a continuous line along the sides of the petioles or are small, interrupted and inconspicuous, sometimes absent. The petioles have proliferating buds in Lophosoria. Blades are bi- to quadripinnate-pinnatifid, gradually narrowed at the base and apex and are often persistently hairy. Pinnules are usually serrate or lobed. Veins are free, simple, paired or forked at base or pinnate. Fertile pinnae are not or slightly reduced. Sori are formed terminally on a single vein. The indusia are basally fused and

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FERNS

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Phylogeny and evolution: Lophosoria was often placed in its own family because it appears morphologically intermediate between Cyathea and Dicksonia. Molecular evidence places it with Dicksonia and Calochlaena in Dicksonioideae. It is distinctive in its flanged spores. Calochlaena was previously considered part of the genus

Genera and species: Dicksonioideae consist of three genera with c. 29 species: Calochlaena (6), Dicksonia (22) and Lophosoria (1).

16h. CYATHEACEAE SUBFAMILY METAXYOIDEAE Silk ferns These terrestrial ferns have creeping or ascending, short rhizomes that are clothed with silky multicellular hairs. Their petioles have a single Ω-shaped vascular bundle in cross-section and have lateral pneumathodes. Petioles are hairy or become glabrous and sometimes have proliferous buds at their base. Blades are once pinnate, the terminal pinnae similar to the lateral ones. Pinnae are stalked and lanceolate. Veins are free, paired or once

POLYPODIALES

FERNS

forked, the ends merging with the thickened margin. Sori are irregularly arranged on the lower side of the pinnae, with one or several on a vein, rounded, without indusia, the receptacle with thin hairs. The sporangia are many per sorus, shortly stalked, with a vertical, complete annulus bypassing the stalk. Spores are trilete and globose. Gametophytes are green, heart-shaped, with reproductive organs on the upper side and a primitive type of antheridium. Juvenile plants have much dissected leaves, looking unlike adult plants.

Distribution: This subfamily is restricted to tropical America from Mexico to Bolivia, Amazonia, the Guianas and Trinidad. Formerly it also occurred on Guadeloupe, where it is now extinct. Phylogeny and evolution: Morphologically Metaxyoideae have the greatest similarity to Lophosoria. The subfamily has been shown to be close to Dicksonioideae in molecular phylogenetic analyses. It is not known from the fossil record.

Metaxya rostrata, Loreto, Peru [16h]

Genera and species: Metaxyoideae consist of the single genus Metaxya, with seven species. Etymology: It is derived from the Greek μεταξύ (metaxy), intermediate, a name in the past also used for silk traders, intermediate between East and West. This fern has silky rhizome hairs, which may be the reason for its name, although the etymology of the name was not documented. Metaxya parkeri, specimen collected by l’Herminier in 1873 from Guadeloupe, where this species is now extinct (Kew Herbarium) [16h]

POLYPODIALES Families 17 to 24 represent the order Polypodiales or the ‘polypod ferns’. This includes some early branching lineages such as Lindsaeaceae, the pteridoid lineage (Dennstaedtiaceae and Pteridaceae) and the two eupolypod lineages, Aspleniaceae and Polypodiaceae. Polypodiales include the vast majority of extant ferns.

17. LONCHITIDACEAE Velvet-fern family

These terrestrial ferns have stout, shortcreeping, fleshy often green rhizomes. All parts are covered in pale hairs. Petioles arise on the upper side of the rhizome only, leaving round scars when withered, and have two vascular bundles that are well developed. Blades are succulent, once to three times pinnate-pinnatifid, the pinnae subopposite, subsessile and adnate to the rachis or shortly stalked, and the margins entire and lobed. Veins are free or irregularly joining,

the areoles without included veinlets. Sori are marginal, joining several veins, and are confined to the sinuses between lobes or occur along the entire blade margin. Indusia are elongate, formed by the reflexed, modified leaf margin. Receptacles are hairy, and sporangia formed among these hairs have three-celled stalks; capsules have a defined annulus. Spores are trilete, globose and granulate. Gametophytes are green and heart-shaped.

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POLYPODIALES

FERNS

Saccoloma dominguense, Guadeloupe [18]

46

Lonchitis hirsuta, Guadeloupe [17]

Saccoloma inaequale, Brazil [18]

Saccoloma dominguense, Guadeloupe [18]

Distribution: The family occurs in tropical America and tropical Africa and Madagascar, usually locally abundant in moist tropical forests, often along streams.

Genera and species: Lonchitidaceae consist of the single genus Lonchitis with two species: L. hirsuta in the Neotropics and L. occidentalis in tropical Africa.

Phylogeny and evolution: Traditionally Lonchitis was placed in Dennstaedtiaceae, close to Blotiella, which it resembles, but these genera differ markedly in rhizome structure, spore morphology and chromosome number. More recently, it was placed in Lindsaeaceae, where some molecular analyses place it, always with weak support. The placement of Lonchitidaceae is among the early polypods, a still unresolved part of the fern phylogenetic tree. It appears that early polypods evolved rapidly, and therefore the independent lineages are difficult to place. Because including Lonchitis in Lindsaeaceae would stretch the morphological circumscription of that family and relationships among early polypods remains uncertain, it has now been accepted as a separate family.

Etymology: Lonchitis is derived from the Greek λόγχ (longch), a lance or spear.

arranged spirally on all sides and have a single vascular bundle. Blades are decompound or (rarely) once pinnate. Veins are free, simple or forked and do not reach the margins. Sori terminate a single vein. The indusium is cupor pouch-shaped and formed close to the margins. Receptacles are glabrous. Sporangia have three-celled stalks and capsules with a defined annulus. Spores are trilete, tetrahedral-globose, prominently ridged and granulate. Gametophytes are green and heart-shaped.

Christenhusz, Fay & Chase

18. SACCOLOMATACEAE Pouch-fern family

Distribution: Saccolomataceae are widespread in the Neotropics, Madagascar, tropical Asia, Australasia and the western Pacific. They are usually found in shady banks in tropical rainforests.

These are terrestrial ferns with trunk-like or short-creeping, scaly rhizomes. Hairs are absent from all parts of the plant. Petioles are

Phylogeny and evolution: This distinctive family has often been placed in Dennstaedtiaceae, but even within that has long been recognised as a subfamily. They form one

POLYPODIALES

FERNS

of the early branching polypod lineages, and its placement is not certain; it is clear though that they are not close to Dennstaedtiaceae. Because of their distinctive morphology and molecular isolation, they are treated in their own family. Genera and species: Saccolomataceae consist of about 12 species that are usually included in the single genus Saccoloma, although species with decompound leaves have been separated into Orthiopteris.

and sparsely hairy. Pinnules are sessile, oblong and serrate. Veins are simple, paired or forked. Fertile pinnules are slightly contracted, the sori are formed marginally, terminating a vein, the vein extending into two small indusium lobes, the larger of the two lobes dome-shaped, covering the sorus, with numerous hairs among the sporangia. The sporangia are stalked with a complete annulus. Spores are globose, ridged and rugulose. Gametophytes are green and heart-shaped.

Genera and species: Cystodiaceae consist of the single genus Cystodium with only one species: C. sorbifolium. Etymology: Cystodium is derived from the Greek κυστός (kystos), a bladder or pouch, in reference to the pouch-shaped indusium.

Distribution: This is a pantropical family with representatives in temperate Africa, Brazil, southern Australia, New Zealand, the

Distribution: Cystodiaceae occur in tropical Asia from Borneo eastwards to the Admiralty and Solomon Islands.

19. CYSTODIACEAE

Phylogeny and evolution: Historically Cystodium was placed among the tree ferns in Dicksonioideae, but morphology and molecular analyses place it clearly in Polypodiales. It had been in Lindsaeaceae, but that is not warranted on the basis of morphology and phylogenetics; it is often found in analyses to have a more isolated position among the early polypods.

These terrestrial ferns have creeping to erect stems with long hairs. Petioles are densely hairy at base and have two vascular bundles separated by an X-shaped bundle in crosssection. Blades are bipinnate-pinnatifid

Cystodium sorbifolium, fiddlehead and rhizome scales, Solomon Islands (CC) [19]

Lace-fern family

These are terrestrial or epiphytic ferns with creeping, scaly rhizomes. Rhizome scales are basally attached and not clathrate. Petioles have a single vascular bundle. Blades are variable in shape, simple or one to four times pinnate, decompound or finely divided, glabrous when mature. Some species of Odontosoria have spiny, vining rachises. Veins are free or anastomosing, the areoles, when present, without included veinlets. Sori form terminally, often uniting several vein ends near the margin. Indusia are basally and laterally attached, opening towards the margin. Receptacles are usually hairy, the sporangia have three-celled stalks and a defined annulus. Spores are trilete, rarely monolete, globose to ellipsoid, smooth or variously ornamented. Gametophytes are green and heart-shaped.

Etymology: Saccoloma is derived from Greek σάκκο (sacco), a bag and λόμα (loma), an edge or border.

Rowan-fern family

20. LINDSAEACEAE

Cystodium sorbifolium, sorus detail, Solomon Islands (CC) [19]

Cystodium sorbifolium, Solomon Islands (CC) [19]

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POLYPODIALES

Odontosoria chinensis, Tahiti [20]

FERNS

Tapeinidium moorei, New Caledonia [20]

Lindsaea orbiculata, Hong Kong [20]

Nesolindsaea kirkii, Seychelles [20]

Odontosoria deltoides, sori, New Caledonia [20]

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Lindsaea quadrangularis subsp. antillensis, Guadeloupe [20]

POLYPODIALES

FERNS

Himalayas and southern Japan. They are found mostly in wet, lowland or montane forests but sometimes grow in more open habitats. Phylogeny and evolution: Lindsaeaceae were traditionally associated with Dennstaedtiaceae, but instead they form one of the early branching lineages in Polypodiales, a sister lineage to all other polypods. Because Dennstaedtiaceae form a clade with Pteridaceae, Lindsaeaceae are not closely related to that family. Genera and species: Lindsaeaceae consist of six genera with about 220 species: Lindsaea (c. 175), Nesolindsaea (2), Odontosoria (c. 20), Osmolindsaea (7), Sphenomeris (1) and Tapeinidium (19). Uses: The leaves of Odontosoria chinensis were collected in Hawaii to make a red dye. Etymology: Lindsaea was named for Jamaican surgeon John Lindsay (1750–1803), who was the first to describe germination of fern spores.

21. DENNSTAEDTIACEAE Bracken family

These terrestrial ferns have usually longcreeping, sometimes ascending erect rhizomes (Blotiella) that are hairy and often also scaly. Petioles are placed on the upper side of rhizomes, with one U-shaped or two parallel, upwardly merging vascular bundles. Blades are often large, bi- to tripinnatepinnatifid, and often hairy. Veins are free, forked or pinnate, rarely merging and then without included veinlets. Sori are formed marginally or close to the margins, are linear or round and terminate one or many veins.

Indusia are linear or cup-like, fixed basally and laterally, opening towards the margins, or the margin is reflexed over the sorus. Spores are trilete or monolete, tetrahedral or reniform. Gametophytes are green and heart-shaped.

Genera and species: Dennstaedtiaceae consist of ten genera with some 240 species: Blotiella (16), Dennstaedtia (c. 70), Histiopteris (c. 7), Hypolepis (c. 50), Leptolepia (1), Microlepia (c. 60), Monachosorum (6), Oenotrichia (2), Paesia (14) and Pteridium (c. 13).

Distribution: Dennstaedtiaceae are a family with a global distribution, species of the genus Pteridium (bracken) occur almost everywhere, except in extremely arid and permanently frozen areas. The diversity of the family is greatest in tropical mountains. Blotiella is most diverse in Africa, Oenotrichia is confined to New Caledonia and Monachosorum is restricted to tropical Asia. By contrast, Histiopteris incisa is widespread, occurring in tropical mountains and in temperate rainforests across the Southern Hemisphere.

Uses: Asian bracken (Pteridium esculentum) is frequently used in Asian cooking, most commonly the young leaves (fiddleheads) are are stir-fried. Starch from the roots is also extracted and made into dumplings considered a delicacy, although they may have a constipating effect. Asian bracken is known to be carcinogenic, so caution should be taken when eating fiddleheads, especially because they are easily confused with other, more toxic bracken species. European bracken (Pteridium aquilinum) produces a dye of various shades of green from the leaf tips, and the rhizome is used in Scotland to make the dark-yellow dye found in some tartans. Dennstaedtia glauca is used as a green manure in the Andes.

Phylogeny and evolution: Dennstaedtiaceae are an old family that were traditionally associated with Lindsaeaceae or Pteridaceae. Molecular analyses place them as sister to Pteridaceae, which together are sister to the ‘eupolypods’ (Aspleniaceae + Polypodiaceae). Pteridium pubescens, New Mexico (DZ) [21]

Etymology: Dennstaedtia is named for German botanist August Wilhelm Dennstädt (1776–1826). Pteridium pinetorum, Finland [21]

Blotiella stipitata, Kenya [21]

Histiopteris incisa, New South Wales, Australia [21]

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POLYPODIALES

FERNS

Hypolepis repens, Guadeloupe [21]

Dennstaedtia punctilobula, New York, USA [21]

22. PTERIDACEAE Ribbon-fern family

Dennstaedtia bipinnata, Mary Selby Botanical Garden, Florida, USA [21]

Sporangia have a vertical interrupted annulus. Spores are trilete, globose or tetrahedral and variously ornamented. Gametophytes are green, heart- or ribbon-shaped and sometimes with filamentous gemmae along the margins. Distribution: Pteridaceae are nearly cosmopolitan with their greatest diversity in dry regions and the wet tropics, but with some taxa extending into higher latitudes.

These terrestrial, epilithic or epiphytic ferns have creeping, ascending or erect rhizomes that are usually scaly (rarely only with hairs). Petioles are hairy or glabrous, have a U-shaped vascular bundle in cross-section and often bear proliferating buds at the base. Blades are monomorphic or dimorphic, simple, pinnate, pedate or decompound, glabrous or covered with hairs and/or scales. Veins are free and forking, flabellate or variously anastomosing and reticulate, the areoles without included veinlets. Sori are formed marginally or along veins and lack a true indusium, although often with a ‘false indusium’, e.g. a modified flap of the leaf margin recurved over the sorus.

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Phylogeny and evolution: Pteridaceae were traditionally variably defined, but molecular studies have helped greatly to redefine this group. The family is monophyletic when the traditional families Adiantaceae, Cheilanthaceae, Parkeriaceae, Platyzomaceae and Vittariaceae are included. Pteridaceae are sister to Dennstaedtiaceae and are composed of five subfamilies (Ceratopteridoideae, Cheilanthoideae, Cryptogrammoideae, Pteridoideae and Vittarioideae), which are discussed below. The family originated in the Late Cretaceous. Genera and species: Pteridaceae consist of about 45 genera, perhaps fewer after generic

recircumscription, with c. 1,150 species. The largest genera are Pteris (c. 300 species) and Adiantum (c. 200 species). The five subfamilies are treated separately below. Uses: The leaves of maidenhair fern (Adiantum capillus-veneris) can be simmered in water for several hours, after which it is boiled into a syrup with sugar called ‘sirop de capillaire’, which can then be mixed with fruit juices to make a refreshing drink. Allosorus pteridioides can be used as a tea substitute. Ceratopteris species are popular aquarium plants, and Ceratopteris cornuta is consumed as a vegetable in Liberia. Fibres from petioles of Adiantum and Pityrogramma have been used by Native Americans to weave black designs in baskets. Many species are grown as ornamental plants, especially species of Adiantum, Pellaea and Pteris, and some species such as Adiantum raddianum, Pteris cretica and P. vittata are commonly naturalised in tropical and temperate regions, especially in urban areas. Etymology: The Greek πτέρης (pteris), a fern, is derived from πτερων (pteron), a feather, in reference to their feathery leaves.

POLYPODIALES

FERNS

22a. PTERIDACEAE SUBFAMILY CRYPTOGRAMMOIDEAE Parsley ferns These terrestrial ferns have creeping or decumbent to nearly erect rhizomes. The rhizome scales are lanceolate, brown and rigid. Leaves are monomorphic (Coniogramme) to strongly dimorphic (Cryptogramma), and then the fertile leaves are more erect with contracted segments, or they are partly dimorphic, with the apical fertile part of the leaf having longer and more slender segments (Llavea). Petioles have one vascular bundle. Blades are one to four times pinnate or two times pinnate-pinnatisect, with the terminal segment similar to the lateral ones, glabrous or sometimes sparsely hairy, sometimes glaucous beneath. Veins are free or anastomosing marginally, the vein ends often being somewhat enlarged. Sporangia are borne on the veins in elongated soral lines, without an indusium, but the sorus is frequently covered by the modified leaf margin. Spores are trilete, angular-globose, verrucate, tuberculate or rugulose. Distribution: These ferns are found in North America, north to Alaska and south to Mexico and Guatemala, southern Chile and adjacent Argentina, throughout Europe, mostly in the north and in mountains, throughout Siberia and the Urals to the Himalayas and East Asia, also in tropical Africa, Java and some Pacific islands. Phylogeny and evolution: Cryptogrammoideae are the first diverging clade in Pteridaceae. They were previously placed in Cheilanthoideae on the basis of superficial

Cryptogramma crispa, Scotland, UK [22a]

morphological similarities, but are not immediate relatives. Cryptogramma occurs around the Arctic, in mountains in the temperate zones and in southern South America, whereas Coniogramme is more a species of humid subtropical lands in the Old World. Llavea cordifolia, endemic to Mexico and Guatemala, was previously not placed in the subfamily, but is now known to belong here as well. Genera and species: Cryptogrammoideae consist of three genera with c. 41 species: Coniogramme (c. 30), Cryptogramma (c. 10) and Llavea (1). Etymology: Cryptogramma is derived from the Greek κρυπτος (kryptos), hidden and γραμμή (grammio), a line, in reference to the linear sorus that is hidden by the reflexed leaf margin.

22b. PTERIDACEAE SUBFAMILY CERATOPTERIDOIDEAE Mangrove ferns These are terrestrial, submerged or floating aquatic ferns. Rhizomes are short, small and erect or stout and creeping. Rhizome scales are thin, few and small or many and large. Petioles have two larger and several smaller or four larger and several smaller vascular bundles. Leaves are partially or completely dimorphic, in Acrostichum once pinnate, the fertile parts covering the entire lower part of the pinna blade, in Ceratopteris the sterile leaves lobed to three times pinnate and fertile leaves contracted, one to five times pinnate. Blades are glabrous, and the veins are

Acrostichum aureum, Seychelles [22b]

reticulate without included veinlets. Sporangia cover the entire lamina (Acrostichum) or form along the veins and covered by a modified marginal indusium (Ceratopteris). Spores are angular, globose, with coarse parallel ridges or with tubercles and papillae. Distribution: This is a pantropical subfamily, usually occurring in flooded or seasonally inundated places or mangroves. Phylogeny and evolution: Difficult to define, Ceratopteridoideae have shifted their morphology as a consequence of adaptation to aquatic habitats. These characters also made the species difficult to place in previous classifications. The genus Ceratopteris was previously placed in the separate family Parkeriaceae, with chromosome numbers and spores showing a distant affinity with Anemiaceae, but the fertile structures show a more clear affinity with Pteridaceae. Acrostichum is a genus that had a broad application in the past, but is now restricted to a few species of semi-aquatic mangrove ferns in the tropics. It was previously believed to be associated with Pteridoideae, but molecular evidence places it with Ceratopteris as sister to Pteridoideae. Genera and species: Ceratopteridoideae consist of two genera with nine species: the large mangrove and swamp ferns Acrostichum (4), and the common fresh water aquatic Ceratopteris (5). Etymology: Ceratopteris is derived from the Greek κέρας (keras), a horn, and πτέρης (pteris), a fern.

Ceratopteris pteridoides, Colombia (MF) [22b]

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POLYPODIALES

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FERNS

Pityrogramma calomelanos, Réunion [22c]

Jamesonia goudotii, Ecuador [22c]

Anogramma leptophylla, Lebanon [22c]

Pteris usambarensis, sori, Kenya [22c]

Taenitis blechnoides, Singapore [22c]

Pteris vittata, Brazil [22c]

22c. PTERIDACEAE SUBFAMILY PTERIDOIDEAE Brakes

absent, but often the sori are covered by the reflexed leaf margin. Spores are trilete, usually angular, globose and variously ornamented.

These terrestrial and epilithic ferns have creeping or erect rhizomes with non-clathrate entire scales or rigid bristles. Petioles have one to four or many (and then usually with two larger) vascular bundles. Leaves can be monomorphic, slightly dimorphic or the fertile leaves can be different from the sterile ones. Blades are simple, trifoliate, or one to five times pinnate, pedately organised or forking from a central point. Veins are free or reticulate without included veinlets. Sori are formed submarginally, joining several vein ends, or along anastomosing veins. The indusium is usually

Distribution: This subfamily occurs throughout the tropics and subtropics of the world, north to the southern USA, southern Europe and East Asia and south to southern South America, Australia and New Zealand. They are uncommon in frost-prone areas.

Christenhusz, Fay & Chase

Phylogeny and evolution: Pteridoideae form with Cryptogrammoideae and Ceratopteridoideae the sister clade to the remainder of the family. They include an expanded Jamesonia (including Eriosorus and Nephopteris) and Pteris, the latter now including Afropteris,

Anopteris, Litobrochia and Neurocallis. Platyzoma, formerly in Platyzomaceae, is also placed in Pteris, although separation as a subgenus is warranted. The relationships between the species of the large genus Pteris need further study. Genera and species: Pteridoideae consist of 15 genera with c. 445 species: Actiniopteris (4), Anogramma (3), Aspleniopsis (1), Austrogramme (5), Cerosora (3), Cosentinia (1), Gastoniella (3), Jamesonia (c. 51, Onychium (c. 10), Pityrogramma (c. 20), Pteris (c. 300), Pterozonium (14), Syngramma (15), Taenitis (c. 15) and Tyronia (4). Etymology: Pteris is Greek for fern.

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Pellaea mucronata, University of California Botanical Garden, Berkeley, USA

[22d]

Doryopteris concolor, Brazil [22d]

22d. PTERIDACEAE SUBFAMILY CHEILANTHOIDEAE Lip ferns These are terrestrial and epilithic ferns, often with adaptations to arid environments. Rhizomes are erect to long-creeping with scales and sometimes hairs. Petioles have one or two vascular bundles. Leaves are undivided or one to five times pinnate, pedately or sometimes palmately compound. The blades are often scaly, sometimes hairy, glandular or glabrous. Veins are usually free or sometimes anastomosing. Sporangia are borne along veins in lines or along the margin and then often covered by a modified marginal indusium. Spores are trilete and often cristate. Distribution: Cheilanthoideae occur throughout the subtropics and tropics, extending into the temperate zones in North America, southern Europe, South Africa, East

Cheilanthes austrotenuifolia, Mt Benia, Western Australia [22d]

Asia and New Zealand. They are especially diverse in the American deserts. Phylogeny and evolution: This subfamily is remarkable in its adaptation to arid conditions, with the leaves being able to revive after desiccation in many species. The lineage forms a natural entity, but the internal generic circumscription is still in great need of study. Molecular work is needed to redefine these genera because two of the best known genera, Cheilanthes and Pellaea in their traditional sense, are widely polyphyletic, and it is possible that all (currently about 25 genera) should be united in a single genus, Hemionitis, the oldest name for this group. The genus Gaga, which received much media attention as it was named for musician Lady Gaga, belongs to the clade that includes Aspidotis, an older name. Genera and species: Cheilanthoideae consist

of c. 310 species in c. 13 genera: Allosorus (c. 50), Aspidotis (24), Bommeria (5), Calciphilopteris (4), Cheilanthes (c. 50), Cheilosoria (c. 50), Doryopteris (c. 40), Hemionitis (8), Mildella (1), Notholaena (c. 30), Pellaea (c. 40), Pentagramma (2) and Pteridella (c. 5). The generic circumscription is far from perfect and needs significant taxonomic and nomenclatural attention because many traditional genera are polyphyletic. The consensus of c. 13 poorly defined genera is an intermediate solution, but it can equally be argued that all are better united into a single genus, with Hemionitis having nomenclatural priority. Both solutions will require large numbers of new combinations. Etymology: Cheilanthes is derived from the Greek χείλος (cheilos), a lip and άνθος (anthos), a flower, in reference to the edge of the pinnules, which form a lip covering the sporangia.

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22e. PTERIDACEAE SUBFAMILY VITTARIOIDEAE Maidenhair and ribbon ferns These are terrestrial, epilithic and epiphytic ferns. Rhizomes are erect, suberect or creeping, sometimes with the leaves widely spaced, sometimes densely clustered. Rhizome scales are thin, ovate-lanceolate and brown, clathrate or not. Leaves are monomorphic or nearly so. Petioles have one or two vascular bundles, which are often short or nearly absent in epiphytic species. Blades are simple, and then entire and linear, sagittate or forked or cleft, or one to six times pinnate, or the petiole divided into two forming a pedate blade, usually glabrous, sometimes pubescent, thin herbaceous or thick and leathery. Veins are simple or forked and free or sparingly anastomosing, sometimes reticulate, but areoles without included veinlets. Sporangia are borne along

Haplopteris ensiformis, Singapore [22e]

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veins in short lines near the margin, often covered by a modified leaf margin (Adiantum) or in simple or branched lines along the veins or immersed in a groove along the midvein. Spores are trilete or monolete, hyaline or not, globose, prominently angled and variously ornamented or smooth. Distribution: Pantropical, with a few representatives in humid temperate regions, north to Alaska and Canada, western Europe, the Himalayas and northeastern Asia, south to southern South America, South Africa, Australia and New Zealand. Phylogeny and evolution: Vittarioideae are composed of the former Vittariaceae plus Adiantum. Apart from Adiantum, they are all epiphytes with finely matted roots, mostly simple blades and sori along the veins or in lateral grooves along the midveins. The association with Adiantum seems odd at first,

Antrophyum boryanum, Réunion [22e]

but they do share some anatomical characters, such as the presence of ‘spicular cells’ in the leaf epidermis. With Cheilanthoideae, Vittarioideae form the crown group in Pteridaceae that together are sister to a clade uniting the other subfamilies. Vittaria in the traditional sense was polyphyletic and has therefore been broken up. Genera and species: Vittarioideae consist of c. 11 genera, with species: Adiantum (c. 200), Ananthacorus (1), Anetium (1), Antrophyum (c. 40), Haplopteris (c. 40), Hecistopteris (3), Monogramma (9), Polytaenium (18), Radiovittaria (5), Scoliosorus (2) and Vittaria (c. 10). It has been debated how many genera should be recognised in Vittarioideae. Etymology: Vittaria is derived from the Latin vitta, a band or ribbon, in reference to the narrowly linear leaves of this genus.

Adiantum capillus-veneris, Sicily, Italy [22e]

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23. ASPLENIACEAE Spleenwort family

These are terrestrial, epiphytic and epilithic (rarely aquatic or climbing) ferns that usually have blackish and wiry roots (rarely fleshy). Their rhizomes are erect, sometimes trunklike or short- to long-creeping, stoloniferous or scandent in some species, branched or not, with lanceolate to linear rhizome scales. The leaves are monomorphic, slightly dimorphic or strongly dimorphic, gemmiferous or not, variously scaly or hairy, brightly coloured when young in several genera. Petioles have (mostly) two vascular bundles. Blades are simple or one to four times pinnate-pinnatifid, in some taxa with conspicuous aerophores along the rachises. Veins are free or anastomosing, reaching the margin or terminating before it (and then often with expanded vein endings), areoles when present usually have no included veinlets. Sori are formed on a flat or raised receptacle, usually along veins on the lower side of the blade, and are often elongate along the vein, placed in vein forks and triangular or circular and placed atop a vein, rarely acrostichoid (covering the entire blade). The indusium can be flat and laterally attached along one (asplenioid) or both sides of the vein (diplazioid) or in the axils of veins and the indusia merged in v- or j-shapes, or hood-like and attached at the base of the sporangium, or reniform and atop a vein, or the indusium can be absent. Sporangia are (long-)stalked, the stalks can be up to three cells wide. Spores are not or occasionally chlorophyllous, monolete, variously ornate. Gametophytes are green, superficial and heart-shaped. Distribution: This family is globally distributed and can be found in all areas except in permanently frozen or arid areas.

Phylogeny and evolution: Also known as ‘eupolypods II’, Aspleniaceae have been expanded to include in this clade the former families Athyriaceae, Blechnaceae, Onocleaceae, Thely pteridaceae and Woodsiaceae. These families are all closely related and share the usually linear (sometimes circular) sori organised along the veins with a laterally attached indusium (when present), the blackish, wiry roots and two vascular bundles in the petioles. Aspleniaceae split from Polypodiaceae in the Late Jurassic, c. 104–107 million years ago. Aspleniaceae diversified into their major subdivisions during the Late Cretaceous and Early Tertiary. Genera and species: Aspleniaceae include 23 genera and c. 2,780 species. The family is divided into eight subfamilies (Asplenioideae, Athyrioideae, Blechnoideae, Cystopteridoideae, Diplaziopsidoideae, Rhachidosoroideae, Thelypteridoideae and Woodsioideae). They are discussed separately below because they are sometimes treated at the family level. The largest genera are Asplenium (c. 700), Athyrium (c. 200), Blechnum (c. 250), Diplazium (c. 350) and Thelypteris (c. 1,100). Uses: The young shoots or fiddleheads of the ostrich fern, Onoclea struthiopteris, are commonly consumed in North America. Onoclea orientalis is also used for this purpose in Asia. The young leaves and croziers of the ‘vegetable fern’, Diplazium esculentum, are commonly consumed in stir-fries or in salads in tropical Asia and the Pacific. The leaves are mildly toxic, but no adverse health effects are known. Young fiddleheads of the painted fern Athyrium nipponicum can also be eaten after being thoroughly boiled to leach out the toxins. The young leaves of Asplenium viviparum are cooked and eaten in New Zealand, and in Asia the young shoots of nest fern (A. nidus) are sometimes consumed. Young croziers of Stenochlaena palustris are eaten in Southeast Asia, and the rhizomes of Blechnum orientale have been eaten in Malaysia. The rhizomes of Onoclea and Cystopteris can be peeled and roasted, but these should be

considered only a famine food. The long stems of Stenochlaena palustris produce a durable fibre after submersion in salt water. They are used in the manufacture of fish traps or are twisted into rope. The cortex of the stem of Blechnum cyatheoides was used in Hawaii to produce a red dye. A number of species are cultivated as garden or houseplant ornamentals. Etymology: From Old French esplen, in turn derived from the Greek σπλην (splin), a spleen, in reference to the former use of spleenwort to cure spleen and liver problems.

23a. ASPLENIACEAE SUBFAMILY CYSTOPTERIDOIDEAE Bladder ferns These terrestrial ferns have blackish, wiry roots that emerge from creeping to suberect, branching, scaly rhizomes. Their leaves are monomorphic and bear scales that are sometimes hair-like. The blades are thinly herbaceous, (one or) two to three times pinnate-pinnatifid and deltate or lanceolate in outline, and occasionally bear bulbils that grow into new plants. Veins are free and terminate at the margin. Sori are formed on a hardened receptacle, along veins on the lower side of the blade and are round or slightly elongate. The indusium is hood-like, attached at the base of the sporangium, or absent. Spores are monolete, echinate, tuberculate or broadly folded. Gametophytes are green and heart-shaped with glandular hairs on the wings. Distribution: Cystopteridoideae are distributed across temperate areas of the Northern Hemisphere and in mountains in the tropics, extending south into the Andes, South Africa, tropical Asia, Australia and New Zealand. Phylogeny and evolution: The genera were in past classifications rarely placed together. Gymnocarpium was usually treated as allied to the dryopteridoid ferns, whereas Cystopteris was usually associated with the athyrioids or woodsioids. Gymnocarpium was found to be sister to a clade uniting Cystopteris and

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POLYPODIALES

Cystopteris fragilis, Yorkshire, England, UK [23a]

Gymnocarpium dryopteris, Michigan, USA [23a]

Gymnocarpium fedtschenkoanum, Hortus botanicus, Leiden, the Netherlands [23a]

Acystopteris, and this trio is sister to the rest of Aspleniaceae (‘eupolypods II’). Cystoathyrium, a native of China, has not been recently found and is probably extinct. It is placed here tentatively. Hybrids between Cystopteris and Gymnocarpium are known.

broadest at the base and gradually taper to the apex. Veins are free, terminating before reaching the hyaline margins. Sori are elongate and formed on a flat receptacle on the lower side of the leaves along one side of veins with a laterally attached indusium. Spores are monolete, echinate, tuberculate or broadly folded. Gametophytes are heart-shaped, superficial and green.

Etymology: Rhachidosorus is derived from the Greek ράχης (rhachis), a spine or ridge, and σορός (soros), a coffin or urn, in reference to the placement of the sori.

Genera and species: Cystopteridoideae consist of four genera with c. 30 species: Acystopteris (3), Cystoathyrium (1), Cystopteris (c. 20) and Gymnocarpium (7). Etymology: From Greek κυστός (kystos), a bladder or pouch, referring to the indusium that looks like an inflated bladder, and πτέρης (pteris), a fern.

23b. ASPLENIACEAE SUBFAMILY RHACHIDOSOROIDEAE Lacquer ferns These terrestrial ferns have creeping, usually unbranched, scaly rhizomes. Their scales are clathrate and entire. Leaves are monomorphic and sparsely scaly. Blades are herbaceous and two to three times pinnate-pinnatifid; they are

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Distribution: Rhachidosoroideae occur in East Asia, often in forests on limestone. Phylogeny and evolution: Plants are similar to and difficult to distinguish from Athyrioideae (to which they were often referred in the past), and the subfamily also shares characters with Asplenioideae. Rhachidosoroideae diverged from their nearest relatives c. 90 million years ago, before Blechnoideae diverged from Athyrioideae. Genera and species: Rhachidosoroideae consist of the single genus Rhachidosorus with four to eight species, depending on taxonomic concept.

23c. ASPLENIACEAE SUBFAMILY DIPLAZIOPSIDOIDEAE Glade ferns These terrestrial ferns have fleshy, rarely wiry, roots that grow radially from erect or shortcreeping, usually unbranched, scaly rhizomes. Rhizome scales are entire, not clathrate and lanceolate. Leaves are glabrous or bear filiform scales. Blades are once pinnate, herbaceous or fleshy; the apex is gradually reduced or similar to lateral pinnae. Venation is free or anastomosing toward the hyaline margins, but areoles, when present, lack included veinlets; the free vein-endings are slightly expanded and raised. Sori are borne singularly along one side of the vein (asplenioid), or two sori are formed back to back along the same vein (diplazioid) with the glabrous or glandular indusium attached laterally along the vein. Spores are monolete and folded with erose

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crests. Gametophytes are heart-shaped and glandular-hairy on the wings. Distribution: This subfamily has a disjunct distribution in eastern North America (Homalosorus) and in East and tropical Asia and some Pacific islands (Diplaziopsis). Phylogeny and evolution: Diplaziopsidoideae are similar to Athyriaceae in many respects, and before molecular data were available all species were placed in the genus Diplazium. The family shares many characters with Hemidictyum, which was originally also placed here, but DNA studies have shown that this would make the family polyphyletic with regards to Asplenioideae. One species, Diplazium flavoviride, also belongs to this lineage because it shares characters with Diplaziopsis, but no molecular data are yet available for this species.

Etymology: Diplaziopsis is derived from the Greek, διπλάσιος (diplasios), double, for its double sori, and -opsis, appearance, in reference to its resemblance to Diplazium.

23d. ASPLENIACEAE SUBFAMILY ASPLENIOIDEAE Spleenworts

the veins, rarely two sori are formed back to back along both sides of the vein, and the indusium is laterally attached along the veins and is glabrous. Sporangia are long-stalked and single rowed. Spores are monolete, reniform, usually with sharp ridges or broad folds or wings. Gametophytes are green and heart-shaped. Distribution: This subfamily occurs nearly globally, except for permafrosted or extremely arid regions.

Genera and species: Diplaziopsidoideae consist of two genera with six species: Diplaziopsis (5) and Homalosorus pycnocarpos.

These are terrestrial and epiphytic ferns with blackish wiry roots. Their rhizomes are scaly, short- to long-creeping or erect and usually unbranched. Rhizome scales are entire, lanceolate and clathrate, sometimes with glandular margins. Leaves are monomorphic or slightly dimorphic, occasionally bearing bulbils that grow into new plants. Petioles are variably coloured, often glossy, with two vascular bundles. Blades are undivided (simple) or one to four times pinnate, usually tapering gradually towards the apex, sometimes with a terminal pinna similar to the lateral. Veins are free or reticulate, areoles, when present, not including veinlets. Sori are elongate along one side of

Homalosorus pycnocarpos, Brooklyn Botanical Garden, New York, USA [23c]

Asplenium marinum, Scotland, UK [23d]

Asplenium septentrionale, Finland [23d]

Phylogeny and evolution: Among ferns Asplenioideae are unusual in showing diversification in both tropical and temperate regions, and there are as many epiphytic as terrestrial species. The family is readily distinguishable by their linear sori with indusia attached to one side of the vein. The numerous segregate genera of Asplenium are now generally not accepted because they made Asplenium polyphyletic, and numerous intergeneric hybrids were frequently observed in the wild. The wider

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POLYPODIALES circumscription resulted in a genus with great diversity in morphology and habitat preference. Only Hymenasplenium, which is sister to Asplenium, is maintained, although some authors have suggested that even this genus should be united with Asplenium. The third genus Hemidictyum has always been difficult to place. It was formerly treated as Asplenium or Diplazium and was considered allied to Thelypteridoideae or Diplasiopsidoideae. Molecular studies have shown that it is sister to the remaining Aspleniaceae, and even though it could be maintained in its own subfamily, it is here united with an expanded Asplenioideae as characters are shared. Hemidictyum diverged from the remaining Asplenioideae in the Late Cretaceous. A great number of segregate genera have been proposed (e.g. Camptosorus, Ceterach, Diellia, Loxoscaphe, Phyllitis, Pleurosorus, Thamnopteris etc.), but intergeneric hybrids are commonly encountered. Most segregate genera were found to be embedded in Asplenium in molecular studies. Asplenium australasicum, Moorea, French Polynesia [23d]

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Genera and species: Asplenioideae consist of four genera with c. 716 species: Asplenium (c. 700), Desmophlebium (1), Hemidictyum (1) and Hymenasplenium (c. 15).

These terrestrial ferns have blackish, wiry roots that emerge from short- to long-creeping or scandent, suberect or erect rhizomes that bear lanceolate, non-clathrate, hairy scales. Their leaves are usually conspicuously hairy, and the hairs are often whitish, acicular (curved at the tip) and unicellular, sometimes combined with various types of multicellular hairs (stellate, forked etc.). Young developing leaves are often covered in mucilage, with conspicuous aerophores (breathing tubes) protruding through the

mucilage, which are persistent when leaves are mature. The leaves are monomorphic or subdimorphic, the fertile leaves then contracted, sometimes bearing bulbils. Petioles have two vascular bundles, rarely more, and are not persistent on the rhizome. Blades are simple or one to three times pinnate-pinnatifid, with or without reduced pinnae at the base, the pinna bases often bearing a conspicuous aerophore. Veins are free, sometimes terminating before reaching the margin, merging below a sinus in lobed pinnae, or anastomosing and sometimes with a conspicuous fish-bone pattern (meniscioid); areoles can have included veinlets, but not always. Sori are formed on top of veins and are circular or elongate, indusiate or exindusiate, on a flat receptacle. Indusia are lateral and reniform when present. Sporangia are stalked, the stalks three cells wide in the middle. Spores are monolete, brown and achlorophyllous, reticulate, echinate or sharply crested. Gametophytes are green and heartshaped, often somewhat elongate and without a distinct midrib, frequently bearing hairs or stalked glands.

Hemidictyum marginatum, Brazil [23d]

Phegopteris hexagonoptera, North Carolina [23e]

Etymology: Asplenium is derived from Greek σπλην (splin), a spleen, in reference to the former medicinal use of spleenwort to cure spleen and liver diseases.

23e. ASPLENIACEAE SUBFAMILY THELYPTERIDOIDEAE Marsh ferns

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Macrothelypteris torresiana, St Petersburg, Florida, USA [23e]

Thelypteris limbosperma, Ireland [23e]

Thelypteris hydrophila crozier showing pneumatophores, Guadeloupe [23e]

Woodsia ilvensis, Nauvo, Finland [23f]

Distribution: This is a nearly cosmopolitan family, with the greatest diversity in (sub-) tropical montane regions. Phylogeny and evolution: Thelypteridoideae have the largest number of species in Aspleniaceae, in which they are deeply embedded. Having unusual hairs and placement of indusia, and sometimes more than two vascular bundles in the petiole, they are not a good morphological match with the remainder of Aspleniaceae, but these are derived characters and the subfamily clearly belongs here. They diverged from other Aspleniaceae c. 80 million years ago. Gener ic deli mit ation in Thelypteridoideae is still disputed, but we opt for a broad generic circumscription subsuming Pseudophegopteris into Phegopteris and

including Cyclosorus and Meniscium in an expanded Thelypteris (if segregated Meniscium has priority over Cyclosorus, causing nomenclatural havoc). Future studies may show that more genera need to be recognised, but with our current knowledge of the group, this reduction in generic names seems the most stable solution. Genera and species: Thelypteridoideae consist of three genera with c. 1,130 species: Macrothelypteris (11), Phegopteris (26) and Thelypteris (c. 1,100). Etymology: Thelypteris is derived from the Old Greek θελης (thelys), female or fruitful, and πτέρης (pteris), a fern, possibly referring to the superficial similarity to the lady fern (Athyrium filix-femina, Athyrioideae)

23f. ASPLENIACEAE SUBFAMILY WOODSIOIDEAE Cliff ferns These are terrestrial ferns that often grow on rocky substrates. Their roots are wiry and blackish as in all other Aspleniaceae. Their rhizomes are scaly, short-creeping or erect and usually not branched. The rhizome scales are lanceolate and not clathrate. Leaves are monomorphic and closely spaced and bear both scales and hairs. The two-stranded petioles are usually persistent on the rhizomes, but are sometimes articulate. Blades are once pinnate to twice pinnate-pinnatifid, usually broadest in the middle, with or without reduced pinnae at base, and apices are reduced to a non-conforming terminal segment. Veins are free and do not reach the margin, and vein endings are usually

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POLYPODIALES expanded, forming hydathodes. Sori are placed dorsally along veins on flat receptacles, and are round and indusiate. Indusia are basal and composed of a series of segments or are globose and sack-like. Sporangium stalks are two or three cells wide in the middle. Spores are monolete, brown and achlorophyllous and variously ornate. Gametophytes are green and heart-shaped.

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23g. ASPLENIACEAE SUBFAMILY ATHYRIOIDEAE Lady ferns

Etymology: Woodsia is named for English botanist Joseph Woods (1776–1864).

These are terrestrial or rarely aquatic ferns with blackish, wiry roots. Their rhizomes are short- to long-creeping, suberect or erect, sometimes branching and scaly. Rhizome scales are lanceolate and usually not clathrate. Leaves are green or tinged pink, red or variously coloured and are usually monomorphic or slightly dimorphic. They can have bulbils, hairs or scales. Petioles have usually two (rarely more) vascular bundles. Blades are simple or one to three times pinnate-pinnatifid, the apex usually pinnatifid and different than lateral pinnae. Veins are usually free, usually terminating before they reach the margins, and often forming hydathodes, sometimes the veins are anastomosing, but areoles formed by this reticulation lack included veinlets. Sori form on a flat receptacle. They are usually elongate, sometimes rounded, and are placed atop a vein or along one (asplenioid) or both sides of the vein (diplazioid), or in the axils of veins and the indusia merged in V- or J-shapes (athyrioid). Sori are rarely marginal; the indusia are usually laterally attached, vaulted or flat. Sporangia are stalked, the stalks two or three cells wide in the middle. Spores are

Athyrium otophorum, private garden, Kingston upon Thames, Surrey, UK [23g]

Athyrium asplenioides, sori, New York, USA [23g]

Distribution: These occur mostly in the Arctic and mountains of the Northern Hemisphere, in open rocky places, forests or arid areas, with one species (Woodsia montevidensis) extending along the Andes into temperate South America. Phylogeny and evolution: Woodsioideae form an isolated lineage in Aspleniaceae, and, in spite of their superficial resemblance to Cystopteridoideae, they are not the immediate relatives of this subfamily. They do occur in similar habitats, but resemblance is due to convergence. They are clearly divided into an Old World and a New World clade. Genera and species: Woodsioideae consist of the single genus Woodsia, with c. 35 species.

Diplazium proliferum, sori, Réunion [23g]

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monolete, achlorophyllous, plain or variously ornate. Gametophytes are green and heartshaped, glabrous or with glandular hairs on the wings. Distribution: This subfamily can be found worldwide, except for extremely arid regions and glaciated or permafrost areas. They are most diverse in the Asian subtropics. Phylogeny and evolution: These four genera have long been associated and were usually included with Cystopteris, Diplaziopsis and Woodsia in the family Woodsiaceae. That assembly was found to be polyphyletic with regards to other lineages in “eupolypods II” and were hence divided. Athyrioideae form a clade with Blechnoideae, which diverged c. 75 million years ago. Anisocampium is sometimes segregated from Athyrium, making the latter genus paraphyletic. Genera and species: Athyrioideae consist of four genera with c. 580 species: Athyrium (c. 216), Deparia (10) and Diplazium (c. 350). Etymology: Athyrium is derived from the Greek αθυρος (athyros), without a door, an ‘atrium’, in reference to the chamber-like, open sorus.

Diplazium sibiricum, Sweden [23g]

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Blechnum ryanii, Guadeloupe [23h]

23h. ASPLENIACEAE SUBFAMILY BLECHNOIDEAE Hard ferns These terrestrial and climbing ferns have blackish wiry roots that emerge from erect, sometimes trunk-like or short- to longcreeping, sometimes scandent, branched or unbranched, sometimes stoloniferous, scaly rhizomes. Rhizome scales are lanceolate to linear, not clathrate. Unfolding leaves are usually colourful, often orange, red or pinkish (green in Onoclea), sometimes covered in mucilage, and are monomorphic or strongly dimorphic, bulbiferous or not and usually scaly and sometimes hairy. Petioles have two large vascular bundles and smaller bundles on the upper side of the petiole; in Onoclea these are starch-filled at the base and sometimes persistent for many years forming a protective sheath. The blades are pinnatifid or one to two times pinnate-pinnatifid, usually once pinnate, the rachis vining in Salpichlaena, the base with or without reduced pinnae, the apical pinna conform or not, pinnae in some species with conspicuous aerophores. Veins are free, simple or forked and only occasionally anastomosing along midveins or fully anastomosing, the areoles in this way lacking included veinlets, the free veins reach the margin or terminate

Woodwardia radicans, private garden, England, UK [23h]

before it. Sori are formed on a flat or raised conical receptacle, elongate along one side of veins parallel to the midvein or round and terminating a vein. The sori are usually indusiate, but are rarely acrostichoid and exindusiate; the indusia are flat and lateral, with the opening facing the midvein, usually elongate, sometimes triangular. Sporangia are stalked, the stalk one to three cells wide. Spores are monolete, not or occasionally chlorophyllous but usually brown, smooth or variously ornate. Gametophytes green, heartshaped, sometimes with glandular hairs on the wings. Distribution: Blechnoideae are cosmopolitan, except for arid, polar and high alpine regions. Phylogeny and evolution: Blechnoideae comprise two subclades, Onoclea s.l. (sometimes treated as Onocleaceae) and Woodwardia+Blechnum s.l. Despite some differences in fertile structures these are closely related and share many morphological characters. Onoclea diverged from the Woodwardia+Blechnum clade at the end of the Cretaceous, some 70 million years ago. The former Onocleaceae include five species that were traditionally divided into two genera, which are at their extremes easily distin-

guishable, the ostrich ferns (often placed in Matteuccia) and the sensitive fern (Onoclea sensibilis), but the other three species are intermediate and have been placed into two additional not very distinctive genera, each genus thus representing one or two species. These species are better united under the single genus Onoclea. The remaining clade has a deep split corresponding roughly with the two Hemispheres: Woodwardia is the first branching, exclusively found on the northern continents. The second clade includes Blechnum, with its segregates Brainea, Doodia, Pteridoblechnum and Sadleria all embedded in it, and Stenochlaena as sister to this clade, but with Salpichlaena and Telmatoblechnum placed with Stenochlaena rather than with core Blechnum. Further phylogenetic research is needed to study character evolution in the morphologically diverse genus Blechnum. Genera and species: Blechnoideae include six (or more) genera with c. 290 species: Blechnum (c. 250), Onoclea (5), Salpichlaena (2), Stenochlaena (c. 15), Telmatoblechnum (2) and Woodwardia (c. 20). Etymology: Blechnum is derived from βλήχνον (blechnon), an ancient Greek word for fern. Plants of the World

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Onoclea struthiopteris, Finland [23h]

Blechnum (Sadleria) cyatheoides, Royal Botanic Gardens, Kew, UK [23h]

Stenochlaena palustris, Singapore [23h]

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Blechnum spicant, Twickel, the Netherlands [23h]

Blechnum paschale, Easter Island [23h]

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24. POLYPODIACEAE Polypody family

sometimes ribbon-like or elongate and straplike to filamentous, glabrous or hairy. Distribution: Polypodiaceae occur worldwide. This is the largest family of ferns, and they occupy most habitats. They are absent from polar, high alpine and arid areas.

This family consists of terrestrial, epiphytic and epilithic, sometimes climbing ferns with erect or long- or short-creeping rhizomes that branch or not. The rhizomes are sometimes viny or climbing, sometimes have stolons or tubers, are often fleshy and are usually scaly. Rhizome scales are clathrate or opaque, hairy or glabrous, basally attached or peltate. Their leaves are variable, monomorphic or dimorphic, simple or one to four times pinnate-pinnatifid, short- or long-petiolate, the petioles cleanly abscising or persistent. Petioles have two to several vascular bundles that are usually arranged in a half-circle, a ring or a U-shape in cross-section, sometimes swollen at the base or with an articulation. Blades can be glabrous, hairy, scaly or glandular, sometimes with proliferous buds, or they can be pedate, flabellate or forked. Veins are free, simple, forked or pinnate, or variably anastomosing or reticulate, areoles, when present, having included veinlets or not, sometimes with thickened endings (hydathodes) before reaching the margin. Sori are superficial or impressed, discrete and round or confluent and acrostichoid (covering all or part of the fertile leaf lamina below), rarely linear and following the veins, indusiate or exindusiate. Indusia, when present are usually centrally attached and reniform, but can be variably laterally attached, reniform or lunate, oblique or round or elongate and linear. Sporangia are stalked, the stalk with up to three cells, the capsule with a vertical annulus. Spores are monolete, ellipsoidal to globose and alate, cristate, verrucate, tuberculate, rugose or echinate, chlorophyllous and green or achlorophyllous and brown. Gametophytes are green and usually heart-shaped, but

Phylogeny and evolution: Polypodiaceae are the broad family formerly recognised as several families placed in ‘eupolypods I’. They include the majority of extant fern diversity. They are especially diverse in the understory of rainforests and as epiphytes on rainforest trees, and they diversified in response to the development of angiosperm rainforests during the Cretaceous. Subfamilies Didymochlaenoideae, Hypodematioideae, Dryopteridoideae, Lomariopsidoideae and Tectarioideae form a grade to the almost exclusively epiphytic or epilithic Oleandroideae, Davallioideae and Polypodioideae. However, epiphytic species also occur in other subfamilies, suggesting that epiphytism has evolved several times independently in Polypodiaceae. Genera and species: Polypodiaceae consist of c. 76 genera with c. 4,070 species, with the largest genera being Dryopteris (c. 400), Elaphoglossum (c. 750), Grammitis (c. 700), Polystichum (c. 370) and Tectaria (c. 250). The family is divided into eight subfamilies (Didymochlaenoideae, Davallioideae, Dryopteridoideae, Hypodematioideae, Lomariopsidoideae, Oleandroideae, Polypodioideae and Tectarioideae), which are sometimes treated as separate families and are therefore treated and discussed separately below. Uses: The tender young leaves of Arthromeris wallichiana are cooked as a vegetable in Nepal. Leaves of some Pyrrosia species are eaten as a vegetable in China. The young fronds of many species of Dryopteris and Polystichum are cooked and eaten in Asia, but caution has to be taken because they are mildly toxic, even when boiled. Rhizomes of Dryopteris expansa and Polystichum munitum were eaten by Native Americans and reportedly taste like sweet potatoes, but

they were only eaten when little else was available. Rhizomes of several Polypodium species, especially P. glycyrrhiza and P. vulgare, contain ostadin, a steroidal saponin that has a sweet taste. They are licorice-flavoured and were commonly chewed by many native North American tribes as an appetiser, especially by children who would not eat. In the past, common polypody, P. vulgare, was employed to flavour tobacco. Rumohra adiantiformis is cultivated on a large scale for the cut-flower industry as the fern leaf in many bouquets of cut flowers. The fibre of Nephrolepis hirsutula is sometimes used to manufacture hats, mats and baskets. Boston fern (Nephrolepis exaltata ‘Bostoniensis’) is a common house plant. This and several other Nephrolepis are frequently planted as ornamentals in the tropics, where they sometimes naturalise. Didymochlaena truncatula and Davallia tyermannii are also frequently grown as houseplants, and many others are used as garden ornamentals. Gametophytes of Lomariopsis lineata are sometimes sold as peculiar aquarium plants, under the German name ‘Susswassertang’. Etymology: Polypodium is derived from the Greek πολύς (polys), many, and ποδιών (podion), feet; in reference to the ‘manyfooted’ branching rhizomes.

24a. POLYPODIACEAE SUBFAMILY DIDYMOCHLAENOIDEAE Mahogany ferns These large terrestrial ferns have erect, thick, scaly rhizomes, with persistent petiole bases. The scales are long, narrow and almost hair-like. Leaves are bipinnate and long-petiolate; petioles have several vascular bundles arranged in a half-circle. Pinnae are articulate to the rachis and more or less similar in size and shape, somewhat rectangular in outline. Veins are free, forked, with endings thickened before reaching the margin. Sori terminate a vein and are often somewhat sunken in the blade, forming bumps on the upper side of the leaf. Indusia are elongate, centrally attached along a line and opening on either

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POLYPODIALES side. Sporangia are long-stalked. Spores are monolete, ellipsoidal to globose, tuberculate and echinate. Gametophytes are heart-shaped and glabrous. Distribution: The single species is found throughout the tropics, often on wet clayey substrates near streams in forests. There is possibly an additional species in Madagascar, but this is yet to be confirmed. Phylogeny and evolution: Didymochlaenoideae are generally treated as related to Dryopteridoideae, but inclusion there would render the latter paraphyletic. In some studies Didymochlaena is sister to Hypodematioideae, but including it in that clade makes Hypodematioideae difficult to define morphologically. Didymochlaena is the first branching clade in Polypodiaceae. Didymochlaenoideae are distinguished from Hypodematioideae in having erect rhizomes, hair-like rhizome scales, bipinnate leaves, with all pinnae of the same shape, not gradually narrowing and confluent towards the apex, veins with thickened ends terminating before the margin, sori terminating a vein, with long and narrow elongate indusia attached along a central line, opening towards the sides and having echinate spores. Didymochlaena truncatula, Brazil [24a]

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Genera and species: This subfamily consists of a single genus Didymochlaena, with one (D. truncatula) or maybe two species. Etymology: Didymochlaena is derived from Greek δίδυμος (didymos), twin, and χλαινα (chlaina), a cloak, in reference to indusia that open on two sides.

24b. POLYPODIACEAE SUBFAMILY HYPODEMATIOIDEAE Bigfoot ferns These are usually terrestrial or epilithic ferns with long- or short-creeping, thick, sometimes fleshy, scaly rhizomes. Rhizome scales are sometimes hairy. Leaves are two to four times pinnate-pinnatifid and long-petiolate. Petioles are often swollen at the base and have two large and several smaller vascular bundles organised in a U-shape in cross-section. Blades and pinnae are gradually reduced and confluent towards the terminal segments. Veins are free, simple, forked or pinnate and end at the margin. Sori are impressed and indusiate, the indusia reniform to nearly round and basally (or also slightly laterally) attached, opening towards the margin, glabrous or hairy. Sporangia are longstalked with stalks one to three cells wide and have a vertical annulus. Spores are monolete, ellipsoidal, coarsely verrucate or tuberculate.

Gametophytes are green and heart-shaped and glandular hairy on the margins. Distribution: They occur mainly in tropical Asia, east to Polynesia, with one species extending into tropical East Africa and Madagascar. Phylogeny and evolution: This subfamily is often associated with Dryopteridoideae, but merging the two would render the latter polyphyletic. The two genera that compose Hypodematioideae have been variously treated in the past: Hypodematium was considered to be associated with the athyrioid ferns, and Leucostegia was previously placed in Davalliaceae. They are, however, each other’s closest relatives. Didymochlaena was found as sister to these two genera, but usually without strong support. It is possibly not directly related. Didymochlaena is here treated in a separate subfamily. Genera and species: Hypodematioideae consist of two genera and 16 species: Hypodematium (13) and Leucostegia (3). Etymology: Hypodematium is derived from the Greek υπό (ypo), under, and δεματίων (demation), a bundle, referring to the hairs on the sori.

Arachniodes aristata, sori, Tahiti [24c]

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Polystichum setiferum, crozier, private garden, Hengelo, the Netherlands [24c]

Elaphoglossum feei, Guadeloupe [24c]

Polybotrya osmundacea, sori, Brazil [24c]

Elaphoglossum crinitum, Botanical Garden, Berlin-Dahlem, Germany [24c]

Polystichum aculeatum, Yorkshire, UK [24c]

Mickelia nicotianifolia, Guadeloupe [24c]

Elaphoglossum peltatum, Mary Selby Botanical Garden, Sarasota, Florida, USA [24c]

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24c. POLYPODIACEAE SUBFAMILY DRYOPTERIDOIDEAE Buckler ferns This subfamily consists of terrestrial and epiphytic ferns. Their rhizomes are creeping or erect, sometimes vining and scaly. Rhizome scales are not clathrate. Leaves are monomorphic or dimorphic, simple or one to four times pinnate-pinnatifid, and petioles have many vascular bundles arranged in a circle. Blades are glabrous, hairy, scaly or glandular. Veins are free, variably anastomosing or reticulate, forking or pinnate, and areoles, when present, can have included veinlets or not. Sori are usually rounded with reniform or round, peltate indusia, or the indusia are absent. In tribe Elaphoglosseae the sori are acrostichoid, i.e. covering the entire lamina of the fertile leaf, without indusia. Sporangia are shortstalked, the stalks with three rows of cells. Spores are monolete, reniform and winged. Gametophytes are green and heart-shaped. Distribution: This is an almost cosmopolitan subfamily, absent from arid and frozen areas. Several species are colonisers on oceanic islands. The subfamily is diverse and includes terrestrial plants, climbers and epiphytes. Phylogeny and evolution: In the traditional sense, this subfamily included a number of genera that have been demonstrated not to

Cyclopeltis presliana, sori, Singapore Botanic Garden [24d]

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belong here, but when Cyclopeltis, Didymochlaena, Dracoglossum, Lomariopsis, Nephrolepis and the genera now in Tectarioideae are excluded and the remaining genera previously in Lomariopsidaceae (e.g. Bolbitis, Elaphoglossum) are included they form a clade that is difficult to define morphologically. Dryopteridoideae are divided into two tribes: Elaphoglosseae, with acrostichoid fertile laminae, and Dryopterideae, the true buckler ferns. Mickelia nicotianifolia was found to have the largest known genome of any leptosporangiate fern. Genera and species: Dryopteridoideae consist of c. 27 genera with c. 2,000 species: Arachniodes (c. 110), Arthrobotrya (3), Bolbitis (c. 60), Ctenitis (c. 120), Cyclodium (10), Cyrtomidictyum (4), Dryopolystichum (1), Dryopteris (c. 400), Elaphoglossum (c. 600–800), Lastreopsis (41), Lomagramma (19), Maxonia (1), Megalastrum (c. 40), Mickelia (10), Olfersia (2), Parapolystichum (5), Phanerophlebia (9), Pleocnemia (23), Polybotrya (35), Polystichopsis (30), Polystichum (c. 370), Pseudotectaria (8), Rumohra (1–8), Stigmatopteris (28) and Teratophyllum (12). Etymology: Dryopteris is derived from the Greek δρυς (drys), oak and πτέρης (pteris), fern, describing the shape of the pinnae of some species.

Nephrolepis biserrata, Brazil [24d]

24d. POLYPODIACEAE SUBFAMILY LOMARIOPSIDOIDEAE Sword ferns These are terrestrial, epilithic and epiphytic ferns. Their rhizomes are variable, often creeping and scrambling over rocks or climbing up trees, sometimes erect, stout and short, in Nephrolepis often with stolons (runners forming buds from which new plants grow), sometimes also with scaly tubers. Leaves are monomorphic or dimorphic, and petioles have two large vascular bundles and several smaller ones arranged in a U-shape. Blades are simple and proliferous at the apex (in Dracoglossum) or once pinnate and not proliferous (bipinnate in some cultivars of Nephrolepis). Pinnae, when present, are articulate to the rachis. Veins are free, parallel, simple and forked or pinnate, sometimes meniscioid-reticulate with included veinlets and in Nephrolepis ending before the margin in thickened hydathodes. Sori are discrete, orbicular, with peltate round or reniform indusia or exindusiate, placed in one to four rows on each side of the costa, in one species of Nephrolepis linear and terminating many veins, or sporangia acrostichoid. Spores are bilateral, monolete, ellipsoid or globular, variously winged or ornamented. Gametophytes are green and heart-shaped or ribbon-like, glabrous or long-hairy.

Dracoglossum plantagineum, Guadeloupe [24d]

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FERNS

Tectaria griffithii, Malaysia [24e]

Hypoderris brownii, Puerto Rico, USA [24e]

Distribution: This is a pantropical subfamily of understory vegetation in rainforests and on rocks along small streams.

Ancient Greek λωμα (loma), an edge or border, in reference to the sori that run in parallel with the leaf margin.

Phylogeny and evolution: Lomariopsidoideae previously included Bolbitis, Elaphoglossum, Lomagramma and Teratophyllum, genera that are now known to belong to Dryopteridoideae. Dracoglossum (previously treated as a species of Tectaria) is most closely related to Lomariopsis, a relationship that has only recently been discovered. Nephrolepis is variously included or excluded, but it shares the articulate leaflets, which are diagnostic for this group. Nephrolepis was traditionally placed in Davallioideae, a result not supported by molecular and morphological analyses. Nephrolepis evolved in the Eocene in the forests of the Laurasian tropical belt, from where two main lineages have dispersed and became isolated, one in the Neotropics and the other in AsiaAustralasia, from where they spread to other tropical regions.

24e. POLYPODIACEAE SUBFAMILY TECTARIOIDEAE Button ferns

Genera and species: Lomariopsidoideae consist of four genera with c. 75 species: Cyclopeltis (7), Dracoglossum (2), Lomariopsis (c. 45) and Nephrolepis (20). Etymology: Lomariopsis is named for its resemblance to Lomaria, a synonym of Blechnum. Lomaria is derived from the

Button ferns are terrestrial with creeping to erect, scaly rhizomes. Their rhizome scales are much longer than wide and not clathrate. Leaves are monomorphic or dimorphic, and petioles have several vascular bundles organised in a ring or semi-circle. Petioles are sometimes articulate (Arthropteris). Blades are simple or pinnate to bipinnate-pinnatifid, sometimes decompound, usually with stubby jointed hairs, and midribs bear proliferous buds in some species. Veins are free or (more often) anastomosing or fully reticulate, usually with included veinlets. Sori are of variable shape and position, usually dorsal or terminal on the veins, indusiate or not, round, oval or (rarely) linear and opening towards the margin, rarely following veins and covering the entire lamina eventually. Indusia, when present, are round, reniform or lunate, sometimes oblique. Sporangia are stalked, and spores are ellipsoidal with wing-like folds or are variously ornamented. Gametophytes are green and more or less heart-shaped.

Distribution: Tectarioideae are pantropical, extending into the subtropics in Asia. Phylogeny and evolution: A number of genera traditionally associated with Tectaria (e.g. Ctenitis, Dryopsis, Lastreopsis, Lomariopsis, Pleocnemia, Pseudotectaria etc.) are not closely related and do not belong in Tectarioideae. Arthropteris is not closely related to Oleandra or Nephrolepis as previously assumed, but is sister to the rest of Tectarioideae. This clade of terrestrial ferns is sister to the predominantly epilithic and epiphytic Oleandroideae, Davallioideae and Polypodioideae. Genera and species: Tectarioideae consist of seven genera with c. 315 species: Aenigmopteris (5), Arthropteris (22), Draconopteris (2), Hypoderris (3), Pteridrys (10), Tectaria (c. 250) and Triplophyllum (23). Etymology: The derivation of Tectaria is obscure, but it is possibly derived from the Greek τεκτα (tekta), mason, because the venation pattern of some species resembles masonry.

24f. POLYPODIACEAE SUBFAMILY OLEANDROIDEAE Stilt ferns These scandent epilithic or epiphytic, rarely terrestrial ferns have long-creeping, frequently

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Oleandra annetii, Seychelles [24f]

branching rhizomes covered with peltate rhizome scales. Leaves are simple, articulate to a short knob-like outgrowth of the rhizome. Petioles have two larger and several smaller bundles organised in a semi-circle. Blades are simple, rounded at the base and acute to acuminate at the apex. Veins are parallel, forked, free or occasionally anastomosing, ending in thickened hydathodes near the margins. Sori are round and form singly on upper vein branches, rarely nearly terminally on the veins, indusiate. Indusia are reniform to nearly round and attached by a broad sinus. Sporangia have stalks that are long and three cells wide, and they emerge mixed with hairs. Spores are monolete, ellipsoid, with prominent wing-like folds and various ornamentations. Gametophytes are green, heart-shaped and often hairy. Distribution: This clade is pantropical, with only a few representatives in the Americas and Africa. Phylogeny and evolution: The subfamily is closely related to Polypodioideae, with which they share several characters. Traditionally Arthropteris was also included, but this genus is now placed in Tectarioideae. Genera and species: Oleandroideae consist of the single genus Oleandra with c. 15–20 species, mostly in the Asian tropics.

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Oleandra sibbaldii, sori, Tahiti [24f]

Etymology: Oleandra is derived from Nerium oleander (Apocynaceae) because of the superficial resemblance of the type species, Oleandra neriiformis, to leaves of oleander.

Distribution: Davallioideae occur throughout the Palaeotropics extending into the Asian subtropics, with a disjunct population in the Canary Islands. They are most diverse in tropical Asia.

24g. POLYPODIACEAE SUBFAMILY DAVALLIOIDEAE Hare’s-foot ferns

Phylogeny and evolution: The subfamily in the strict sense (excluding Leucostegia, now in Hypodematioideae, and Gymnogrammitis, now part of Selliguea, Polypodioideae) has only two genera. There were several other genera traditionally accepted in this group, but they have been found to be embedded in one or other of the two genera now recognised.

These are epiphytic and epilithic ferns with long-creeping, branching, densely scaly rhizomes. Rhizome scales are peltate and cover the entire rhizome, making it look like the foot of an animal. Leaves are usually monomorphic, rarely dimorphic, and petioles have two larger and three smaller vascular bundles. Blades are simple or one to three times pinnate-pinnatifid and are hairy or glabrous when mature. Veins are free, simple, forked or pinnate, terminating before reaching the margin. Sori are formed marginally or away from the margin but always terminating veins. They are triangular or round, with a cup-shaped, reniform or lunate indusium, rarely forming a confluent marginal sorus. Sporangia are long-stalked, the stalk threecelled and the capsule with a vertical annulus. Spores are monolete, ellipsoid, winged, cristate or variously ornamented. Gametophytes are green, heart-shaped with short, occasionally branched hairs.

Genera and species: Davallioideae consist of two genera with about 45 species: Davallia (c. 30) and Davallodes (c. 15). Etymology: Davallia is named for English botanist Edmond Davall (1763–1798), who bequeathed his herbarium to his friend James E. Smith, who named this genus for him.

24h. POLYPODIACEAE SUBFAMILY POLYPODIOIDEAE Polypodies These mostly epiphytic and epilithic ferns are rarely found terrestrially (and then mostly in temperate regions). Rhizomes

POLYPODIALES

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Davallia mariesii, Royal Botanic Gardens, Kew, UK [24g]

Pyrrosia lingua, Jardin des Plantes, Paris, France [24h]

Microsorum scolopendrium, Réunion [24h]

Davallia solida, Tahiti [24g]

Platycerium coronarium, Singapore [24h]

Pleopeltis polypodioides, St Petersburg, Florida, USA [24h]

Loxogramme lanceolata, Kenya [24h]

Grammitis serrulata, Guadeloupe [24h]

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POLYPODIALES are creeping, often long and branching, scaly or (rarely) glabrous, sometimes short and stubby or suberect. Rhizome scales are clathrate or opaque. Most species (except Grammitis) have leaves that leave a clean scar or knob after falling off. Leaves are monomorphic or dimorphic with petioles that have a variable number of vascular bundles arranged in a U-shape. Blades are simple, pinnatifid or once pinnate, rarely more divided, sometimes pedate, flabellate or forked, glabrous or with hairs or scales. Veins are free, anastomosing or reticulate, sometimes the areoles with included veinlets. Sori are formed underneath the lamina or (rarely) marginally, usually rounded or elliptic, sometimes sunken in the lamina or conf luent to completely acrostichoid, never indusiate, but sori sometimes covered with scales. Sporangia are short or longstalked, with three cells; the capsule has a vertical, interrupted annulus. Spores are monolete, bean-shaped, white, yellow, green or brown, or trilete, thin-walled, globosetetrahedral and chlorophyllous, and are variously ornamented. Gametophytes are green and heart-shaped, broader than long, often with a prominent midrib and often with

Polypodium vulgare, Turku, Finland [24h]

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various hair-types, or the gametophytes are strap-like to filamentous, sometimes with proliferating buds. Distribution: Polypodioideae are a cosmopolitan subfamily of evergreen ferns with the greatest diversity in the tropics. Like most ferns, they are absent from arid or frozen areas. Phylogeny and evolution: Polypodioideae are sister to Davallioideae and Oleandroideae, with which they share their preference for growing epiphytically and epilithically. Polypodioideae differ from those subfamilies in their lack of an indusium. Polypodioideae are divided into five tribes: Loxogrammeae including the peculiar Loxogramme; Drynarieae with Drynaria and Selliguea; Platycerieae, with Pyrrosia and Platycerium (stag horn ferns); Microsoreae, with Microsorum, Goniophlebium and relatives; and Polypodieae, including the true polypods and Grammitis. Grammitis differs in having green, trilete spores and persistent petiole bases and often filiform gametophytes. Previously these ferns were treated in a separate family

Grammitidaceae with a great number of genera, but they are better treated as the single genus Grammitis in subfamily Polypodioideae. The genera in Polypodioideae are still in need of further study and recircumscription. Genera and species: Polypodioideae include about 29 genera, sometimes more depending on taxonomic preference, with c. 1,600 species: Campyloneurum (55), Dendroconche (2), Dictymia (2), Drynaria (27), Goniophlebium (27), Grammitis (c. 700), Lecanopteris (13), Lemmaphyllum (13), Lepidomicrosorium (27), Lepisorus (c. 90), Leptochilus (c. 50), Loxogramme (44), Microgramma (25), Microsorum (c. 40), Neocheiropteris (c. 9), Paragramma (5), Pecluma (30), Phlebodium (4), Phymatosorus (31), Platycerium (16), Pleopeltis (c. 35), Pleurosoriopsis (1), Podosorus (1), Polypodium (c. 70), Pyrrosia (c. 70), Selliguea (c. 145), Serpocaulon (43), Thylacopteris (2) and Tricholepidium (14) Etymology: Polypodium is derived from the Greek πολύς (polys), many, and ποδιών (podion), feet, refering branching, creeping rhizomes.

Grammitis nanodes, Kenya [24h]

GYMNOSPERMS The first seed plants were gymnospermous in the sense that ovules did not develop inside an ovary, but rather were ‘naked’ on the sporophylls (fertile leaves). The ovules develop on modified leaves or scales, often fused into coneshaped structures. The earliest seed-like fossils are known from Upper Devonian deposits, c. 385–359 million years ago. All seed plants are heterosporous, i.e. they produce two kinds of spores, microspores (pollen grains) and megaspores (ovules in the ovaries). Hence, it is assumed that the ancestors of seed plants must have been heterosporous, but heterospory in seedless vascular plants (ferns and lycopods) is mostly restricted to aquatic lineages, which have evolved heterospory as a secondary trait. The fossil record of gymnosperms includes many distinctive taxa that do not belong to the four modern lineages, including seed-bearing trees with a fern-like vegetative morphology, the seed ferns or pteridosperms. It has been suggested that angiosperms are derived from a larger group of gymnosperms that were much more heterogeneous than extant gymnosperms and undoubtedly were not a single clade. However, all extant gymnosperms are sister to the angiosperms. Early molecular results (e.g. Hasebe et al. 1992) were the first to demonstrate that the extant group was a clade, but most workers at the time considered that the sparse sampling of the early DNA studies made the results somewhat unconvincing. However, all major DNA studies to date have demonstrated this relationship, so it has slowly gained acceptance. Prior to DNA studies, it was thought likely that each of the four extant clades would be related to different groups of sporebearing taxa, thus making the concept of “gymnosperm” similar to that of “dicot”

in referring to a stage of evolution (a grade) rather than a monophyletic group. As indicated above, the stillprevailing notion is that gymnosperms in the broadest sense, including fossil taxa, are not monophyletic and thus the term refers to a grade. However, use of “gymnosperm” is nonetheless made acceptable (unlike “dicot”) by the fact that all extant taxa form a clade. Given that this was an unexpected result, how the other (all fossil) taxa are related to these is more speculative than previously thought. Therefore, until such time as definitive results have been obtained, we advocate continued use of “gymnosperm” (whereas the term “dicot” must be abandoned). Seeds are thought to have evolved to keep plant embryos from drying out. The development of seeds allowed plants to be less dependent on water for fertilisation, and enclosure of female gametes in an ovule increased this success rate. Formation of seeds also allowed storage of nutrients that were packed together with the embryo. These seeds could germinate in periodically more hospitable environments. Early seed plants (gymnosperms) still depended on the wind to bring pollen to the ovaries, but with the development of flowers, animals became involved in this process, although some modern gymnosperms are known to be insect-pollinated as well. Insects, especially beetles, were attracted by flowers to eat from fleshy structures (tepals) and pollen, but soon more specialised associations between flowers and their pollinators developed, resulting in the enormous diversity of flowers and pollinators we see today. Diversif ication of angiosperms resulted in a loss of gymnosperm diversity. At present the slightly more than 1,000 species of extant gymnosperms are

arranged in 12 families. Even though the group was more diverse in the fossil record than today, they occur throughout the world, forming dominant stands in boreal forests, subtropical swamps, tropical mountains, cloud and rain forests, coastal habitats and semi-deserts. Against the background of their paucity of species numbers, they exhibit a great deal of heterogeneity and specialisation, including at least one (half-)parasite (Parasitaxus) and species adapted to unusual substrates (e.g. ultramafic rocks in New Caledonia) and conditions (e.g. waterlogged conditions, as for Taxodium and Retrophyllum), arid regions and annually deep-frozen boreal forests. The general manner in which they are described leads one to conclude that they are a relict group, which has been collectively excluded from many habitats by the angiosperms, but this is misleading and ignores the fact that, like most groups of ferns and angiosperms, there has been a recent radiation of species adapted to modern environmental conditions. In fact, in the more archaic cycads, nearly all extant species and genera are the products of a radiation that took place in the past 20 million years. Although they clearly exhibit ancient morphologies, the modern species are not “living fossils”, except perhaps for Ginkgo biloba, the only extant member of its clade with a long fossil history. In terms of their inter-relationships, cycads are sister to the rest, followed successively by ginkgo and the conifers (the term conifer is often used for the pines and their relatives, but cones are also found in cycads), in which “gnetoids” (Gnetaceae, Ephedraceae and Welwitschiaceae), formerly thought to be transition groups between gymnosperms and angiosperms, are collectively sister to Pinaceae (see results of Chaw et al.

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Here we use the classif ication of Chase & Reveal (2009), who did not use a formal name for the gymnosperms, stating that, given the speculative nature of the hypothesised relationships of the fossil gymnosperms and their still-likely grade status, such formal taxonomic recognition would be unwise and premature. Each of the four major groups was given the rank of subclass, although it must now be considered that Gnetidae should now be treated as three orders in Pinidae. The angiosperms were given equal rank to each of these, as the single subclass Magnoliidae. As Chase & Reveal (2009) pointed out, past classifications that recognised the angiosperms as several subclasses in a single class (such as the system of Cronquist) forced other members of the land flora and the various groups of aquatic plants (“algae”, not a monophyletic concept) to be pushed into ever higher ranks, leaving “no room at the top”.

General references Byng JW. 2015. The gymnosperms handbook. A practical guide to extant families and genera of the world. Plant Gateway, Hertfort. Chase MW, Reveal JL. 2009. A phylogenetic classification of the land plants to accompany APG III. Botanical Journal of the Linnean Society 161: 122–127. Chaw SM, Parkinson CL, Cheng Y, Vincent TM, Palmer JD. 2000. Seed plant phylogeny inferred from all three plant genomes: monophyly of extant gymnosperms and origin of Gnetales from conifers. Proceedings of the National Academy of Sciences (USA) 97: 4086–4091. Chaw SM, Zharkikh A, Sung HM, Lau TC, Li WH. 1997. Molecular phylogeny of extant gymnosperms and seed plant evolution: analysis of nuclear 18S rRNA sequences. Molecular Biology and Evolution 14: 56–68. Christenhusz MJM, Reveal JL, Farjon A, Gardner MF, Mill RR, Chase MW. 2011. A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70. Eckenwalder JE. 2009. Conifers of the world. Timber Press, Portland. Farjon A. 2010. A handbook of the world’s conifers. Brill, Leiden. Foster AS, Gifford EM. 1974. Comparative morphology of vascular plants. Freeman, San Francisco. Hasebe M, Kofuji R, Ito M, Kato M, Iwatsuki K, Veda K. 1992. Phylogeny of gymnosperms inferred from rbcL gene sequences. Botanical Magazine (Tokyo) 105: 673–679.

2000, a study that has created a great deal of controversy). Other studies using various methods of analysis and/or downweighting of particular categories of DNA positions that change frequently have produced other topologies, but the one most favoured and most commonly obtained is with the gnetoids as sister to Pinaceae, the so-called “gnepine hypothesis”. Given that the previously most favoured hypothesis for gnetoids was as the sister group to the angiosperms, the “gnepine hypothesis” is revolutionary and will still require some years of additional investigation before it becomes more widely accepted. Relative to what we know of these taxa morphologically, such a relationship is bizarre and calls into question relationships hypothesised for fossil groups such as Bennettitales (Cycadeoidales) and “seed ferns” (e.g. Archaeopteris) on morphological g rounds. Sim i larly unex pected results of DNA studies are those for Psilotum and Equisetum, which were long held to be only distantly (and non-exclusively) related to ferns, but they have been clearly shown to be part of the fern clade, calling into question relationships proposed for the former to the rhyniophytes (the earliest vascular plants known) and the latter to Calamites and related sphenophylls (Foster & Gifford 1974).

Etymology: The word gymnosperm is derived from γυμνός (gymnos), naked, and σπέρμα (sperma), seed, referring to the seeds that are formed on the sporophylls without a covering. Part of the gymnosperms are called conifers, from κώνος (conos), a cone, and φέρουν ( feroun), to bear, referring to the cones in which seeds and pollen are formed.

The tallest trees in the world are a gymnosperm, Sequoia sempervirens, Muir Woods, California, USA

Maarten Christenhusz holding cones of Pinus coulteri, one of the largest cones of any pine, Rancho Habitat dominated by a gymnosperm, Pinus Santa Ana Botanical Garden, California, USA palustris, Green Swamp, North Carolina, USA

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CYCADALES

GYMNOSPERMS

CYCADALES Cycads are an ancient group that is sister to all other living gymnosperms. Fossil cycads are known from the Lower Permian of China, c. 275 million years ago, with their greatest diversity during the Jurassic/Cretaceous periods. It is thought that cycads arose from an extinct Palaeozoic lineage of seed ferns, but this hypothesis needs re-examination in the light of the DNA-based assessments of their relationships. Cycads radiated and dispersed during the Permian and early Mesozoic, leading to the Jurassic sometimes being referred to as the ‘age of cycads’, during which bennettites (cycadeoids), cycads, ginkgos and conifers dominated the vegetation of the world. The extant families (Cycadaceae and Zamiaceae) do not have such an extremely long fossil history, but most extant genera can be recognised from early Tertiary deposits (c. 50–60 million years old). Fossil genera of Cycadaceae date back to the Permian, but the extant genera of Zamiaceae evolved since the end of the Cretaceous, thus rejecting the role of dinosaurs in producing modern cycad diversity and the designation of any extant species as a living fossil.

25. CYCADACEAE Sago family

These are unisexual plants with a subterranean or emergent trunk clothed with persistent leaf bases and new leaves formed in flushes. Young parts of the plants are hairy. The leaves are pinnate and not coiled when emerging, but the leaflets emerge in a circinnate way (coiled

Cycas revoluta, female cone, California, USA [25]

with the tip in the centre, like watch springs) and are not articulate to the midrib. The lower leaflets are usually reduced to paired rigid thorns. Male sporophylls (fertile leaves) are arranged in a cone. Female sporophylls are free, not forming cones, consisting of a linear stalk and an expanded apical lobe, usually bearing more than two (up to eight) ovules. Sperm is free-swimming. Seeds have a hard inner and a fleshy outer seed coat. Distribution: This family occurs in tropical Asia, Melanesia, Fiji, Tonga, Australia, Madagascar and coastal tropical East Africa. Pollen is eaten by beetle larvae, and pollination is generally via insects, although some wind pollination may also occur. Most species have a brightly coloured sarcotesta

Cycas seemannii, New Caledonia [25]

(seed coat) and are dispersed by animals, but some coastal species like Cycas rumphii and C. seemannii have floating seeds that are dispersed by sea currents; these species have a wider distribution as would be expected for this mode of dispersal. Phylogeny and evolution: Cycas is the oldest extant genus of cycads, with fossils dating back to the Permian of China and the Eocene of Japan, and molecular estimates of its divergence from the lineage leading to all other extant cycads date to the early Jurassic. In this sense, at least Cycas is a living fossil, although this has been claimed for Cycadales as a whole. Unexpectedly, a molecular clock study demonstrated that none of the species in Cycas and the other cycad genera is older than five million years.

Cycas revoluta, male cone, Irvine, California, USA [25]

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CYCADALES

GYMNOSPERMS

Microcycas calocoma, female cone, Montgomery Botanical Center, Miami, Florida, USA [26]

Dioon edule, female cone, St Petersburg, Florida, USA [26]

Ceratozamia hildae, Matthaei Botanical Garden, Ann Arbor, Michigan, USA [26]

Stangeria eriopus, male cone, Helsinki Botanical Garden, Finland [26]

Lepidozamia peroffskyana, female cones, Royal Botanic Gardens, Sydney, Australia [26]

Encephalartos woodii, male cones, Royal Botanic Gardens, Kew, UK [26]

Genera and species: Cycadaceae include the single genus Cycas, which has c. 91–107 species.

26. ZAMIACEAE

circinnate vernation). Male and female sporophylls (fertile leaves) are fused into cones. Pollen cones are soon shed, but seed cones persist for one to several years. Sperm is free-swimming. Seeds have a hard inner seed coat and a fleshy outer layer, aiding dispersal by animals.

Coontie family

Uses: Starch from the pith of many Cycas species is used to make sago in Asia, but has to be carefully washed to leach out toxins. All untreated parts of cycads are extremely poisonous to humans. Cycas revoluta is a popular house and garden plant. Cultivation of various Cycas species is common throughout the tropics and subtropics. Etymology: Cycas is derived from the Greek κυκας (kykas), a word first used by Theophrastus, thought to be a scribal error for κούκας (koikas), meaning ‘palm trees’, the accusative plural of koix, a word from an unknown pre-Greek language. 74

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Distribution: This family is patchily distributed in tropical America, Sub-Saharan Africa and Australia. These are unisexual plants with subterranean or emergent, simple or irregularly branched trunks that are clothed with persistent leaf bases. New leaves are formed in flushes. Leaves are pinnately compound (but bipinnate in Bowenia); the leaves are not coiled, except in Bowenia and Stangeria (the latter with

Phylogeny and evolution: Stangeria is peculiar in its circinnate vernation, resembling a fern, and it was originally described as one. It was placed with Bowenia in Stangeriaceae, both sharing this feature. Molecular phylogenetic studies have shown these two genera are embedded in Zamiaceae.

GINKGOALES

GYMNOSPERMS

Extant species diversity is the result of a recent radiation within the past five million years, and Zamiaceae should therefore not be considered living fossils, even though some are remarkably similar to fossil taxa. Genera and species: Zamiaceae include nine genera with c. 200 species: Bowenia (2), Ceratozamia (27), Dioon (15), Encephalartos (65), Lepidozamia (2), Macrozamia (c. 30), Microcycas (1), Stangeria (1) and Zamia (c. 55).

Uses: Young seeds of Dioon edule are ground and cooked into tortillas. Seeds of some Encephalartos species are also eaten in South Africa, and the exudate from cones is eaten by children and birds. Zamiaceae leaves are used for decoration, especially in some religious ceremonies in Central America. Nearly extinct: Encephalartos woodii is known only from a single male plant from which cuttings are growing in botanic gardens.

It is extinct in the wild and cannot be reintroduced in the absence of a female plant. The oldest specimen is in the Temperate House of the Royal Botanic Gardens, Kew. Etymology: Zamia is derived from the Latin ‘nuces zamiae’, in translation of Pliny the Elder; this is an erroneous transcription of the phrase ‘nuces azaniae’, referring to pine nuts, probably from Classical Greek αζάινειν (azainein), to dry.

GINKGOALES This order, which originated c. 265 million years ago, occurred world-wide during the Mesozoic. Some authors have hypothesised that they were derived from Palaeozoic pteridosperms, but the morphology of these Carboniferous Ginkgo-like plants is difficult to interpret. Leaves resembling Ginkgo are known from Permian fossils onwards.

27. GINKGOACEAE Maidenhair-tree family

These are unisexual deciduous trees up to 30 m, with irregularly furrowed grey bark. They branch irregularly with long shoots and brachyblasts (stubby short shoots) at regular intervals, each bearing a cluster of leaves. Leaves are fan-shaped, with flabellate venation (radiating out from the petiole), and are slightly fleshy, light green, turning bright yellow in autumn, and deciduous. Male cones are borne on short catkin-like shoots. Pollen is spherical, and sperm is free-swimming. Seeds are formed on Ginkgo biloba, male, Royal Botanic Gardens, Kew, UK [27]

short leaf-like shoots, obovoid to ellipsoid, and c. 2 cm in diameter, yellow to orange, glaucous, with an apical scar, maturing in a single season, usually one (rarely two) per peduncle. The outer seed-coat softens and gives off a foul odour of butyric acid when ripe. Distribution: Ginkgo biloba is native to China, but its occurrence in the wild is uncertain. It is reported to occur naturally in mountain valleys in Zhejiang Province, but because this area has seen human inhabitation for some 1,500 years it is plausible that its persistence is due to ancient cultivation near Buddhist monasteries. The great majority of ginkgos have been planted as ornamental trees, and the species is nearly cosmopolitan. Hardy to -30˚C, it is frequently planted in hemiboreal, temperate and subtropical areas around the world. Phylogeny and evolution: The fossil record shows Ginkgo to have been widespread, diverse and abundant in the Mesozoic. Because Ginkgo biloba, female cones, Ann Arbor, Michigan, USA [27]

nearly identical plants are known from fossils some 200 million years old, the single extant species is often referred to as a living fossil. It is sister to the conifers plus gnetoids. Genera and species: Ginkgo biloba is the sole species in its order (Gingkgoales), family (Ginkgoaceae) and genus (Ginkgo). It is a relict of a once diverse and widespread lineage of plants. Uses: Fresh or canned seeds (with the soft outer layer removed) are a delicacy in East Asian cuisine. Due to their high levels of tolerance to pollution ginkgos are commonly planted along roads and in city parks. Six Ginkgo trees near the blast centre of the atomic bomb in Hiroshima (August, 1945) resprouted without deformation and are still alive today. Etymology: Ginkgo is derived from the Chinese 銀杏 (yínxìng), meaning ‘silver apricot’. The same characters are used for this tree in Japan, but there they are read as ‘ginkyō’. This was how Engelbert Kaempfer (1712), the first Westerner to describe this species in 1690, transcribed its name. Kaempfer’s name was misread by Linnaeus (1753), who erroneously changed the y into a g and published the genus as Ginkgo, which became the accepted name, albeit difficult to pronounce. Plants of the World

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WELWITSCHIALES

Welwitschia mirabilis, male cones with pollinator in Namibia (FF) [28]

GYMNOSPERMS

Welwitschia mirabilis, female cones, Namibia (FF) [28]

Welwitschia mirabilis, female, Namibia (FF) [28]

WELWITSCHIALES This order is known from the fossil record in which there is a seedling of c. 110 million years old. It was once more widespread, pollen being known from North America, Portugal and Brazil.

28. WELWITSCHIACEAE Tumbo family

Tumbos are dioecious plants with short woody, unbranched, cup-shaped stems that widen to a concave disc up to 1 m across with a long taproot. The two (rarely three) strapshaped leaves grow from a basal meristem throughout the life of the plant (estimated to be at least 1,000 years in some individuals), the leaves dying at the tip and fraying. These remarkable leaves have a subparallel venation that occasionally anastomoses or terminates 76

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blindly. From the woody stem, near the leaf bases, small branches arise that bear either male or female cones. Male cones are terminal in groups of two or three on each branch. Female cones consist of a single ovule enclosed in an integument and another layer derived from two confluent primordia with two bracts. There is one seed per cone, which has a wing and is dispersed by wind. Seeds germinate only when water is available, but seedlings are susceptible to fungi, making establishment from seed difficult in cultivation. The two cotyledons photosynthesise for 1.5 years, after which they wither away and are replaced by two leaves. Distribution: This family has two disjunct populations, one in northern Namibia, the other in southern coastal Angola. Phylogeny and evolution: A fossil seedling called Cratonia cotyledon from northeastern

Brazil has a similar vasculature to that of Welwitschia and is perhaps 114–112 million years old. Other fossils with putative relationships to Welwitschia have been found in the same area. Genera and species: Welwitschiaceae include a single genus with a single species, Welwitschia mirabilis, which is divided into two subspecies: W. mirabilis subsp. mirabilis in Angola and W. mirabilis subsp. namibiensis in the southern part of its range. The subspecies differ in characters of the male cones. Etymology: Welwitschia was named by Joseph Hooker for its discoverer, Austrian explorer and botanist Friedrich Martin Josef Welwitsch (25 February 1806–20 October 1872), who, when he discovered it in Angola named it Tumboa, after its local name. Welwitschia has been conserved over Tumboa.

GNETALES

GYMNOSPERMS

GNETALES This order sometimes includes Welwitschiales and Ephedrales, but they are here restricted to the single family Gnetaceae. They have an estimated age of c. 160 million years in most studies, although there is a probable gnetalean fossil of c. 260 million years old.

29. GNETACEAE Emping family

drupe-like; they are enclosed in a red, orange or yellow, usually fleshy false seed coat, with two cotyledons. Distribution: This family occurs in tropical regions, the Amazon Basin, tropical West Africa, India, Southeast Asia and Malesia.

These are unisexual lianas, sometimes trees, that have stems with swollen nodes. Leaves are opposite, petiolate, without stipules, simple, the margins entire, the venation pinnate and apices usually acuminate (driptips), resembling leaves of angiosperms. Both male and female strobili are compound, spike-like, terminal or lateral structures (megastrobili), sometimes emerging from old stems. Each spike consists of a straight axis above a basal pair of opposite, connate bracts, the axis bearing three to six superposed cupules, each containing male or female strobili. Male strobili consist of a stamen and a bract. Female strobili consist of an ovule with two integuments and a bract. Seeds are Gnetum gnemon, seeds, Singapore [29]

Phylogeny and evolution: Gnetaceae may have evolved some 250 million years ago, but the current radiation diverged during the Oligocene and Miocene, late in the history of this ancient lineage. The leaves and fertile structures are angiosperm-like. Nonetheless, DNA studies have shown that Gnetaceae (with Welwitschiaceae and Ephedraceae) are more closely related to Pinaceae than to angiosperms, and hence they are placed here next to the conifers. Genera and species: Gnetaceae consist of the single genus Gnetum with c. 30–35 species. Uses: Gnetum gnemon is cultivated commercially in Indonesia, where the kernels of the large nutritious seeds are beaten flat and fried, becoming the popular snack called ‘emping’.

The young leaves, flowers and fruits are used as vegetables, eaten raw, boiled or roasted. Fibres from the bark provide durable cordage for, for example, fishing nets, rope and bags. The wood is used for paper and house construction. Topical application reduces biting by mosquitoes, and enzymic inhibition prevents insect predation of foliage, which may be applicable in pest control. Scleroderma sinnamariense, a fungus usually associated with Gnetum gnemon, produces an edible fruiting body. Gnetum gnemon is frequently planted as a shade tree and has been used for reforestation of dryland habitats. The trees are sometimes interplanted on farms to provide living stakes for the cultivation of yams. The leaves of Gnetum africanum, are commonly consumed as a vegetable throughout Africa, locally called fumbwa. It is not cultivated, but leaves are gathered from the wild, cut into small strips, boiled and mixed with palm oil or peanut butter. Etymology: The name Gnetum is derived from Maluku or Malay gnemon utan, following the description of the plant by Rumphius, Herbarium Amboinense 1: 183–184, plates 71–73. 1741.

Gnetum gnemon, Singapore [29]

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EPHEDRALES

Ephedra californica, male cones, California, USA [30]

GYMNOSPERMS

Ephedra equisetina, female cones at the time of pollen transfer, Royal Botanic Gardens, Kew, UK [30]

Ephedra americana, mature female cones bearing seeds, Copenhagen Botanical Garden, Denmark [30]

EPHEDRALES The distinctively ridged pollen of Ephedrales is found in the fossil record at least since the Late Triassic. It is sometimes included under Gnetales, but here it is used for the single family Ephedraceae.

30. EPHEDRACEAE Jointfir family

are enclosed by a pair of fused bracteoles that become fleshy or leathery; these are shed with the seed. Female cones have one or two seeds that are yellow to dark brown and have two cotyledons. Distribution: This family occurs in North America, western South America, Mediterranean Europe, North Africa and warm temperate Asia, usually in arid habitats.

These are dioecious (or sometimes monoecious) shrubs and trailing vines with photosynthetic, green, round, jointed branches. Leaves are mostly non-photosynthetic, scale-like, simple, opposite or in whorls of three. They are connate at the base to form a sheath and superficially resemble the sheath-like leaves of Equisetum. Male or female cones are formed singly or in whorls at nodes. Opposite or whorled membranous bracts each subtend a small cone. Ovules 78

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Phylogeny and evolution: Ephedraceae evolved before the early Cretaceous, when they comprised at least two genera. The extant genus Ephedra dates back to the Cretaceous, making the extant species relicts, that were well adapted when semi-arid climates expanded across the World during the Oligocene. Genera and species: Ephedraceae consist of the single extant genus Ephedra, which has c. 40 species.

Uses: In many native cultures where Ephedra occurs it is used for medicinal purposes, such as treatment for cough, poor circulation and allergies. This is because the plants are rich in tannins and often the alkaloid ephedrine. It is commonly applied in herbal treatments and is an ingredient of modern cold and allergy medicines (usually as pseudephedrine). It is also used as a dietary supplement in energy drinks and weight-loss preparations. Ephedrine is not present in the New World species, and the psychotropic properties ascribed to Mormon tea (E. funerea) remain uninvestigated. Etymology: Ephedra is a Greek name given by Pliny to the horsetail, which in turn is derived from έφεδρος (ephedros), ‘sitting upon’, from επί (epi), on, and έδρα (hedra), a seat, probably referring to the segments that sit upon each other.

PINALES

GYMNOSPERMS

PINALES Fossils assigned to Pinales have been dated to c. 155 million years ago, but molecular clock estimates date this order to over 200 million years. The order consists of the single family Pinaceae, which were formerly widespread and now dominant in some parts of the Northern Hemisphere.

31. PINACEAE Pine family

These bisexual, evergreen and deciduous trees, occasionally shrubs, are resinous and aromatic in all parts. Terminal branches are radially symmetrical, lateral branches are well developed and similar to leading shoots or are reduced to short shoots. Leaves are simple, needle-like to linear, and sessile to

short petiolate. They are borne singly and then spirally arranged on long shoots or in fasciculate tufts on short shoots. Male cones are axillary, solitary or clustered, and ovoid to ellipsoid or cylindrical. Sporophylls are overlapping, bearing two pollen sacs. Female cones are compound, axillary, solitary or grouped, and scales are overlapping and free from the subtending bracts for most of their length. Each scale bears two ovules on the upper side. Seeds are winged, the wing free from subtending bracts. The number of cotyledons can vary from two to 15 (rarely up to 24). Distribution: This family has a mostly temperate Northern Hemisphere distribution. It occurs in North America south to Nicaragua, the West Indies, throughout temperate (including North Africa) and

Picea likiangensis near Lijiang, Yunnan, China [31]

tropical Eurasia south to Sumatra and the Philippines. Pollination is by wind, causing allergies in some humans where Pinaceae are abundant. Seeds are dispersed by wind. Some species become aggressive invasives in parts of the world where they have been planted for timber. Phylogeny and evolution: The family is known from the fossil record since the Cretaceous. Originally the family included all conifers, but it is now restricted to the clade including the genus Pinus. There is good evidence to support the hypothesis that there has been a recent radiation of species in the three largest genera, Abies, Picea and Pinus. Some molecular studies have controversially indicated that the family is sister to the three families of gnetoids.

Pinus tabuliformis var. mukdensis, Royal Botanic Gardens, Kew, UK [31]

Abies forrestii, Royal Botanic Gardens, Edinburgh, Scotland, UK [31]

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PINALES Genera and species: Pinaceae include 11 genera with 224 species: Abies (49), Cathaya (1), Cedrus (4), Keteleeria (3), Larix (12), Nothotsuga (1), Picea (33), Pinus (c. 115), Pseudolarix (1), Pseudotsuga (7) and Tsuga (8). Uses: Most commercial softwood timber is from members of Pinaceae. They are also a major source of pulp for paper production and for tar and turpentine, essential oils etc. In Scandinavia, pine bark flour (Pinus sylvestris) is used in local cuisine, and young twigs of

GYMNOSPERMS

spruce (Picea abies) are used for tea or to flavour liquor. Many species of Pinus produce edible seeds. Commercially, pinenuts (piñones or pignolias; Pinus pinea) are harvested for this purpose. Several species of Picea and Abies are commercially grown as Christmas trees. In Mexico, some species of long-leaved pine are used for basketry. Lebanon cedar (Cedrus libani), now a rare species, was once abundant in the Levant and was harvested for its fragrant wood; the Temple of Solomon was built of this timber (Old Testament; Kings 5: 6). Larix has hard, heavy, decay-resistant

Larix decidua, young female cones, Royal Botanic Gardens, Kew, UK [31]

Cedrus libani forest in Lebanon [31]

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wood, which when knot-free (as ‘boatskin larch’) is in great demand for yacht building. Pines and other species are commonly used in dendrochronological studies to date buildings and ruins accurately. Pinaceae are planted as ornamentals throughout temperate and subtropical regions of the world, and cones of Pinaceae are often used in dried flower arrangements and painted for use as Christmas decorations. Etymology: Pinus is Latin for a pine tree.

Pseudotsuga menziesii, female cone with three-parted bracts, California, USA [31]

Pinus banksiana, male cones, Royal Botanic Gardens, Kew, UK [31]

ARAUCARIALES

GYMNOSPERMS

ARAUCARIALES Podocarpaceae fossils are known since the Early Triassic, c. 240 million years ago, whereas Araucariaceae fossils date back to the Mid Jurassic, c. 160 million years ago. The order consists of two families that are predominantly found in the Southern Hemisphere and are believed to be of Gondwanan origin.

Distribution: This mostly Southern Hemisphere family is found in southern South America, Southeast Asia, Malesia, the Philippines, New Caledonia, Australasia and some Pacific Islands.

Phylogeny and evolution: Originating in the Triassic, the family diversified and spread to both Hemispheres during the Jurassic and Early Cretaceous, becoming a significant component of Gondwanan forests until the late Cenozoic. Angiosperms dominated forests in the Cretaceous, which probably resulted in the extinction of some araucarian lineages, but some new genera also evolved. Continental separation and associated climatic drying and cooling reduced the ranges of these predominantly tropical trees, but volcanic activity provided new habitats in Australasia where Araucariaceae diversified. Now mostly restricted to some Southern Hemisphere areas, the family was formerly more widespread, Araucaria fossils being known from both Hemispheres from the Jurassic. Most extant Araucariaceae must have evolved from the Early Tertiary and dispersed to the islands of Australasia where they diversified; these islands provide exactly the tropical rainforest habitats where angiosperms were thought to have most successfully replaced conifers. This has more to do with the adaptability of the more recently evolved species of Agathis than their ability to compete effectively with angiosperms.

Araucaria angustifolia forest in Santa Catarina, Brazil [32]

Wollemia nobilis, male and female cones, Royal Botanic Gardens, Kew, UK [32]

32. ARAUCARIACEAE Kauri-tree family

These evergreen bisexual and unisexual trees have verticillate branches and spirally arranged leaves. Leaves usually have parallel venation. Male cones are cylindrical, and the sporophylls are numerous, each bearing about a dozen pollen sacs. Female cones are subglobose to ovoid and mature in two years. They disintegrate on the tree upon maturity, the cone scales one-seeded, without distinct bracts. Seeds have four cotyledons that are fused into two double cotyledons.

Genera and species: Extant Araucariaceae consists of three genera with 41 species: Agathis (21), Araucaria (19) and Wollemia (1). Uses: Many species are important timber trees because they are usually among the largest trees in the forest. Seeds of Araucaria angustifolia and A. bidwillii are locally consumed. Many species are common ornamentals, especially Araucaria heterophylla, the Norfolk Island pine, a common houseplant, and A. araucana, the monkey-puzzle tree, which got its name because when it was first introduced, it was said to be a puzzle how a monkey could climb such a prickly tree. The recently discovered Wollemi pine (Wollemia nobilis) is a peculiar conifer from Wollemi National Park, New South Wales, Australia, that has been saved from possible extinction by being brought into horticulture and distributed to gardens worldwide. The species thrives in mild temperate climates. Etymology: Araucaria is derived from Araucania, a region inhabited by the Mapuche (or Spanish: Araucana) people, now in Chile, where Araucaria araucana is native. Araucaria angustifolia, seeds, Curitiba, Brazil [32]

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Distribution: This family occurs in Neotropical mountains from Mexico and the Caribbean to Chile, and from tropical Africa, India, China and Japan to Australia, New Zealand and New Caledonia. The family is most diverse in Australasia.

Phylogeny and evolution: Podocarpaceae are most closely related to Araucariaceae, with which they form a clade. Both families are predominantly found in the Southern Hemisphere, but this may be more because they are prevalent in tropical and subtropical forests, which are currently mostly situated in the Southern Hemisphere continents. Fossils from the Northern Hemisphere are known. The family includes some peculiar members, such as the only parasitic gymnosperm, Parasitaxus usta, growing usually on the roots or branches of another Podocarpaceae, Falcatifolium taxoides, both endemic to New Caledonia. Even though Parasitaxus makes connections with the xylem of Falcatifolium by which it extracts water and nutrients, it gains carbon from fungi, so it is partly mycoheterotrophic, partly parasitic. Retrophyllum minus is unusual in its preference for an aquatic habitat, growing as an emergent small tree in streams also in New Caledonia. The genus Phyllocladus was, due to its unusual foliage morphology (phylloclades), presence of an aril-like structure and different pollination mechanism, segregated in a separate family, but apart from these differences it shares numerous characters with other Podocarpaceae (winged pollen, an epimatium etc.) and molecular studies show this genus to be closely related to other Podocarpaceae.

Prumnopitys andina, female and male cones, National Botanic Gardens, Glasnevin, Ireland [33]

Podocarpus nivalis, female cone, Royal Botanic Gardens, Edinburgh, Scotland, UK [33]

33. PODOCARPACEAE Yewpine family

These evergreen bisexual and unisexual shrubs and trees are usually terrestrial, rarely aquatic (Retrophyllum) or epiparasitic (Parasitaxus). Leaves are spirally arranged, sometimes opposite, and they are scale- or needle-like or flat and leaf-like, linear to lanceolate. Male cones are catkin-like with numerous crowded stamens, imbricate, each stamen with two pollen sacs. Female cones mature in one year and are reduced to a few fleshy bracts or scales (epimatia), borne on a thin peduncle containing a single ovule. Seeds are covered by a fleshy structure and have two cotyledons.

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Genera and species: Podocarpaceae consist of 19 genera with 185 species: Acmopyle (2), Afrocarpus (5), Dacrycarpus (9), Dacrydium (21), Falcatifolium (5), Halocarpus (3), Lagarostrobos (1), Lepidothamnus (3), Manoao (1), Microcachrys (1), Nageia (6), Parasitaxus (1), Pherosphaera (2), Phyllocladus (5), Podocarpus (104), Prumnopitys (9), Retrophyllum (5), Saxegothaea (1) and Sundacarpus (1). Uses: The family includes some minor timber trees, and many are threatened by overharvesting. Several species are used for cabinetry, furniture, and carving and turning because of their fine grain, soft wood and fragrance. An essential oil, methyl eugenol, from the wood of the longlived tree Lagarostrobos franklinii is now chemically synthesised and used as a wood preservative and an insecticide. Oil from seeds of Nageia nagi is edible and contains eicosatrienoic acid, a compound that can reduce swellings. Stems are used in floristry as everlastings. Etymology: Podocarpus is derived from Greek ποδός (podos), a foot, and καρπός (karpos), a fruit, referring to the obliquely stalked fruit that resembles a foot kicking a ball.

Retrophyllum minus, New Caledonia [33]

CUPRESSALES

GYMNOSPERMS

CUPRESSALES With an estimated age of some 245 million years, this is another old order that was formerly more diverse and widespread than it is at present. It consists of three families, Cupressaceae, Sciadopityaceae and Taxaceae.

34. SCIADOPITYACEAE Umbrella-pine family

These are evergreen bisexual trees with whorls of brown scales (which are the actual leaves) from which green needle-like cladodes (which are short stems) emerge. These cladodes resemble the needles of Pinaceae, but are composed of stem tissue and have two vascular bundles. Male cones are formed in dense terminal clusters. Female cones are subsessile, ovoid and disintegrate soon after

the seeds have been released. Scales are green, thin and flat, with five to nine seeds per scale. Seeds are flattened and ovoid with a narrow wing along each side. They are orange-brown and have two cotyledons. Distribution: This family is currently only found in southern Japan (southern Honshu, Kyushu, Shikoku) in mid-elevation cloud forests. It is listed as vulnerable, due to harvesting of the water-resistant wood for boat-making. Phylogeny and evolution: Sciadopityaceae are sister to Taxaceae and Cupressaceae. The family was widespread in the past and, before its discovery in Japan in the 19th century, it was only known from fossils from the Early Cretaceous 230 million years ago. It was more diverse in the past; Sciadopitys was, for instance, diverse in Tertiary Europe.

Sciadopitys verticillata, male cones, Royal Botanic Gardens, Kew, UK [34]

Genera and species: Sciadopityaceae consist of the single genus Sciadopitys that has only one extant species, S. verticillata. Uses: The koyamaki or Japanese umbrella pine is a popular garden plant throughout the temperate zones, despite its relatively slow growth. It is remarkably hardy and grows well even in cold climates. The wood is sometimes used for timber, but not on a commercial scale. Baltic amber is composed of the resin of extinct Sciadopityaceae. Amber has been appreciated for its colour and is often used for jewelry, but amber is also used in perfumes and folk medicine. Etymology: Sciadopitys is derived from the Greek σκιαδος (skiados), a whorl, and πίτυς (pitys), a fir tree.

Sciadopitys verticillata, female cone, Royal Botanic Gardens, Kew, UK [34]

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35. CUPRESSACEAE Cypress family

These are bisexual and unisexual, resinous and aromatic terrestrial trees and shrubs, without or with knee-like roots in aquatic Taxodium. Lateral branches are similar to leading shoots and are densely clothed by scale-like leaves or by leaf bases. Leaves are simple, persistent or deciduous, alternate and spirally arranged (sometimes twisted and appearing two-ranked), opposite in four ranks, or whorled, deltate to linear, sessile or petiolate; twigs are often heterophyllous. Male cones are axillary or terminal, solitary or in clusters of two to five (to 20), sometimes in panicles. They are spherical to oblong, and the sporophylls overlap, bearing two to ten pollen sacs. Female cones are compound, axillary, terminal, solitary or in clusters of two to five (to c. 100); the scales are usually overlapping and fused to subtending bracts, but sometimes with only the bract apex free, the fused scale/ bract complex peltate, oblong or cuneate, woody or fleshy at maturity, bearing one to 20 ovules. Seeds are wingless or with two or three wings, and the number of cotyledons varies from two to five, occasionally up to nine. Distribution: This family is nearly cosmopolitan. It is found on all continents except Antarctica. The generic diversity is greatest in the Southern Hemisphere and shows a relictual distribution, except for the largest genus Juniperus, which is widespread across the Northern Hemisphere, extending from the Arctic to the subtropics. Phylogeny and evolution: Fossils of Cupressaceae have been known since the Jurassic. The family was formerly divided into two

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families: Cupressaceae sensu stricto, with whorled or opposite (four-ranked) leaves, and Taxodiaceae, with mostly alternate leaves. The two former families are united in having female cones with fused bract/scale complexes. The leaf orientation has proven not to be an important character to distinguish between families and molecular studies have confirmed a close relationship between these groups, which is why they are now united. The clade of Cupressus s.l. has been contested in the past, but the group appears to be best treated as five genera, the Vietnamese golden cypress Xanthocyparis and the Nootka cypress Callitropsis forming a grade leading up to the New World Hesperocyparis, the Old World Cupressus and panboreal Juniperus. In principle these can all be merged into a single genus, but this will result in the loss of the well-established and large genus Juniperus, destabilising the taxonomy of this economically important group. Callitropsis nootkatensis, a commonly cultivated species has, however, been taxonomically unstable, being formerly placed in Cupressus, Chamaecyparis and Xanthocyparis. Actinostrobus and Neocallitropsis have been merged with Callitris. Genera and species: Cupressaceae consist of 30 genera with c. 146 species: Arthrotaxis (3), Austrocedrus (1), Callitris (22), Callitropsis (1), Calocedrus (4), Chamaecyparis (5), Cryptomeria (1), Cunninghamia (2), Cupressus (9), Diselma (1), Fitzroya (1), Fokienia (1), Glyptostrobus (1), Hesperocyparis (16), Juniperus (c. 50), Libocedrus (5), Metasequoia (1), Microbiota (1), Papuacedrus (1), Pilgerodendron (1), Platycladus (1), Sequoia (1), Sequoiadendron (1), Taiwania (2), Taxodium (2), Tetraclinis (1), Thuja (5), Thujopsis (1), Widdringtonia (4) and Xanthocyparis (1). Uses: Cupressaceae include the largest (Sequoiadendron giganteum) and tallest (Sequoia sempervirens) trees in the world. Both species (and numerous other members of the family) are commonly harvested for timber. Wood of many species is resistant to fungal decay and termite damage and is therefore often used in construction.

Chamaecyparis and Cunninghamia are in demand in Japan and China, respectively, for coffin wood. Wood of many species is aromatic and therefore often called ‘cedar wood’, and is then usually confused with the wood of Cedrus (Pinaceae) or Cedrela (Meliaceae), which are all aromatic timbers. Fleshy cones (‘berries’) of juniper (Juniperus communis) are commonly used to flavour English gin, Dutch jenever and German sauerkraut. Many species are popular in horticulture as architectural elements in the garden. Because of the association with grieving, Cupressaceae have been planted in graveyards in various cultures in Europe (Juniperus), the Middle East (Cupressus) and East Asia (Platycladus, Thuja). The columnar Cupressus sempervirens or arborvitae (the ‘tree of life’) has been a traditional plant in graveyards since ancient times throughout the Mediterranean, and since Roman times it has been grown in gardens, providing a picturesque ‘Italianate’ feature. Taxodium grows in swamps and is remarkable in forming knee roots (the function of which is unknown). Etymology: Cupressus is the Latin name for a cypress tree, from Greek, κυπάρισσος (kuparissos), probably derived from an unknown Mediterranean language. This was a tree sacred to the god Pluto, who changed his grieving boy companion Cyparissos into a cypress tree, hence its abundance in Mediterranean graveyards. Juniperus communis, female cones, Finland [35]

CUPRESSALES

GYMNOSPERMS

Callitris columellaris, female cone, Western Australia [35]

Cryptomeria japonica, female cones, Royal Botanic Gardens, Kew, UK [35]

Cupressus sempervirens, male and female cones, France [35]

Papuacedrus papuana var. arafakensis, Glasgow Botanic Gardens, Scotland, UK [35]

Taxodium distichum, Twickel Estate Gardens, the Netherlands [35]

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36. TAXACEAE Yew family

GYMNOSPERMS

bear a single seed, and the hard seed coat is partially or wholly covered with a fleshy or leathery aril, resembling a berry. Seeds have two cotyledons. Distribution: This family occurs in North and Central America, North Africa and throughout Eurasia to Malesia and New Caledonia.

These are bisexual and unisexual evergreen trees and shrubs, usually not resinous or aromatic. Lateral branches are similar to leading shoots. Leaves persist for several years, are shed singly, are arranged alternately and spirally but are often twisted so as to appear two-ranked, rarely opposite. Leaves are simple, needle-like and linear to linearlanceolate. Male cones are solitary or clustered, axillary. They are globose to ovoid, with the sporophylls bearing two to 16 pollen sacs. Female cones are axillary and are reduced to one or two ovules subtended by bracts. Cones

Phylogeny and evolution: Genetic relationships among Taxaceae have proven difficult to assess, but it seems that Amentotaxus is part of core Taxaceae, whereas Cephalotaxus is sister to the rest, although with weak support. Tertiary fossils of Amentotaxus differ little from extant species, making these ‘living fossils’. Members of Taxaceae are characterised by slow rates of evolutionary change. The family is sometimes divided into two or more subfamilies. Genera and species: Taxaceae include six genera with 32 species: Amentotaxus (6), Austrotaxus (1), Cephalotaxus (11), Pseudotaxus (1), Taxus (7) and Torreya (6).

Uses: The wood of some species of Taxaceae is used for carving. The fleshy aril of Taxus is edible, but it has to be consumed with caution because the seeds of all species of Taxus are poisonous, containing the alkaloid taxol; this compound inhibits mitosis and is used in synthesised form (paclitaxel) in chemotherapy treatment for various types of cancer. Clippings from yew hedges are collected from gardens around Western Europe to harvest this compound. Like cypress in the Mediterranean, yew is evergreen and often planted in churchyards and ancient burial grounds in Britain, Ireland and Normandy (perhaps as a remnant of druidic tradition). Yew is also the preferred wood for making longbows due to its flexibility. It was also a prized wood to use for making musical instruments, especially lutes. Seeds of the Japanese kaya, Torreya nucifera, are edible and pressed to produce cooking oil. The yellow wood of kaya is used for go boards, a game played in East Asia. Etymology: Taxus is Latin for a yew tree.

Cephalotaxus harringtonia subsp. drupacea, female cone, National Botanic Gardens, Glasnevin, Ireland [36]

Ancient yew, Taxus baccata, in the churchyard of Downe Village, Kent, UK [36]

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Taxus baccata, male cones, Box Hill, England, UK [36]

Taxus baccata, female cones, Royal Botanic Gardens, Kew, UK [36]

ANGIOSPERMS Imagine a world without flowers – it was for the great majority of its existence. Some 245–160 million years ago a group of plants emerged that had their reproductive leaves organised in whorls with sterile leaves enclosing them and large female organs covered in tissue to protect the developing seed. The precise way flowers developed and from what pre-existing group of gymnosperms (seed plants without flowers) flowering plants evolved is still a mystery. Evolutionary developmental biologists have identified the genes that control floral development in angiosperms and their counterparts in gymnosperms, but they have not unravelled the way in which these genes changed and first started being able to produce flowers. We thus know the “before and after” but not the details of the route taken. Darwin referred to the origin and rapid rise to dominance of the angiosperms as an “abominable mystery”, but how the first flowers evolved is an equally great conundrum. Early f lowers co-evolved with the appearance of a greater diversity of insect groups, particularly beetles, which fed on the f lowers, pollen and seeds, but also pollinated these organs. From herbivory, plants and insects co-evolved, and flowers came to host their pollinators, providing food rewards while not being greatly harmed. Flowers with lots of stamens evolved, dusting the visitors while they ate fleshy petals with pollen then transported to a nearby flower. The developing seeds were located deep in tissues, f irst for their protection but later as fruits to attract animals to consume and disperse the seeds. One benefit of seeds is that plants became less dependent on permanent water. Seeds could survive in dormant stages when

conditions were unsuitable for growth, due to drought, fire or frost. Thus, seeds allowed early flowering plants to disperse more widely, and interactions of seeds and fruits with animals resulted in diversification of forms. This diversity of plants spurred greater diversity in animals, which evolved new ways of using plants for food and shelter. Flowering plants became so successful that they rapidly replaced other vegetation formed by earlier groups of seed plants. The first rainforests formed, modifying the climate more than the earlier gymnosperms ever managed, and these new habitats acted as a spur to further diversification. Angiospermdominated landscapes became the most diverse the planet has ever witnessed. With the evolution of large grazing mammals and grasses (Poaceae), grasslands evolved, where the growing stems of the plants are below ground, not exposed to grazing mouths and trampling hoofs. Annual life histories developed as another method of overcoming periods of the year when growing conditions were unsuitable (too wet or dry, hot or cold), and these plants put all their accumulated resources into the seeds that would produce their next generation. When finally a peculiar bipedal ape came on the scene, the diversity of flowering plants was enormous, and with its large brain and hands with opposable thumbs it made good use of the existing resources, especially the annuals, which were more easily domesticated and could form the basis of settled agriculture. Humans invented language, agriculture, art, science, religion and culture, all based on the diversity of f lowering plants and the resources they provide. Do we still need plants? If you see the big financial centres of

A selection of angiosperm seeds and fruits

glass and steel, you might wonder, but without plants there would be no food, medicine, paper, clothes, houses, fuel or clean air. Moreover, plants sustain people, making them happy and healthy. Flowers are important cultural symbols as well — a rose for England, a thistle for Scotland, an orchid for Singapore, a tulip for Holland and Turkey, a protea for South Africa and the sacred lotus for Buddhism. Give a rose to a loved one or a lily as a sign of sympathy when a loved one dies. A world without flowers would not be quite the same. Etymology: The word angiosperm is derived from the Greek αγγείων (angeion or aggeion), a vessel or capsule, and σπέρμα (sperma), a seed, in reference to the fact that the seeds of flowering plants are enclosed in a layered structure as opposed to those of gymnosperms (which are naked).

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THE ANA GRADE FAMILIES

AMBORELLALES

THE ANA GRADE FAMILIES (AMBORELLANAE, NYMPHAEANAE, AUSTROBAILEYANAE) We refer to these families as the ANA grade because they do not form a clade. These three superorders sucessively are sister to the three major clades of flowering plants: magnoliids, monocots and eudicots. They are composed of Amborellales, the sister of the remainder of the flowering plants; Nymphaeales, an order of water lilies and associated families that have a long fossil history; and Austrobaileyales, an order of vines, shrubs and trees with spirally arranged flower parts. Previously these were placed among the “dicots” (they do indeed have two seed leaves or cotyledons), but as previously treated by most taxonomists, the monocots are embedded among dicots, and thus we no longer refer to dicots as a meaningful evolutionary unit. The ANA grade is treated as series of separate entities apart from the magnoliids, monocots and eudicots.

AMBORELLANAE AMBORELLALES This order diverged c. 130 million years ago and is probably sister of the rest of the angiosperms. They are now only found on New Caledonia, which emerged from the ocean c. 37 million years ago. Amborella must have lived elsewhere for a long time before it dispersed to New Caledonia and became extinct in its original area of occurrence.

37. AMBORELLACEAE Amborella family

These dioecious irregularly branching, typically sprawling shrubs can sometimes appear vine- or tree-like and have simple, alternate, two-ranked leaves without stipules. Leaf margins are undulate and coarsely dentate. Venation is pinnate, the veins connecting near the margin. Inflorescences are axillary and cymose. Flowers are small, white, the perianth undifferentiated, spirally arranged, basally connate, with five to eight petals and several petaloid bracteoles. Male flowers have six to 25 sessile anthers. Female flowers have a few staminodes, superior ovaries and several (three–six) free carpels that develop into a cluster of stalked, fleshy red drupes. Seeds are pitted with pockets of resinous substances. 88

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Distribution: The sole species occurs in forests of New Caledonia at between 200 and 1,000 m on schistose soils and is locally common between the Dogny Plateau and the Tipindjé River basin. Phylogeny and evolution: Amborella was formerly included in Monimiaceae or, if recognised as a separate family, was often included in Laurales. DNA studies have since shown it to be sister to all other angiosperms. As a result, the family has now been placed in its own order Amborellales. Despite its isolated position outside the main angiosperm clade, this family should not be considered primitive and clearly exhibits some derived characters, including unisexual

Amborella trichopoda, male flowers, New Caledonia [37]

flowers. Pollination is by wind and insects. Amborellaceae lack developed wood vessels, but if this is to be interpreted as primitive or derived is not certain. There has been some controversy about its phylogenetic position, and some studies have instead placed it in a clade with Nymphaeales (including Hydatellaceae; see below), but the majority of studies support its position as stated above. Genera and species: Amborellaceae consist of the single species, Amborella trichopoda. Etymology: Amborella is the diminutive form of ambora, a native Malagasy plant name for Tambourissa (Monimiaceae), to which Amborella was originally thought to be related.

Amborella trichopoda, female flowers, New Caledonia [37]

NYMPHAEALES

THE ANA GRADE FAMILIES

NYMPHAEANAE NYMPHAEALES Families 38–40 form the order Nymphaeales, which are the waterlilies and relatives. They are all aquatic rhizomatous plants that have boat-shaped pollen and fused cotyledons. Fossils of Nymphaeales date back to the Cretaceous, and it has been suggested that the fossil Archaefructus, thought to be about 124 million years old, belongs to this order. Archaefructus is putatively one of the earliest known fossil flowering plants, although opinion about its position and relationships as well as its age have been subject to controversy.

38. HYDATELLACEAE Watertufts family

These minute, annual and sometimes perennial, usually (semi-)aquatic, clumpforming herbs have a short upright branching rhizome and narrow linear leaves with a single vein and no petiole. Leaves are glabrous and lack sheaths and ligules, distinguishing them from grasses that may grow nearby. Inflorescences are capitate, with a sessile scape and bracts, resembling a single

Trithuria submersa, inflorescence, Western Australia (KD) [38]

inverted flower with centrally placed stamens (male flowers) surrounded by ovaries (female flowers) or inflorescences unisexual. Flowers are minute and reduced, without a perianth, and unisexual. Male flowers have a single stamen with a long slender filament and an anther that is fixed at its base. Female flowers are a single stalked ovary with a single style, beaded stigmas and a superior ovary. The fruit is a capsule, indehiscent or splitting into three valves. Seeds are extremely small and difficult to see. Distribution: These peculiar diminutive plants are found in microherb communities in seasonally wet sites in Australia, New Zealand and India.

it has now been shown that Hydatellaceae are most closely related to the water lilies and hence are placed with Cabombaceae and Nymphaeaceae in Nymphaeales. The unusual floral characteristic of ovaries surrounding the stamens, otherwise found only in Lacandonia (Triuridaceae), is now thought to be due to these “flowers” actually being heads composed of stamens and pistil-like structures that each represent reduced individual flowers. The genus Hydatella, traditionally separated on the basis of its indehiscent fruits and unisexual plants, is now merged with Trithuria, the name that has nomenclatural priority. Genera and species: Hydatellaceae consist of the single genus Trithuria with c. 12 species that are difficult to distinguish.

Phylogeny and evolution: These plants were first believed to be grass-like monocots (originally placed in Centrolepidaceae), but

Etymology: Hydatella is a diminutive of the ύδωρ (hydor), water; e.g. a small water plant.

Trithuria filamentosa, Lake Dobson, Mt Field National Park, Tasmania, Australia (CD) [38]

Trithuria bibracteata, Perth, Western Australia [38]

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NYMPHAEALES

THE ANA GRADE FAMILIES

Cabomba aquatica, Singapore Botanic Gardens [39]

Brasenia schreberi, Warm Lake, Valley County, Idaho, USA (CD) [39]

39. CABOMBACEAE Fanwort family

These f loating aquatic perennial herbs have horizontal rhizomes or stems that are freely rooting at the nodes with two vascular bundles and long internodes. Submerged parts of the plants can be covered in a gelatinous sheath. Leaves are simple, petiolate, opposite or seemingly alternate, the floating leaves peltate with entire or lobed margins and dichotomously branching venation. Submerged leaves are palmately dissected, the parts dichotomously branching with linear segments (in Cabomba), or

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Cabomba caroliniana, Royal Botanic Gardens, Kew, UK [39]

entire (in Brasenia). Flowers are emergent from the water, solitary, small, hexamerous and actinomorphic. Sepals and petals are usually three, the petals with nectaries in Cabomba. Stamens are three or six (to 36), with slender filaments. Ovaries are superior, the carpels three to 18, with a capitate (in Cabomba) or elongate (in Brasenia) stigma. Fruits are aggregate and contain one, two or three seeds. Distribution: These are widely distributed in the Americas from Canada to Argentina and in Sub-Saharan Africa, temperate and tropical East Asia and eastern Australia. Cabomba is frequently introduced and naturalised outside its native range. Phylogeny and evolution: Fossils assignable to Cabombaceae are known from Lower Cretaceous (115 Mya) sediments in Brazil. Brasenia and Cabomba diverged from each other c. 20 million years ago. The family was formerly more widespread than at

present, based on fossil evidence of Brasenia schreberi from Europe. Cabombaceae share many characters with Nymphaeaceae, especially seed anatomy of Brasenia and Nymphaeaceae. Genera and species: Cabombaceae consist of two genera with six species: Brasenia (1) and Cabomba (5). Uses: Water shield, Brasenia schreberi, is cultivated in East Asia as a vegetable called junsai (Japanese) or chun cai (Chinese). Submerged parts of this species are covered with a mucilaginous jelly, which has potential to control algal and bacterial growth in ponds. Cabomba species are common oxygenating plants in the aquarium trade from which they may naturalise. Etymology: Cabomba is derived from an Amerindian name for the plant, which came into Latin through Spanish. Its origins have not been traced.

NYMPHAEALES

THE ANA GRADE FAMILIES

40. NYMPHAEACEAE Waterlily family

These are annual and perennial aquatic herbs with vertical stems or creeping rhizomes with complex vascular tissues. Tubers are often present. Leaves are simple, alternate, spirally arranged, peltate or cordate, submerged, floating or emergent. Flowers are solitary, usually emergent and replace a leaf in the spiral arrangement, and they are often large, usually with some distinction between outer (four to six) sepals and inner (0–70) petals, usually numerous in many spiral whorls, decreasing in size towards the centre, becoming staminodial. Stamens are numerous (up to 200), not all fertile and sometimes grading into petals, filaments stout, often flattened or petaloid. The ovary is superior, with many (three to 35) whorled carpels that are laterally or completely fused. The fruit is a spongy berry with many large seeds. Distribution: This is a globally distributed family, but absent from glaciated or dry Nymphaea lotus, private garden, Irvine, California, USA [40]

regions.They are absent from New Zealand, southern Australia and southern South America.

of complex origin. Some Barclaya and Nymphaea species are popular aquarium plants.

Phylogeny and evolution: The family is believed to have been much more diverse in the past. It is believed that they dominated aquatic habitats. The family has been suggested to be 121 million years old, but crown group diversification is believed to have occurred in the Northern Hemisphere during the Tertiary. Wind-pollinated, apetalous Ondinea purpurea from Western Australia is possibly embedded in Nymphaea and is now known as Nymphaea ondinea. Euryale and Victoria are closely related and could be considered part of the same genus, pending further evidence.

The biggest and the smallest: Discovery of the largest waterlily, the annual Victoria amazonica from South America, caused a sensation. It was named for Queen Victoria, and after it was first cultivated in 19th century Britain the leaf venation served as an inspiration for a then novel form of glasshouse architecture, most famously the Crystal Palace in London. With a diameter of 2.65 m (area 5.51 m 2) this is the largest undivided leaf on the planet. The smallest waterlily, Nymphaea thermarum, was discovered in 1987 in a hotspring in Rwanda but was thought to be extinct in its native habitat. It also gained widespread notice when it was revived from seeds at the Royal Botanic Gardens, Kew, from where it has now been reintroduced into the wild and distributed to other botanical gardens around the world. Theft of some of the plants on display at Kew also gained public notoriety.

Genera and species: Currently Nymphaeaceae consist of five genera with c. 85 species: Barclaya (4), Euryale (1), Nuphar (17), Nymphaea (c. 60) and Victoria (2). Uses: Nuphar polysperma seeds were used as a flour substitute by some native American tribes, and seeds of Euryale ferox are used as food in East Asia. Rhizomes and seeds of some Nymphaea species can also be eaten. Flowers of some Nymphaea species were used as a narcotic by shamans and priests to induce ecstasies. Nymphaea and Nuphar are commonly grown as pond plants, with many hybrids and cultivars Victoria cruziana, Helsinki Botanical Garden, Finland [40]

Etymology: Nymphaea is derived from Greek mythological creatures called νύμφη (nymphe), nymphs, who were divine female spirits often associated with natural features, most often with the life-giving flow of springs. Nuphar polysepala, Royal Botanic Gardens, Kew, UK [40]

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AUSTROBAILEYALES

THE ANA GRADE FAMILIES

AUSTROBAILEYANAE AUSTROBAILEYALES Families 41–43 represent the order Austrobaileyales. This order is difficult to characterise in morphological terms, but the three families share the presence of tiglic acid (a skin and eye irritant), a compound not unique to this order, but setting it apart from the magnoliid orders. They are woody lianas and trees and have bisexual flowers with undefined numbers of parts. The order has an estimated age of c. 150–200 million years.

41. AUSTROBAILEYACEAE Austrobaileya family

The single species is a liana with simple, opposite, leathery, oblong to narrowly ovate leaves that are glabrous and produce essential oils in spherical oil glands. Leaf margins are entire, and petioles are short, without stipules. Flowers are bisexual, axillary or terminal, solitary, pendulous, c. 5 cm across and exude a smell of rotting fish to attract flies for pollination. There are 11–24 tepals

that are spirally arranged and spreading, the outer ones green and sepal-like, the inner ones cream with red or purple dots, variable in size and shape, but the innermost smallest. Flowers have six to 11 boat-shaped to flat, spotted stamens, with the anthers facing inward, embedded in the connective. There are nine to 16 flat, overlapping staminodes that surround the carpels. The ovary is superior, and carpels are eight to 13 and free with c. 6 mm long styles. The fruit is a stalked, ellipsoid, globose or pear-shaped berry that is 5–7 cm wide, fleshy in texture and orange-yellow in colour. The seeds are whitish and resemble a chestnut. Distribution: They are found in wet tropical rainforests between 380 and 1,100 m in northeastern Queensland (Australia). Phylogeny and evolution: The family is

Austrobaileya scandens, Queensland, Australia (CD) [41]

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suggested to have an age of c. 100–200 million years, depending on the analysis. There are no known fossils, but the pollen resembles that of fossil pollen Clavatipollenites from the Lower Cretaceous. Previously placed in Laurales as one of the most ‘primitive’ members, Austrobaileyaceae are instead the sister to the other families of Austrobaileyales (Schisandraceae and Trimeniaceae) in DNA-based analyses. Genera and species: Austrobaileyaceae include a single genus with one species: Austrobaileya scandens. Etymology: Austrobaileya is derived from Latin australis, southern, and two botanists called Bailey: British-Australian botanist Frederick Manson Bailey (1827–1915) and American wood anatomist Irving Widmer Bailey (1884–1967).

Austrobaileya scandens, habit, Royal Botanic Garden, Sydney, Australia [41]

AUSTROBAILEYALES

THE ANA GRADE FAMILIES

Trimenia neocaledonica, New Caledonia (JM) [42]

42. TRIMENIACEAE Bittervine family

Trimenia papuana, New Guinea (JM) [42]

round, fleshy berries. Seeds are hard, smooth or ridged.

43. SCHISANDRACEAE Star-anise family

Distribution: The family is restricted to eastern Malesia and northern Australasia from Sulawesi through New Guinea, the Solomon Islands, south to northern New South Wales (Australia) and from New Caledonia to Fiji in the Pacific.

These trees, shrubs and lianas have young shoots that are often hairy and accumulate aluminium in their tissues. Leaves are opposite, petiolate and without stipules. Leaf margins are entire or serrate, and the blade usually has translucent dots. Veins are pinnate and connected near the margin. Inf lorescences are axillary or terminal cymes or panicles. Flowers are unisexual or bisexual, small and wind-pollinated. Tepals soon fall off after opening of the flower, but they are spirally arranged, numerous (to 38), the outer ones swollen and somewhat peltate, the inner ones reduced and more membranaceous. Flowers have seven to 25 spirally arranged stamens with short filaments and a connective protruding apically. The ovary is superior, the carpel solitary (rarely two), but rudimentary in male flowers, the stigma sessile and tufted. Fruits are small,

Phylogeny and evolution: Trimenia was originally thought to belong to Cunoniaceae, and when segregated from that family it was first placed in Ternstroemiaceae (a family now included in Pentaphylacaceae). It was associated with Monimiaceae until 1950, when Trimenia was placed in its own family in Laurales. It has now been shown to form an isolated lineage in Austrobaileyales. A second genus, Piptocalyx, is now merged with Trimenia. Fossil seeds were found in 118 million year old deposits in Hokkaido, Japan. Genera and species: Trimeniaceae consist of the single genus, Trimenia, with eight species. Etymology: Trimenia is named for British botanist Henry Trimen (1843–1896), who was director of the Royal Botanical Garden Peradeniya (Sri Lanka, then Ceylon). The genus does not, however, occur in Sri Lanka.

These are small trees, shrubs and twining lianas, with some (Illicium) accumulating aluminium. Leaves are aromatic, alternate, spirally arranged, often in false whorls or clusters at the ends of branches, simple and glabrous. Leaf margins are entire, sinuate or denticulate, petiolate and without stipules. Flowers are solitary, paired or a few clustered in the leaf axils. Flowers are bisexual (Illicium) or unisexual, with five to 30 undifferentiated tepals in one to several whorls, the outer ones small and bract-like, the middle ones larger and more colourful, the inner ones often reduced and sometimes transitional to stamens. Stamens are four to 50 or more numerous, free to completely fused, the anthers basifixed with the connective linear and protruding at the tip, often swollen and broader. The ovary is superior with five to 100 or up to 300 carpels arranged in a single Plants of the World

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AUSTROBAILEYALES whorl (Illicium) or spirally arranged, free, but laterally compressed and attached at the base of the flower or forming a subglobose mass. Styles are short, slender or conical. The fruit is an aggregate of explosive follicles (Illicium) or fleshy berries (Kadsura, Schisandra). Seeds have a hard seed coat and a circular cap. Distribution: The family is disjunctly distributed from the southeastern USA and Mexico into the Antilles and in East Asia from the Russian Far East and Japan to Sri Lanka, Java and Sulawesi.

THE ANA GRADE FAMILIES

included in a broad Magnoliaceae, but were later segregated into two families (Illiciaceae and Schisandraceae), still in the order Magnoliales. With increasing knowledge of morphology and anatomy, the two families were then placed together in the order Illiciales. DNA analyses placed these families together in Austrobaileyales, in which they are treated as a single family. Illicium separated from the other two genera some 75–90 million years ago. Genera and species: Schisandraceae consist of three genera with 85 species: Illicium (44), Kadsura (16) and Schisandra (25).

Phylogeny and evolution: The family had a continuous distribution in the Northern Hemisphere in the Tertiary, but died out in Europe and western Asia during the ice ages. Members of this family were previously

Uses: Fruits of some Schisandra species are edible and can be made into jams and juices. The sour fruits of Kadsura scandens

Etymology: Schisandra is derived from the Greek σκίζει (skizei), to split, and άνδρες (andres), men, in reference to the wellseparated anther cells.

Illicium henryi, Caerhays Estate, England, UK [43]

Illicium anisatum, National Botanic Gardens, Glasnevin, Ireland [43]

Kadsura longipedunculata, female, Helsinki Botanical Garden, Finland [43]

Schisandra rubriflora, male flowers, Royal Botanic Gardens, Kew, UK [43]

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are consumed in Malesia. Unripe fruits of Illicium verum are harvested as star anise, a culinary spice, also used in teas, liqueurs and medicine (as an antiviral). Illicium was used in ancient China as an insecticidal fumigant. Oil from Illicium anisatum from Japan and Korea is used for perfume, but it is toxic and locally used as a fish poison. Fruits of this species are common in ornamental flower arrangements and potpourri mixes. They are toxic and easily confused with true star anise. Some species of Illicium, Kadsura and Schisandra are cultivated as ornamentals.

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Illicium simonsii, fruits, Royal Botanic Gardens, Kew, UK [43]

CANELLALES

MAGNOLIIDS CANELLALES Families 44 and 45 are placed in Canellales, an order that can be recognised by the chemical composition of the leaves. The bark of several species of this order were used for various purposes but most notably as a cure for scurvy. An age of 122–125 My is estimated for this order.

44. CANELLACEAE Canela-bark family

tube. The ovary is superior, unilocular with two to six carpels and a short fat style that bears a two- to six-lobed stigma. The fruit is a berry.

Genera and species: Canellaceae consist of six genera with 18–21 species: Canella (1), Capsicodendron (2), Cinnamodendron (8), Cinnamosma (3), Pleodendron (3) and Warburgia (1–4).

Distribution: The family is disjunctly distributed in the New World (Florida, Central America, West Indies and the Atlantic Forest in Brazil) and in the Old World (only eastern Africa and Madagascar).

Uses: Bark of Canella winterana is used as a condiment and stimulant, for flavouring tobacco, as a fish poison and for timber (as Bahama whitewood) in the Caribbean. The bark of Cinnamodendron corticosum is used as a spice in the West Indies. Leaves of Warburgia ugandensis are eaten in curries in East Africa, and its resin is used to preserve wood. Fragrant wood of the Madagascan endemic Cinnamosma fragrans is used for ceremonial purposes in India, threatening natural stands of this species.

These trees, sometimes shrubs, have aromatic, alternate, simple, entire, petiolate leaves without stipules. They are commonly dotted with glands. Inf lorescences are axillary or terminal panicles or racemes, but flowers are sometimes solitary. Flowers are bisexual and regular, with three thick, leathery sepals and five to 12 petals. Petals are in one or two whorls, or spirally arranged, usually free, fused in Cinnamosma. Six to 12 stamens are connate into a tube, and the anthers are fixed to the outside of this filament

Phylogeny and evolution: The structure of the seed coat resembles that in Winteraceae and Illicium, and the family was thus long thought to be related to these. However, their parietal placentation confused early botanists, and these anatomical characters putatively shared with Myristicaceae also complicated placement. Their chemical properties (drimane-type sesquiterpenoids) are similar to those in some Winteraceae, and this relationship has been corroborated by molecular analyses. The two families are now placed in the separate order Canellales.

Warburgia ugandensis, National Museums of Kenya, Nairobi [44]

Cinnamosma fragrans, Orangea Forest Reserve, Antsiranana, Madagascar (CD) [44]

Etymology: Canella is a diminutive form of the Greek κάννα (kanna), a reed or cane, originally of Irish Celtic origin. The Spanish name ‘canelo’ is often used for plants in Canellaceae and Winteraceae that produce Winter’s bark. Canella winterana in fruit, Pointe-aux-Châteaux, Guadeloupe [44]

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CANELLALES

Tasmannia insipida, Australian National Botanic Gardens, Canberra [45]

MAGNOLIIDS

Tasmannia lanceolata, private garden, Kingston upon Thames, Surrey, UK [45]

45. WINTERACEAE Winter’s bark family

These aromatic shrubs, trees and rarely lianas are usually terrestrial, but sometimes epiphytic. Leaves are simple, spirally arranged and petiolate, lacking stipules, and they are usually dotted with glands and often whitish beneath. Inflorescences are terminal or axillary and compound in dichasia or the flowers solitary. Flowers are bisexual or unisexual (in Drimys), actinomorphic or bisymmetric. Sepals are fused into a cup-shaped calyx, rupturing when a f lower opens. Petals are two to many, free or the outer ones fused and rupturing as well. Three to many stamens are free, the filaments short and thick, and the anthers opening with lateral slits, the connective usually not developed. The ovary is superior, and one to many carpels are free or fused. Stigmas are sessile and circular,

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Pseudowintera colorata, private garden, Kingston upon Thames, Surrey, UK [45]

linear or cup-shaped. Fruit is a dry or fleshy berry, sometimes with stone cells. Distribution: This is a family with a widespread but patchy distribution in the montane tropics and temperate rainforests in the Southern Hemisphere, occurring from southern North America to northern South America (Mexico to Ecuador and Venezuela), eastern Brazil, coastal Chile, the Juan Fernandez Islands, Madagascar, the Philippines, Borneo, Sulawesi, New Guinea, the Solomon Islands, New Caledonia, eastern Australia, Tasmania and New Zealand. Phylogeny and evolution: Takhtajania perrieri from Madagascar (sometimes placed in its own family because of its parietal placentation) possibly diverged from other Winteraceae c. 45 million years ago, although morphological analysis has shown it is similar to Pseudowintera. The age of the family is suggested to be c. 125 million years old. Fossils assigned to Winteraceae have been found outside the current range of the family, suggesting it was more widespread in the past and that the current distribution is relictual. Drimys in the broad sense is paraphyletic, and hence Tasmannia needs to be applied to the Old World taxa. The genus Zygogynum is still in need of taxonomic revision but includes Belliolum and Bubbia.

Drimys winteri in fruit, private garden, Kingston upon Thames, Surrey, UK [45]

Genera and species: Winteraceae consist of five genera with c. 65 species: Drimys (6), Pseudowintera (3), Takhtajania (1), Tasmannia (5) and Zygogynum (c. 50). Uses: Tasmannia lanceolata was used as pepper substitute by colonial Australians and after introduction to England became established as ‘pepperleaf’ in Cornish cuisine. Dorrigo pepper (Tasmannia stipitata) is used for the same purpose in Australia. Winter’s bark (Drimys winteri) was used as an effective remedy for scurvy in the 17th and 18th centuries, until vitamin C could be synthesised. Etymology: This family is named for John Winter, captain of the ship Elizabeth that accompanied Sir Francis Drake’s journey around the world in 1577–1580. Winter sent a boat ashore to search for medicinal herbs after sailing around Cape Horn, and he discovered Drimys bark, which became a remedy for scurvy for centuries afterwards. Drimys winteri is called ‘canelo’ in Spanish, and it was often confused or substituted by Canella winterana from the Caribbean, which can be used for similar purposes. Δριμύς (Drimys) means tart or pungent in Greek, referring to the bitter taste of the bark.

CANELLALES

MAGNOLIIDS

Hero of Labour — Armen Leonovich Takhtajan (1910–2009) In 1940, Soviet-Armenian botanist Armen Takhtajan published his classification scheme for flowering plants, emphasising phylogenetic relationships between plants. It took a decade before his classification became known to scientists in the west, and he began collaborating with Arthur Cronquist, whose system was influenced by him. Takhtajan’s system, last updated in 2009, divided flowering plants into the classes Magnoliopsida (dicots) and Liliopsida (monocots), which he subdivided into subclasses. Similar in general to the Cronquist system, those of Takhtajan had greater complexity with smaller orders and families, to allow families to be easily morphologically defined. It sometimes blurred the distinction between family and genus. During Soviet times the teaching of classical genetics could lead to imprisonment, but Takhtajan openly fought this, calling the government ignorant, and his correspondence with the West must have made him something of a maverick among Soviet scientists. He was fired in 1948 from his two botanical jobs in Armenia, but was hired by the more independent Leningrad University a year later, where he ran a lab that explicitly defied the doctrines of Lysenko. He even fired Lysenkoists in his laboratory. He was director of the Komarov Institute of Botany from 1976 to 1986. An endemic Winteraceae from Madagascar, Takhtajania perrieri, was named in his honour, a species that was only found once in 1909 and was thought to be extinct until it was rediscovered in 1994.

Zygogynum pomiferum, Tchamba River Valley, New Caledonia (CD) [45]

Photo of Armen Takhtajan (public domain)

Drimys granadensis, San Francisco Botanical Garden, California, USA [45]

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PIPERALES

MAGNOLIIDS

PIPERALES Families 46–50 form the order Piperales, a clade of plants with sympodial growth and usually heart-shaped leaves with secondary palmate venation, often with oil glands. Anthers are usually in threes, and the perianth is either strongly reduced or the petals are fused. The group is estimated to have originated 110–175 million years ago.

46. SAURURACEAE Lizard’s-tail family

perianth. The three, four or six stamens are free or fused with the ovary. The three to five carpels bear distinct styles. Flowers produce no nectar, and the fruit is fleshy, indehiscent or dehiscent. Distribution: This family is found in North America and temperate East Asia (China, Indochina, Japan, Taiwan, Luzon, Java), often in swamps or other wet habitats.

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Members of this family are perennial, aromatic herbs with stoloniferous rhizomes. They have simple alternate leaves with stipules that are fused with the petiole. Inf lorescences are terminal spikes with bracts that subtend the flowers, the lowest bracts sometimes enlarged and petal-like (the entire inflorescence then resembling a single flower). Flowers are small and lack a

Phylogeny and evolution: Saururaceae are closely related to Piperaceae, with which they share many characters. The family has an estimated age of 75–78 million years. A fossil from the Eocene was attributed to the genus Saururus, but it differs in stamen number. Morphologically intermediate, Zippelia has been moved back and forth between the two families, but it is now firmly placed in Piperaceae on the basis of

Anemopsis californica, Royal Botanic Gardens, Kew, UK [46]

Gymnotheca chinensis, Cuc Phuong, Vietnam (CD) [46]

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molecular studies. Differences between the families are in stem vasculature and placement of ovules. Genera and species: Saururaceae consist of four genera with six species; Anemopsis (1), Gymnotheca (2), Houttuynia (1) and Saururus (2). Uses: Houttuynia cordata is often cultivated as a garden plant, but can become invasive. Shoots and rhizomes of this species are eaten as a vegetable in China and have a fish-like flavour. Saururus chinensis is used in Chinese medicine. Anemopsis californica, Saururus cernuus and S. chinensis make attractive plants for large aquaria or ponds. Etymology: Saururus is derived from the Greek σαύρα (saura), a lizard, and ουρά (oura), a tail.

Saururus cernuus, private garden, Kingston upon Thames, Surrey, UK [46]

PIPERALES

MAGNOLIIDS

Distribution: A pantropical family extending into the temperate regions in the Himalayas, Japan, South Africa and New Zealand, but mainly restricted to tropical forests.

47. PIPERACEAE Pepper family

Phylogeny and evolution: Piperaceae are the largest magnoliid family. Closely related to Saururaceae they are estimated to be 70–100 million years old. Diversification was initiated in the Lower Cretaceous, but much of the current diversity dates from the Tertiary and later. Herbaceous Verhuellia from the Greater Antilles, previously treated as a synonym of Peperomia, is sister to the remaining Piperaceae. Zippelia and Manekia are sister to a clade including both Peperomia and Piper. Piper is now generally treated in its broad sense, including Arctottonia, Macropiper, Lepianthes, Ottonia, Pothomorphe and Trianaeopiper. Sarcorhachis is a synonym of Manekia.

worldwide), from which the dried unripe fruit gives black pepper and white pepper when the peppercorn skin is first removed. Piper longum (long pepper) is also used as a spice and medicinally. Peppercorns were the main reason for trade between Europe and the East Indies in the 16th and 17th centuries and sparked the Chinese and European colonisation of the region. Piper aduncum and P. angustifolium are similarly used as minor spices in the Neotropics. Piper betle is the leaf chewed with lime and betel nuts (Areca, Arecaceae). Piper cubeba (cubebs), P. clusii (African cubebs) and P. guineense (Benin pepper) are used medicinally and to flavour tobacco. The roots of Piper methysticum (kawa pepper) are fermented in several Pacific countries to make a narcotic sedative drink. Kawa-kawa (Piper excelsum) is used as an aphrodisiac in New Zealand. Many species and cultivars of Peperomia are popular houseplants, especially P. argyreia, P. caperata and P. velutina.

These are herbs, shrubs, trees and lianas, sometimes epiphytic, with stems that have swollen nodes. Leaves are simple, entire, alternate or opposite, often aromatic, succulent or membranaceous, petiolate, the petiole in some genera sheathing the stem, sometimes having a ligule-like structure or with a cap-like prophyll (a leaf protecting the leaf bud). Blades are often oblique at the base, often cordate, sometimes peltate. Venation is variable, often palmate, sometimes pinnate. Inflorescences are axillary, opposite the leaves or terminal. Flowers are formed on a spike or raceme that can be singular or several together organised into umbels or racemes. Flowers are tiny and lack a perianth, but each is subtended by a bract. Two (to six) stamens are free. There are two to five carpels, the ovary is superior and the fruit is a berry.

Uses: Many Piper species are used as spices, most notably P. nigrum from India, Sri Lanka and Madagascar (now cultivated

Etymology: Piper is derived from the Sanskrit pippali, a berry or peppercorn, which came via Middle Indic pippari into Greek πιπέρι (piperi) and Latin piper, the name of the spice.

Piper nigrum, Brooklyn Botanical Garden, New York, USA [47]

Zippelia begoniifolia, fruits, Cuc Phuong, Vietnam (CD) [47]

Peperomia fraseri, Royal Botanic Gardens, Kew, UK [47]

Manekia incurva, Guadeloupe [47]

Genera and species: Piperaceae consist of five genera with c. 3,700 species: Manekia (4), Peperomia (c. 1,600), Piper (c. 2,100), Verhuellia (3) and Zippelia (1).

Peperomia emarginella, Serra de Maranguape, Ceará, Brazil [47]

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PIPERALES

48. ARISTOLOCHIACEAE Birthwort family

These are lianas, small trees, shrubs and rhizomatous perennial herbs with alternate, mostly entire, occasionally palmately lobed leaves that are usually cordate at their base, or they are achlorophyllous, leafless, root parasites, with rhizome-like stems that have haustorial roots connecting to the host plant (Hydnoroideae). Leaves have petioles and no true stipules, but stipule-like prophylls may be present and are fused and ochrea-like in Lactoris. Flowers are solitary, axillary and often grow from old wood. In Lactoris there are two to four male, female or bisexual flowers from short condensed shoots (brachyblasts), and in Hydnoroideae the solitary flowers develop underground, emerging only when they open. The perianth is formed in one (or two) whorls, actinomorphic and trimerous,

Hydnora africana, South Africa (DK) [48]

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MAGNOLIIDS

irregularly lobed, or zygomorphic and all parts fused into various pipe-shaped or other tubular structures. Stamens are five, six or numerous (>40) and arranged in one to four whorls. Filaments are free or fused or entire stamens are fused with the style forming a structure called a gynostemium. The anthers open by lengthwise slits, or pollen is extruded in threads (Hydnoroideae). The three to six carpels are fused (but nearly free in Lactoris), and the ovary is inferior, sometimes superior. The style is three- to six-lobed or variously branched, or sessile and flat in Hydnoroideae. The fruit is usually a dehiscent (basketlike) capsule but sometimes follicular or a dry berry, or in Hydnoroideae a leathery, circumscissile indehiscent berry with many seeds and a strong smell.

southern South America (Prosopanche) and in savannas and arid parts of the southern Arabian Peninsula, Sub-Saharan Africa and Madagascar (Hydnora).

Distribution: The family is widespread in the Americas (Canada to Argentina), throughout mainland Europe and North Africa, Sub-Saharan Africa, Madagascar, Sri Lanka, temperate and tropical East Asia to New Guinea and northern Australia. Lactoridoideae are restricted to the Juan Fernández Archipelago (Chile), where they grow on wet or foggy slopes usually in the forest understory at 400–600 m elevation. Parasitic Hydnoroideae are found in semi-arid regions of Costa Rica, Peru and

Phylogeny and evolution: Aristolochiaceae were previously thought to be allied to Annonaceae on the basis of some superficial floral characters (e.g. floral parts in threes), but this proved to be incorrect. Molecular studies have shown that Aristolochiaceae are sister to Piperaceae and Saururaceae, and they are therefore included in Piperales. Hydnoraceae and Lactoridaceae are included in Aristolochiaceae because they make the latter paraphyletic. Fossil pollen of Lactoris was found in the Cretaceous of southern Africa dated c. 69 million years old, but fossil pollen is also known from South America, India, Australia and Antarctica, which means that it was a widespread Gondwanan element that became extinct on the continents. The remaining wild population of Lactoris fernandeziana is restricted to Robinson Crusoe Island (Más a Tierra) and consists of c. 1,000 plants with low genetic diversity. Hydnoraceae were originally described as a group of fungi, even though the floral nature of the reproductive organs was recognised

Aristolochia anguicida, capsule, near Turbaco, Bolívar, Colombia [48]

Aristolochia pilosa, Ecuador [48]

PIPERALES

MAGNOLIIDS

Asarum speciosum, Royal Horticultural Society Garden, Wisley, UK [48]

Saruma henryi, Royal Botanic Gardens, Kew, UK [48]

Thottea siliquosa, Royal Botanic Gardens, Kew, UK [48]

Aristolochia arborea, New York Botanical Garden, USA [48]

Asarum caudatum, northern California, USA [48]

from the start. Because of their parasitic nature, an association with Rafflesiaceae was also assumed, but DNA studies have shown that Hydnoraceae are embedded in Aristolochiaceae. This association is morphologically supported by the inferior ovary and general fusion of flower parts. Aristolochia is here treated in its broad sense, including Einomeia, Endotheca, Euglypha, Holostylis, Howardia, Isotrema and Pararistolochia, and has an estimated age of 104–122 million years. The Asarum–Saruma clade is of more recent origin, c. 36–44 million years old. Genera and species: Aristolochiaceae consist of seven genera with c. 500 species in four subfamilies: Asaroideae – Asarum (c. 100) and Saruma (1); Hydnoroideae – Hydnora (c. 6) and Prosopanche (4); Lactoridoideae – Lactoris (1); Aristolochioideae – Aristolochia (c. 360) and Thottea (30).

to make a dessert, but the fruit is astringent on its own. Plants are high in tannins and have been used for tanning and preserving fish nets, and infusions have been used as a face wash to treat acne. Birthwort (Aristolochia clematitis) was used as an abortifacient in former times, and Virginian snakeroot (Aristolochia serpentaria) has snake-like roots that were thought to be effective in the treatment of snake bites. Neither of these is, however, fit for purpose due to their ineffectiveness and toxicity. Wild ginger, Asarum canadense, has been used as an inferior ginger substitute in North America. Aristolochia petersiana is used as an arrow poison in East Africa. Some species, such as pipevine or Dutchman’s pipe (Aristolochia durior), calico flower (A. littoralis) and pelican flower (A. gigantea, A. grandiflora, A. ringens), several asarabacca or wild ginger (Asarum) species and ‘upright wild ginger’ (Saruma henryi) are occasionally grown as garden ornamentals.

consisting of only a flower and root-like stems. Hydnoroideae are the only known angiosperms with no leaves or scales of any sort. Hydnora africana and H. triceps parasitise succulent Euphorbia species (Euphorbiaceae) and H. johannis only parasitises roots of Acacia and Albizia (Fabaceae). Prosopanche americana and P. bonacinae grow on the roots of Prosopis and other legumes (Fabaceae). Little is known about H. esculenta from Madagascar, which has unisexual flowers and has not been collected since 1947. Heat production (thermogenesis) occurs in the flowers of Prosopanche and in some Hydnora, attracting beetles and flies for pollination. Some species are easily overlooked, especially H. triceps, which produces subterranean flowers and fruits. Due to their inconspicuous nature and seasonal appearance, Hydnoroideae remain poorly known plants.

Uses: The fruits of jackal food, Hydnora africana, are deliciously sweet when baked on a fire. Fruit pulp can be mixed with cream

Parasitism: Hydnoroideae (Hydnora and Prosopanche) are achlorophyllous root parasites that are extremely reduced,

Etymology: Aristolochia is derived from the Greek άριστος (aristos), best, and λοχεία (locheia), childbirth, because some Mediterranean species were applied to cure infections after childbirth or to induce labour.

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MAGNOLIIDS

MAGNOLIALES Families 49 to 54 form the order Magnoliales. These woody plants can be recognised by their often two-ranked or spirally arranged leaves. Their petals are whorled (or spirally arranged), and their medium-sized seeds have an irregular ruminate endosperm (like nutmeg).

49. MYRISTICACEAE Nutmeg family

exposing woody seeds that are usually covered in a lacy or entire, leathery or fleshy aril. Distribution: This is a pantropical family that are often canopy trees in rainforests.

These aromatic, often dioecious trees, sometimes shrubs, have red sap and red, long terminal buds. Leaves are simple, alternate, often oriented in a plane, shortly petiolate and without stipules. Leaf margins are entire, and hairs on the leaf surfaces and stems are usually branched or stellate. Inflorescences are panicles or fascicled racemes. Flowers are small, actinomorphic and funnel-, bell- or urn-shaped. Tepals are usually three, basally fused and often fleshy. Male flowers have two to 40 stamens with fused filaments. Female flowers have a single carpel, superior ovary and bilobed stigma. The fruit is a fleshy to woody capsule, usually splitting in half, Myristica fragrans, Singapore (MC) [49]

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Phylogeny and evolution: Myristicaceae clearly belong to core Magnoliales, which probably evolved >100 million years ago. They are well supported as sister to Degeneriaceae. Fossils from upper Cretaceous deposits in the Sahara are known, and Eocene fossil seeds are known from Europe, e.g. from the London clay. Diversification of modern lineages happened fairly recently, c. 15–20 million years ago. Genera and species: Myristicaceae include 21 genera with c. 520 species: Bicuiba (1), Brochoneura (3), Cephalosphaera (1), Coelocaryon (4), Compsoneura (c. 19), Doyleanthus (1), Endocomia (4), Gymnacranthera (7), Haematodendron (1), Horsfieldia (c. 100), Iryanthera (20), Knema (c. 90), Mauloutchia (10), Myristica (c. 170), Osteophloeum (1), Otoba (8), Paramyristica (1), Pycnanthus (4), Scyphocephalium (4), Staudtia (1) and Virola (c. 65). Virola surinamensis, fruit, French Guiana [49]

Uses: Myristica fragrans is a tree with apricotlike fruits in which nutmeg (the seed) and mace (the aril) are formed. This native of the Banda Islands in the Maluku Archipelago (Moluccas) in Indonesia was important in the 17th century spice trade, giving the name “Spice Islands” to this region. Ground nutmeg is used as a culinary spice but with excessive use is addictive, toxic and potentially hallucinogenic. Nutmeg oil is used medicinally and for flavouring tobacco and toothpaste. Bark of Iryanthera, Virola elongata and Osteophloeum platyspermum is used locally as a hallocinogen. Gymnacranthera, Horsfieldia and Knema seeds have oils that are used to make candles. Fat from Otoba seeds is used to make soap, and Virola sebifera contains oils that are suitable for candle and soap making. Caihuba, Virola surinamensis, produces an edible oil that is similar to cocoa butter. Horsfieldia iryaghedhi, Pycnanthus angolensis, Staudtia stipitata and Virola koschnyi produce fine timbers. Etymology: Myristica is derived from the Greek μύρων (myron), a balm or ointment, probably derived from a Semitic root m’rr, meaning bitter, a cognate with myrrh. Compsoneura excelsa in fruit, Los Mogos, Osa, Puntarenas, Costa Rica (CD) [49]

MAGNOLIALES

MAGNOLIIDS

Tuliptree, Liriodendron tulipifera, Royal Botanic Gardens, Kew, UK [50]

Magnolia macrophylla, fruit, Nichols Arboretum, Ann Arbor, Michigan, USA [50]

Magnolia ×soulangeana (a hybrid of M. denudata and M. liliiflora), Royal Botanic Gardens, Kew, UK [50]

Magnolia stellata, private garden, Kingston upon Thames, Surrey, UK [50]

Magnolia doltsopa, Royal Botanic Gardens, Melbourne, Australia [50]

Magnolia campbellii, Royal Botanic Gardens, Kew, UK [50]

50. MAGNOLIACEAE

dissemination of scent and attracting pollinators. The fruit is cone-like with free or fused follicles. In many species the carpels dehisce, and the pendent seeds exhibit a red aril.

flowers, Pachylarnax with few carpels but many ovules per carpel, and Talauma with fused carpels. These genera have, however, been found to be embedded in Magnolia sensu lato, expanding that genus to > 250 species, a number that is still growing. The two species of Liriodendron are well-supported as sister to Magnolia. Magnoliaceae are probably sister to the rest of Magnoliales.

Tuliptree family

Distribution: The family has a disjunct distribution in eastern North America, tropical America (Mexico to Brazil and Peru), southern India, Sri Lanka, the Himalayas and throughout temperate and tropical East Asia (Japan and Korea to New Guinea). These trees and shrubs have simple, entire and lobed, spirally arranged and petiolate leaves and stipules that enclose the bud and sheath the stem; these soon fall off leaving a scar. Stalked flowers are formed singly on the end of branches or short axillary shoots. Petals are free, six or more, spirally or whorled, sometimes differentiated into sepal-like outer petals and petal-like inner ones. Numerous stamens are free and spirally arranged, the filaments short or elongate, often flattened, and the anthers are elongate with the connective produced into a tip. Ovaries are superior, often stalked. Carpels are usually numerous, sometimes few, spirally arranged and free. Beetles are the most frequent pollinators, and some species create heat in their flowers (thermogenesis), increasing the

Phylogeny and evolution: The 98 million year old fossil flower Archaeanthus and fossil fruits of Lesqueria have been assigned to Magnoliaceae. Liriodendron in particular was widespread across the Northern Hemisphere during the late Cretaceous and Tertiary. Numerous now extinct lineages have been recorded from fossils. More modern representatives appeared in the late Miocene in Eurasia, especially when compared to North American extant taxa, which are considerably older. Several modern genera were described on the basis of deviating morphological characters not found in Magnolia sensu stricto; these include Elmerillia with sessile ovaries, Kmeria with unisexual flowers, Manglieta with four or more ovules per carpel, Michelia with axillary

Genera and species: This is now a family consisting of just two genera with c. 267 species: Liriodendron (2) and Magnolia (c. 265). Uses: Essential oils from Magnolia champaca are used for perfumery; its leaves are used to feed silk worms. Timber of Magnolia is used for boxes, matches, engraving, flooring, broom handles, traditional Japanese shoes etc. Wood of Liriodendron (whitewood) is used for furniture, shingles, latches and formerly canoes. Many species are highly valued ornamentals. Etymology: Magnolia was named in honour of French botanist Pierre Magnol (1638–1715), who was the first to publish plant families in an intrinsic ‘natural’ classification. Plants of the World

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51. DEGENERIACEAE Masiratu family

MAGNOLIIDS

Distribution: This family occurs only in Fiji. Phylogeny and evolution: Degeneriaceae are closely related to Myristaceae. Wood anatomy is similar to Eupomatiaceae, but the two families appear not to have an exclusive relationship. Genera and species: The single genus Degeneria has two species: D. roseiflora and D. vitiensis, which due to the removal of the natural barrier between the two populations by human activity, are forming hybrid swarms.

These glabrous aromatic trees have simple, alternate, spirally arranged, entire, petiolate leaves without stipules. Flowers are formed solitarily on axillary peduncles and are large and actinomorphic. Three sepals are free. The 12–25 petals are free, in several whorls (or are nearly spirally arranged) and reduced in size toward the centre, merging into the numerous, spirally arranged flattened stamens. The ovary is surrounded by staminodia that are spoon-shaped and longer than the stamens. The ovary is superior, with a single carpel that is not fully closed when the flower opens. Stigmatic surfaces flare outwards. Flowers are pollinated by beetles; the staminodes spread and protect the stamens upon opening, the staminodes closing over the ovary the following day, exposing the stamens and protecting the ovary. The fruit has orange or red fleshy seeds that are embedded in fleshy tissue. The fruit dehisces after falling from the tree, and seeds are distributed by fruit pigeons and parrots that eat the fleshy pulp. Degeneria vitiensis, Monasavu Dam, Viti Levu, Fiji (CD) [51]

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Etymology: Degeneria was named for American botanist Otto Degener (1899–1988), who discovered it in Fiji in 1942.

52. HIMANTANDRACEAE Pigeonberry-ash family

actinomorphic but lack a perianth. The flower bud is instead protected by two closed bracts that form a cap over the flower. Strap-like staminodes and stamens are numerous, and the stamens are placed between staminodes; inner staminodes and some stamens bear glands. The superior ovary has seven to 28 free carpels that fuse in fruit. The fruit is a drupe with several flat hard seeds; it is dispersed by fruit pigeons. Distribution: The family is only known from Sulawesi, the Moluccas, New Guinea and northeastern Queensland (Australia), where it can be found in rainforests. Phylogeny and evolution: The family is probably most closely related to Eupomatiaceae/Annonaceae, but this relationship is not well supported. Genera and species: Himantandraceae consist of the single genus Galbulimima with two species: G. baccata and G. belgraveana, sometimes considered to be a single polymorphic species. Uses: The aromatic bark and leaves are boiled and used as a hallucinogen by Papuans. The trees are also used for timber.

These aromatic trees have simple, alternate leaves that are organised in a plane (tworanked). Leaves are petiolate, entire and without stipules. Lower leaf surfaces and young twigs are covered with copper-coloured peltate scale-like hairs. Solitary, axillary flowers are Degeneria vitiensis with fruit cut open to show seeds, Monasavu Dam, Viti Levu, Fiji (CD) [51]

Etymology: Himantandra is derived from Greek ιμάς (himas), a strap or thong, and άνδρες (andres), men, in reference to the strap-shaped stamens: It is a later synonym of Galbulimima. Galbulimima baccata, Queensland, Australia (CD) [52]

MAGNOLIALES

MAGNOLIIDS

53. EUPOMATIACEAE Bolwarra family

superior ovary has numerous, fused, spirally arranged carpels without styles. The fruit is a fleshy berry.

54. ANNONACEAE Soursop family

Distribution: The family occurs in the tropical rainforests of eastern New Guinea and the temperate rainforests of eastern Australia (Queensland, New South Wales, Victoria).

These are shrubs with root tubers that rarely become small trees. Leaves are aromatic, simple, alternate, two-ranked, entire and petiolate and lack stipules. Flowers are formed in an axillary position or terminally on shoots, sometimes caulif lorous, actinomorphic, without a perianth, but flower buds are protected by one or two fused bracts that form a cap over the bud. Numerous stamens have short, broad filaments, and basifixed anthers open with slits. The inner staminodes are numerous, up to 80, and bear glands. The petal-like staminodes open fully when in the female stage, but cover the ovary when in the male stage of the flower. Stamens and staminodes are basally fused, the entire structure falling off after the male phase of the flower. The stigmas are pollinated by Elleschodes beetles. The fallen staminal structures are eaten by the same beetles, which in doing so bring pollen from the ground back up to new flowers. The Eupomatia laurina, Royal Botanic Gardens, Kew, UK [53]

Phylogeny and evolution: Eupomatiaceae are morphologically most similar to Himantandraceae, which share characters of the f lower structure, but molecular results place Eupomatiaceae as the sister of Annonaceae, with which they share vegetative characters, pollen with a circular aperture and syncarpous berries. The family is estimated to have an age of c. 100 million years.

Etymology: Eupomatia is derived from the Greek ευ (eu), well, and πωματιών (pomation), a cap, referring to the cap that covers the flower bud.

Large trees, shrubs and lianas with fibrous bark and zigzag branches are typical for this family. Simple, alternate, usually two-ranked (in a plane) leaves are usually petiolate and lack stipules. Flowers are terminal or axillary, solitary or in fascicles, sometimes caulif lorous or (rarely) sprouting from underground runners. Flowers are usually bisexual with open development: the perianth enlarges when the flower has already opened. Sepals are usually three and free for most of their length, but they are often basally fused. Three to six (rarely 12) petals are usually placed in two whorls, but sometimes in a single whorl. The numerous stamens are usually spirally arranged, sometimes whorled, and staminodes are present in some genera. Filaments are usually short and free, sometimes elongate and fused into a tube. The superior ovary has one to many carpels that are free or partly to completely fused, and styles are also free or fused. The fruit

Eupomatia laurina, Royal Botanic Gardens, Kew, UK [53]

Annona muricata, fruit, Helsinki Botanical Garden, Finland (MC) [54]

Genera and species: Eupomatiaceae consist of the single genus Eupomatia, which has three species. Uses: The sweet bolwarra fruit of Eupomatia laurina has a strong, aromatic flavour and is used in cooking, jams, desserts and beverages.

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Cananga odorata, Singapore [54]

Anaxagorea javanica (WA) [54]

Uvaria curtisii (WA) [54]

is composed of one to several free, stalked berries, arranged in an umbel, or a fleshy fused berry. Seeds are hard and usually glossy.

Genera and species: Annonaceae include 111 genera in four subfamilies with c. 2,300 species: Anaxagoreoideae – Anaxagorea (30); Ambavioideae – Ambavia (2), Cananga (2), Cleistopholis (4), Cyathocalyx (7), Drepananthus (26), Lettowianthus (1), Meiocarpidium (1), Mezzettia (3) and Tetrameranthus (6); Annonoideae – Afroguatteria (2), Annona (162), Anonidium (4), Artabotrys (102), Asimina (7), Asteranthe (3), Bocagea (2), Boutiquea (1), Cardiopetalum (3), Cleistochlamys (1), Cymbopetalum (27), Dasymaschalon (25), Desmos (24), Diclinanona (3), Dielsiothamnus (1), Disepalum (9), Duckeanthus (1), Duguetia (93), Exellia (1), Fissistigma (48), Friesodielsia (51), Froesiodendron (3), Fusaea (2), Gilbertiella (1), Goniothalamus (136), Guatteria (210), Hexalobus (5), Hornschuchia (10), Isolona (20), Letestudoxa (3), Melodorum (10), Mischogyne (2), Mitrella (8), Mkilua (1), Monanthotaxis (56), Monocyclanthus (1), Monodora (16), Neostenanthera (4), Ophrypetalum (1), Porcelia (7), Pseudartabotrys (1), Pyramidanthe (1), Sanrafaelia (1), Schefferomitra (1), Sphaerocoryne (3), Toussaintia (4), Trigynaea (12), Uvaria (190), Uvariastrum (8), Uvariodendron (15), Uvariopsis (16) and Xylopia (157); Malmeoideae – Alphonsea (25), Annickia (8), Bocageopsis (4), Crematosperma (29), Dendrokingstonia (2), Desmopsis (14), Ephedranthus (6), Fenerivia (10), Fitzalania (2), Greenwayodendron (2), Haplostichanthus

(11), Huberantha (27), Klarobelia (12), Maasia (6), Malmea (6), Marsypopetalum (6), Meiogyne (16), Miliusa (52), Mitrephora (47), Monocarpia (1), Monoon (56), Mosannona (14), Mwasumbia (1), Neo-uvaria (5), Onychopetalum (2), Orophea (50), Oxandra (29), Phaeanthus (10), Phoenicanthus (2), Piptostigma (14), Platymitra (2), Polyalthia (120), Polyceratocarpus (8), Popowia (26), Pseudephedranthus (1), Pseudomalmea (4), Pseudoxandra (23), Pseuduvaria (58), Ruizodendron (1), Sageraea (9), Sapranthus (6), Sirdavidia (1), Stelechocarpus (3), Stenanona (14), Tridimeris (1), Trivalvaria (4), Unonopsis (48), Wangia (1) and Winitia (2).

Distribution: Annonaceae have a pantropical distribution, with a single temperate genus (Asimina) in the eastern USA. Phylogeny and evolution: Fossil Annonaceae are known from the Late Cretaceous. Annonaceae diverged from Eupomatiaceae c. 100 million years ago. Anaxagorea, often placed in a separate subfamily, diverged from the remaining Annonaceae c. 88 million years ago, after which the remaining Annonaceae diversified 70–85 million years ago, with the late Cretaceous fossil flower Futabanthus (c. 89 million years old) being one of the early members of modern Annonaceae. Recently some genus-level reorganisation was needed after DNA studies. The family is now divided into four subfamilies: Anaxagoreoideae, Ambavioideae, Annonoideae and Malmeoideae, the vast majority of genera belonging to the last two subfamilies. Rollinia is now included in Annona, and Haplostichanthus is now united with Polyalthia. Oxandra and Polyalthia were found to be polyphyletic, resulting in the recognition of the genera Hubera and Monoon, the latter including Enicosanthum.

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Christenhusz, Fay & Chase

Uses: Several species are cultivated for their fruits, especially Annona muricata (soursop, custard apple or guanábana), A. cherimola (cherimoya), A. squamosa (sweetsop or custard apple) and Asimina triloba (pawpaw or Michigan banana). Guanábana seeds have insecticidal properties. The fragrant flowers of ylang-ylang (Cananga odorata) are used in the perfume industry. A columnar form of Monoon longifolium (previously known as Polyalthia) is commonly planted as a street tree in the tropics. Etymology: Annona is derived from anona, the Taino (Amerindian) name for the fruit, not based on the Latin annus, year, as is sometimes claimed.

LAURALES

MAGNOLIIDS

LAURALES Families 55 to 61 comprise Laurales. These woody shrubs and trees are recognised by their opposite often aromatic leaves, flowers with a hypanthium and more or less spirally arranged perianth, concave receptacle and endotestal seeds. The age of divergence of this order has been estimated at 108–114 million years.

55. CALYCANTHACEAE Spicebush family

staminodes are situated on the inside edge of the flower base. The semi-inferior ovary holds one to many carpels that are fused if there is more than one. The fruit is a dry indehiscent capsule formed by the flower base and contains one to several glossy brown seeds. Distribution: Calycanthaceae are disjunctly distributed between North America, temperate East Asia and Australia (northern Queensland).

These are aromatic shrubs and trees with scaly buds. Leaves are simple, opposite, entire and petiolate with ethereal oil cells and no stipules. Flowers are solitary or in few-flowered clusters. The perianth consists of 15–40, spirally arranged tepals that are attached to the outside of the cup-shaped flower base (receptacle). Five to 30 stamens are inserted spirally on the rim of the flower base, and their filaments are short or the anthers sessile. The numerous

Phylogeny and evolution: Calycanthaceae are sister to all other Laurales and are believed to have diverged about 114–171 million years ago. Several fossils are thought to share characters with Calycanthaceae, but most are disputed because they also show characters not now known to occur in the family. Sinocalycanthus is morphologically intermediate between Calycanthus and Chimonanthus, but is now considered a synonym of the former genus,

Chimonanthus praecox, Royal Botanic Gardens, Kew, UK [55]

Calycanthus chinensis, Mt Tianmu, Anhui, China [55]

with which it forms hybrids in cultivation that are of ornamental value. Genera and species: Calycanthaceae include three genera with ten species: Calycanthus (3), Chimonanthus (6) and Idiospermum (1). Uses: Carolina allspice, Calycanthus floridus, was formerly used as a cinnamon substitute. Flowers of Chimonanthus praecox are used in East Asia to scent linen. Calycanthus oil is used in high-quality perfumes. Idiospermum australiense is an endangered rainforest tree from Queensland that was thought to be extinct but is now being cultivated in botanical gardens in Australia to ensure its survival. Calycanthus and Chimonanthus are frequently cultivated in gardens for their scented flowers. Etymology: Calycanthus is derived from Greek καλυξ (kalyx), crown or calyx, and άνθος (anthos), flower. Idiospermum australiense, Royal Botanic Gardens, Sydney, Australia [55]

Calycanthus floridus, North Carolina, USA [55]

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LAURALES

56. SIPARUNACEAE Fevertree family

MAGNOLIIDS

male flowers in search of a nesting site and in doing so transfer the pollen. Several drupes form together and are enclosed in a large false fruit that splits at maturity.

57. GOMORTEGACEAE Keule family

Distribution: This is a family from tropical America and tropical West Africa.

These are monoecious and dioecious shrubs, small trees and, sometimes, lianas. Leaves are simple, opposite and entire; the second leaf of a pair is represented by a midrib only in Glossocalyx, but is normally developed in Siparuna. Flowers are axillary and unisexual. Four to six tepals are fused, sepal-like or obscure (appearing to be absent). Male flowers have two to many stamens inside a ball-, cup- or urn-shaped flower base, which sometimes protrude from the valved sepals at anthesis. Female flowers have few to many, often sunken carpels, with the styles free at the tip and connate basally, usually protruding through the flower opening. Flowers are pollinated by gall midges, which lay their eggs inside male flowers, but visit both female and Siparuna guianensis, New York Botanical Garden, USA [56]

Siparuna nicaraguensis, Guatemala (MV) [56]

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Phylogeny and evolution: Previously Sipar unaceae were included in the polyphyletic Monimiaceae, which are now better understood, and the genera have been reorganised. Siparunaceae are sister to Gomortegaceae+Atherospermataceae. The oldest fossils of Siparunaceae are c. 88 million years old and have been recorded from deposits in the Late Cretaceous of Antarctica and the Eocene of Germany, so the family must have been widespread previously. Dioecy evolved several times from monoecy in Siparuna.

Etymology: Siparuna is derived from an Amerindian name (possibly Wayampi) for the plant in French Guiana.

These grey-barked, aromatic, evergreen trees have simple, opposite, entire, leathery, petiolate leaves without stipules. Inf lorescences are terminal or axillary racemes with a terminal flower. Peduncles are hairy and bear small bracts that drop off after opening of the flowers. Seven to ten spirally arranged tepals are hairy and intergrade with the seven to 13 stamens, the outer ones resembling petals, often with partly developed anthers; fully developed stamens have filaments with two stalked glands on each side. The one to three staminodes are sterile and often petal-like. The inferior ovary has two or three carpels, a short style and two or three stigmas. The fruit is a yellow, singleseeded drupe, similar to an avocado.

Glossocalyx brevipes, Monts de Cristal, Gabon (CD) [56]

Gomortega keule, Adelaide Botanic Garden, South Australia [57]

Genera and species: Siparunaceae consist of two genera and 78 species: Glossocalyx (4) in Africa and Siparuna (74) in the Neotropics.

Gomortega keule, Adelaide Botanic Garden, South Australia [57]

LAURALES

MAGNOLIIDS

Atherosperma moschatum, Australia (PW) [58]

Gomortega keule, habit, Adelaide Botanic Garden, South Australia [57]

Nemuaron vieillardii, Tchamba River Valley, New Caledonia (CD) [58]

Laurelia sempervirens, Royal Botanic Gardens, Kew, UK [58]

Distribution: Coastal central Chile (regions VII and VIII), where the single species grows in forests on the south-facing slopes of humid ravines. It is endangered due to overharvesting and clearing of forests for agriculture.

58. ATHEROSPERMATACEAE

Distribution: Atherospermataceae occur disjunctly around the southern Pacific: southern South America (Chile), New Guinea, eastern Australia, Tasmania, New Caledonia and New Zealand.

Southern-sassafras family

Phylogeny and evolution: Cretaceous fossils are known from the Antarctic Peninsula and taking their current distribution into account, the family may have a Gondwanan origin. The oldest fossils are 88 million years old. The family is sister to Gomortegaceae.

Phylogeny and evolution: Morphologically clearly part of Laurales, Gomortegaceae are sister to Atherospermataceae. Genera and species: Gomortegaceae consist of the single species, Gomortega keule. Uses: The sweet fruit of keule is edible and considered a delicacy in Chile, but it is rarely found outside that country. It is used to produce marmalade and liqueur. The easily worked wood was previously used to make high-quality furniture, but harvesting wood of this rare, slowly growing species is not sustainable. Etymology: Gomortega was named for Spanish botanist and physician Casimiro Gómez Ortega (1741–1818), first director of the Real Jardin Botánico de Madrid. Keule or queule is the Mapuche name for the plant.

These are aromatic trees and shrubs with T-shaped or simple hairs and simple, opposite, coarsely serrate leaves without stipules. Lateral veins of the leaves join to form a vein along the margin. Flowers are solitary and axillary or aggregated into cymose inflorescences. Flowers are bisexual or unisexual, more or less actinomorphic, with four to 20 tepals in two whorls; the perianth is poorly differentiated or clearly separable into sepals and petals. The 12 to numerous stamens and staminodes are fused, each with a pair of glands at the base. The superior or half-inferior ovary (the carpels then sunk in the flower base) have three to numerous free carpels with a lateral or gynobasic style. The fruit is an aggregate of indehiscent plumed achenes.

Genera and species: Atherospermataceae consist of seven similar genera and 20 species: Atherosperma (1), Daphnandra (9), Doryphora (2), Dryadodaphne (4), Laurelia (2), Laureliopsis (1) and Nemuaron (1). Uses: Laurelia sempervirens fruits are used as a spice in South America. The strongly scented bark of Atherosperma moschatum is made into an infusion for tea in Tasmania, and the fragrant wood of Doryphora sassafras is used in Australia to repel insects. Etymology: Atherosperma is derived from the Greek άθερ (ather), awn and σπέρμα (sperma), seed, referring to the plumed achenes.

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LAURALES

59. HERNANDIACEAE Lantern-tree family

These trees, shrubs and lianas are rich in essential oils. When they are lianas, they climb using petioles (Illigera) or specialised stem hooks (Sparattanthelium). Leaves are simple or palmately compound, entire or palmately lobed (Illigera), peltate or cordate at the base, palmately veined, petiolate, without stipules, sometimes with secretory cavities or cystoliths in the leaf blades.

MAGNOLIIDS

Flowers, formed in terminal cymes, are bisexual or unisexual and actinomorphic. Six to ten free tepals form two whorls and are similar in shape and size. Three to five stamens have filaments that usually bear a pair of glands at the base or sides. The inferior ovary has a single carpel with a terminal style and an apical stigma. Fruits are indehiscent dry samaras or enclosed in an inflated balloon formed from fused bracteoles, which are dispersed by sea currents, wind or animals. Distribution: The family has a pantropical distribution, often assocated with coastal habitats or rainforests. Phylogeny and evolution: Hernandiaceae are divided into two subfamilies that diverged c. 76 million years ago, one (Hernandioideae) morphologically more

Hernandia nymphaeifolia, fruit, Seychelles [59]

Gyrocarpus americanus in fruit, Cintalapa, Las Minas, Chiapas, Mexico (CD) [59]

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similar to Monimiaceae and the other (Gyrocarpoideae) morphologically similar to Lauraceae, but pollen morphology is shared by both subfamilies. In molecular studies, Hernandiaceae are sister to Monimiaceae, and this pair are sister to Lauraceae. Genera and species: Hernandiaceae include five genera and 58 species: Gyrocarpus (3), Hazomalania (1), Hernandia (23), Illigera (19) and Sparattanthelium (13). Uses: Gyrocarpus produces a light timber. Hernandia nymphaeifolia is sometimes used as a street tree in cities in the tropics. Etymology: Hernandia was named for Spanish naturalist and court physician Francisco Hernández de Toledo (1514–1587), who took part in the first scientific mission to the Americas.

Hernandia nymphaeifolia, Seychelles [59]

Sparattanthelium botocudorum, Recife, Brazil [59]

LAURALES

MAGNOLIIDS

60. MONIMIACEAE Boldo family

These hermaphroditic, monoecious and dioecious, aromatic trees, shrubs and lianas have simple, opposite, entire or serrate, gland-dotted, pinnately veined, petiolate leaves without stipules. Bisexual or unisexual flowers are solitary or produced in cymes or racemes; they are actinomorphic or somewhat oblique. The flower base (receptacle) is hollow, the perianth has distinct tepals in two whorls or the perianth is entirely absent. The outer tepal whorl (when present) has four parts that are sepal-like and sometimes fused and cap-like, the inner whorl (when present) has seven to 20 (or more) free petal-like parts. Ten to numerous free stamens form one or two whorls, and these surround one to 50

staminodes. Filaments do not bear glands. The superior to partly inferior ovary has (one to) three to many free carpels, with sessile, apical stigmas. Flowers are pollinated by small insects such as thrips that lay their eggs in the fig-like flowers or are visited by other insects when the flowers are of a more conventional structure. The fruit has indehiscent achenes aggregated in heads fusing to produce a false fruit formed by the fleshy flower base that splits irregularly when ripe. Distribution: This pantropical family extends into the temperate zones in southern South America (Chile), eastern Australia, Tasmania and New Zealand. Phylogeny and evolution: The family used to include Atherospermataceae and Siparunaceae, which are morphologically similar but make the family polyphyletic with regard to Hernandiaceae and Gomortegaceae. Monimiaceae are sister to Hernandiaceae and are estimated to have an age of c. 90 million years. Fossil wood dated at 85 million years has been used to calibrate phylogenetic trees of this clade. The family is thought to be of eastern Gondwanan origin, but the current

Tambourissa purpurea, Royal Botanic Gardens, Kew, UK [60]

distribution has been achieved by several long-distance dispersal events. Genera and species: Monimiaceae include 24 genera and c. 275 species: Austromatthaea (1), Decarydendron (3), Ephippiandra (6), Hedycarya (11), Hennecartia (3), Hortonia (3), Kairoa (2), Kibara (45), Kibaropsis (1), Lauterbachia (1), Levieria (7), Macropeplus (1), Macrotorus (1), Matthaea (6), Mollinedia (90), Monimia (3), Palmeria (15), Parakibara (1), Peumus (1), Steganthera (c. 17), Tambourissa (45), Tetrasynandra (3), Wilkiea (6) and Xymalos (1). Uses: The leaves of boldo, Peumus boldus, can be made into a digestive hot drink, its fruits are also edible and its bark is used as a dye in Chile. Hedycarya angustifolia is sometimes used for furniture making in Australia. Boldo and lemonwood (Xymalos monospora) produce good timbers. Etymology: Monimia is derived from Μόνιμη (Monime), the wife of King Mithridates VI of ancient Pontus (died 63 BC). In modern Greek the word is used to mean ‘permanent’.

Peumus boldus, Adelaide Botanic Garden, South Australia [60]

Hedycarya angustifolia, Australian National Botanic Gardens, Canberra [60]

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LAURALES

61. LAURACEAE Bay-laurel family

This family consists of aromatic trees and shrubs, except Cassytha, which are parasitic twiners with scale-like leaves and stem suckers (haustoria). In all genera other than Cassytha, leaves are simple, alternate, opposite or whorled, entire or lobed, petiolate and without stipules. Inflorescences are mostly axillary panicles or false umbels, rarely a head or flowers solitary. Flowers are actinomorphic, bisexual or unisexual, trimerous. The small, flat to enlarged and urn-shaped flower base is free from the ovary. Tepals are formed in two equal whorls, usually undifferentiated. The many stamens are usually in four whorls, the innermost consisting of sterile staminodia or this whorl absent, usually one whorl bearing glands at the base of the filaments. The ovary is usually superior, rarely inferior (Hypodaphnis) and has a single carpel. The fruit is a one-seeded berry or drupe, usually with the flower base and flower stalk enlarged to form a leathery or woody cup surrounding the base of the fruit, rarely completely enclosing it. Distribution: This widespread temperate and tropical family occurs in the Americas (eastern Canada and California to Argentina and southern Chile), Macaronesia, the Mediterranean, Sub-Saharan Africa, Madagascar, the Mascarenes, Asia (India to Korea and Japan), Australia (including Tasmania), New Zealand and many Pacific islands east to Hawaii. Phylogeny and evolution: Lauraceae were a dominant family in Middle Cretaceous floras, and many Triassic fossils from North America can be assigned to modern genera, especially Sassafras, which has a remarkably

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variable leaf shape. Fossilised Sassafras-like wood has, however, been found in Antarctica, implying a broader distribution of this genus. The oldest fossils are c. 100 million years old. Despite the generally large seeds of this family, most clades in recent phylogenetic analyses include species from different continents, implying frequent long-distance dispersal across oceans. The Macaronesian species, for instance, are closely related to American species, but not to each other, implying two independent dispersal events from the Americas to the Canary Islands, where the family is emblematic due to the ancient ‘laurisilva’ forests, believed to be similar to European forests before the ice ages. Hypodaphnis, with its inferior ovary, is sister to the rest of the family, followed by parasitic Cassytha on a long branch, although some analyses also show Cassytha to be sister to a clade comprised of Caryodaphnopsis and Neocinnamomum; all analyses place the parasitic Cassytha firmly in Lauraceae. The species-rich genus Ocotea diversified recently. Phylogeny and classification at the generic level are not yet fully understood, and it is likely that Actinodaphne, Lindera, Litsea, Persea and Phoebe will need to be redefined when better molecular sampling of these genera is available. The family is sister to Hernandiaceae/Monimiaceae. Genera and species: Lauraceae include c. 45 genera and about 2,850 species: Actinodaphne (100), Aiouea (21), Alseodaphne (50), Aniba (41), Apollonias (2), Aspidostemon (c. 15), Beilschmiedia (c. 250), Caryodaphnopsis (16), Cassytha (20), Cinnadenia (2), Cinnamomum (c. 350), Cryptocarya (c. 200), Dehaasia (35), Dicypellium (2), Dodecadenia (1), Endiandra (106), Endlicheria (40), Eusideroxylon (2), Hypodaphnis (1), Kubitzkia (1), Laurus (2), Licaria (40), Lindera (100), Litsea (c. 400), Mezilaurus (20), Micropora (1), Nectandra (120), Neocinnamomum (6), Neolitsea (100), Nothaphoebe (40), Ocotea (c. 350), Paraia (1), Parasassafras (2), Persea (c. 200), Phoebe (100), Phyllostemonodaphne (1), Pleurothyrium (45), Potameia (30), Povedadaphne (1), Rhodostemonodaphne (20),

Sassafras (3), Sinosassafras (1), Umbellularia (1), Urbanodendron (1), Williamodendron (4) and Yasunia (2). Uses: Avocado, Persea americana, is a major crop around the world known for its tasty, nutritious fruits that are rich in vitamins, potassium and oils. Avocado oil is used for cooking (high in monounsaturated fats), cosmetics and lubrication. Enkala (Litsea garciae) is a minor fruit crop in tropical East Asia. Litsea cubeba has an edible fruit, and its leaves repel insects, although they can be used as food for silkworms. The bark of several Cinnamomum species is used as a culinary spice, predominantly true cinnamon (C. verum), cassia bark or bastard cinnamon (C. aromaticum), korintji cinnamon (C. burmannii) and kulilawan (C. culitlawan). Cinnamon oil is also used in incense, toothpaste and medicines. Camphor (C. camphora) is of economic importance as an insecticide, but the camphor used in mothballs is usually chemically synthesised. Cinnamaldehyde, the compound that gives cinnamon its flavour and odour, has numerous applications. Apart from flavouring beverages, food and chewing gum, it has been used as an effective fungicide in agriculture. It has also proved to be antimicrobial and has anticarcinogenic properties. It has even been used to prevent corrosion of steel. Bay (Laurus nobilis) is a well-known culinary herb used to flavour soups and stews. The plant was important in Classical Greek and Roman times when it was a symbol of victory, made into a wreath offered to gods, emperors and Olympic athletes. Cryptocarya moschata yields a spice known as Brazilian nutmeg. The bark and seeds of Madagascar cloves (Cryptocarya agathophylla) are used to flavour rum. Essential oils from the bark and seeds of several Cryptocarya species are extracted. Dicypellium caryophyllaceum bark produces a spice similar to cloves, but the species is not cultivated and now threatened with extinction due to overharvesting from the wild. Seeds of Actinodaphne hookeri, Licaria pulcheri (puchurin nut) and Nectandra pichurim (purchury bean) are used for (medicinal) oil. Oils of Nectandra

LAURALES

MAGNOLIIDS

Lindera umbellata var. membranacea Royal Botanic Gardens, Kew, UK [61]

Umbellularia californica, fruit, California, USA [61]

Cassytha filiformis, Singapore [61]

Cinnamomum verum, Helsinki Botanical Garden, Finland [61]

Persea americana, fruit, Kenya [61]

Cassytha filiformis, St Petersburg, Florida, USA [61]

Laurus nobilis, Hampton Court, England, UK [61]

elaiophora and Neolitsea sericea can be used for burning and making soap. Brazilian sassafras (Aniba roseodora) produces the famed bois-de-rose oil used in the perfume industry. Fragrant wood of Orinoco sassafras (Ocotea cymbarum) and camphorwood (O. usambarensis) are similarly exploited. Because of excessive harvesting from the wild, these species are now threatened with extinction. Spicebush (Lindera benzoin) and sassafras (Sassafras albidum) were used as teas by North American natives. Traditionally, they were used as a flavouring, but they are now banned because they have been linked to kidney cancer. Sassafras oil is still used as

a topical treatment for insect bites. Cassytha filifomis yields a brown dye. Many species produce good timber, and Persea nanmu is prized as a coffin wood in China. Endiandra palmerstonii wood is especially valued due to its varied colouration.

stems. They attach to the host via sucker-like cups that are formed singly or in files along the stem of the parasite and dissolve the underlying stem or leaf tissues of the host plant. They do make roots initially, but these die when a connection with the host plant is established.

Parasitism: The only parasitic genus is Cassytha, which has its centre of diversity in Australia where they are commonly referred to as dodder-laurels. They can become pests, especially in urban parks where vegetation remains free from fire; Cassytha species are fire-sensitive. The leafless stems coil tightly around the stems of a wide range of host plants, often forming dense mats of intertwining wiry

Etymology: Laurus is Latin for a bay tree. Crowns of bay branches were an emblem of victory or distinction, hence the term ‘laureate’. The word is a cognate with Δάφνη (Daphne), a Greek mythological nymph who flirted with the god Phoebus (hence the genus Phoebe), but when pursued, her mother Gaia changed her into a laurel bush.

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CHLORANTHALES

UNPLACED

CHLORANTHALES This order has long been controversial in terms of their relationships, with most authorities stating that their affinities are uncertain. They do not necessarily belong to the magnoliids, but in several molecular studies they are sister to them.

62. CHLORANTHACEAE Pearl-orchid family

This family is composed of aromatic herbs, shrubs and soft-wooded trees, with or sometimes without vessels in their stems. Their simple, opposite, petiolate leaves have dentate margins. Petioles are fused across the node at the base, and the stipules are fused with the petioles, sheathing the swollen stem nodes. The inflorescence is a terminal or axillary head, spike, thyrse or raceme that is usually compound. Flowers are usually subtended by up to three bracts and are unisexual (in Ascarina and Hedyosmum) or bisexual with the stamens adnate to one side of the ovary (in Chloranthus and Sarcandra). Three fused tepals are typical but sometimes absent. Male flowers have one to five stamens, with the connective

Ascarina rubricaulis, Plateau de Dogny, New Caledonia (CD) [62]

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often enlarged. Female and bisexual flowers have inferior ovaries or are naked. Bisexual flowers have one to three fused anthers that are connected to and sometimes envelop the ovary. The ovary has a single carpel with a short style and (nearly) sessile stigma. The fruit is a berry or drupe, and the seeds have a hard seed coat. Distribution: This family occurs in tropical America, Madagascar, tropical and temperate East Asia, New Guinea, New Caledonia, New Zealand and on Pacific islands. Chloranthaceae are absent from mainland Africa. Phylogeny and evolution: Affinities of this family have been much disputed in the past. They were placed in Piperales, Magnoliales and Laurales, and even an association with Trimeniaceae or the monocots has been suggested. Molecular analyses (based mostly on plastid DNA) have placed them in an isolated position as sister to the magnoliids, and they are now placed in their own order Chloranthales. Fossils of this family are common, world-wide in distribution and morphologically diverse. The spiraperturate pollen is unique to the family and easily recognised in pollen deposits; the oldest

Chloranthus holostegius, Kunming Botanical Garden, China [62]

pollen fossils are from the Early Cretaceous, c. 125 million years old, and can be attributed to Hedyosmum. All modern genera diverged c. 90 million years ago, and diversification of Hedyosmum in the New World is estimated to have occurred c. 35–45 million years ago, coinciding with the uplift of the Andes. Chloranthaceae were, however, more diverse in earlier times and show a pattern of extinction and recent radiation. Genera and species: Chloranthaceae include four genera with 77 species: Ascarina (12), Chloranthus (18), Hedyosmum (45) and Sarcandra (2). Uses: The leaves of Chloranthus glabrus and C. spicatus are used to scent tea or added to tea for bulk. Chloranthus erectus was used in Java instead of tea, before tea (Camellia) was introduced there. The leaves have detoxifying and anti-inflammatory properties, and several species of Chloranthus are used in Asian traditional medicine. Etymology: Chloranthus is derived from Greek χλωρός (chloros), green, and άνθος (anthos), flower.

Hedyosmum mexicanum, male, Mexico [62]

Hedyosmum mexicanum, female, Guatemala [62]

MONOCOTS Monocots or monocotyledons were first recognised as a natural group of plants by John Ray (Methodus Plantarum Nova, 1682), who was the first to emphasise the differences between the number of seed leaves (cotyledons, one versus two) among flowering plants. The dicots or dicotyledons (usually) have two seed leaves, but they are not monophyletic because they include the ANA grade magnoliids and eudicots; basically dicots include all angiosperms minus the monocots. The monocots are a clade and are recognised by the APG classification (2009), but there is no single or set of morphological characters that are guaranteed to inform an observer that a plant is a monocot. Having made this

point, most monocots have leaves with parallel veins, an embryo with a single cotyledon (which may be retained in the seed as an absorbtive organ, so you cannot see it, as in grasses), adventitious roots (the tap root aborts at an embryonic stage), stems with scattered vascular bundles (not in a ring) and a particular type of sieve cell plastids. One can add to this set that monocot flowers are often in threes, but this also occurs widely in the magnoliids and some eudicots. Exceptions to these are frequent among monocots; for example, most members of Pandanales have tetramerous flowers, and several forest-adapted monocots have leaves with reticulate venation. Possession of

the last must be qualified by stating that all monocot leaves are derived from the apex of the leaf primordium, which perhaps could be considered the best candidate for an unqualified monocot synapomorphy. It is not, however, a feature that you can see with the naked eye or even a hand lens, so it is really not all that helpful to the field botanist. Much has been made about a possible aquatic origin for the monocots, and it seems clear that among the early diverging lineages are a large number of taxa that are either aquatic (many Alismatales) or inhabit wet habitats. The same argument has been advanced for the angiosperms as a whole, pointing in particular to

A landscape dominated by monocot families, Chapada Diamantina, Bahía, Brazil , including Arecaceae, Bromeliaceae, Cyperaceae, Orchidaceae, Poaceae and especially Velloziaceae

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MONOCOTS

Nymphaeales, but it is also clear that the taxa comprising these early-diverging clades are relictual and perhaps not representative of the characteristics of their ancestors. The answers to these questions can probably be established by further study of the fossil record and perhaps by understanding patterns of gene expression, but at present it seems unwise to come down clearly on one side of this debate. We prefer to see these as exciting directions for further research. From the study of whole plastid genome sequences, the position of the monocots seems to be as sister to Chloranthaceae, eudicots or the magnoliid clade, and studies of many low-copy nuclear genes are not yet far enough advanced to provide a robust answer to the question of what are the closest relatives of monocots. What is clear is that monocots represent the earliest major radiation of angiosperms; they have collectively conquered all ecological zones on the planet, including marine and freshwater systems, deserts and edges of glaciers. Monocots occur in both polar zones and dominate landscapes across the planet. They include some of the smallest and some of the largest angiosperms – some of the largest palms can individually be seen from orbiting satellites, which is even more remarkable when realising that they are really giant herbs that do not have secondary growth in their trunks. There are tree-forming monocots with secondary growth, albeit a type termed ”anomalous”, such as in agaves, aloes, cordylines, Joshua trees and other yuccas. Finally, some monocots are among the most adaptable and productive domesticated organisms on earth, providing us with food, fibres, medicines and building materials. It is difficult to imagine how seven billion people could be fed if it were not for the efficient production of starch and sugar by grasses (wheat, rice, maize, sugarcane and others). Other monocots 116

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General references: Chase MW. 2004. Monocot relationships: an overview. American Journal of Botany 91: 1645–1655. Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall MR, Price RA, Hills HG, Qiu YL, Kron KA, Rettig JH, Conti E, Palmer JD, Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD, Hedrén M, Gaut BS, Jansen RK, Kim KJ, Wimpee CF, Smith JF, Furnier GR, Strauss SH, Xiang QY, Plunkett GM, Soltis PS, Swensen SM, Williams SE, Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn GH Jr, Graham SW, Barrett SCH, Dayanandan S, Albert VA. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Annals of the Missouri Botanical Garden 80: 528–580. Chase MW, Fay MF, Devey D, Maurin O, Rønsted N, Davies J, Pillon Y, Petersen G, Seberg O, Tamura MN, Asmussen

CB, Hilu K, Borsch T, Davis JI, Stevenson DW, Pires JC, Givnish TJ, Sytsma KJ, McPherson MA, Graham SW, Rai HS. 2006. Multigene analyses of monocot relationships: a summary. Aliso 22: 63–75 Columbus JT, Friar EA, Porter JM, Prince LM, & Simpson MG (eds). 2006. Monocots: comparative biology and evolution, excluding Poales. Rancho Santa Ana Botanical Garden, Claremont (Aliso 22). Dahlgren RMT, Clifford HT, Yeo PF. 1985. The families of the monocotyledons. Springer, Berlin. Duvall MR, Clegg MT, Chase MW, Lark WD, Kress WJ, Hills HD, Eguiarte LE, Smith JF, Gaut BS, Zimmer EA, Learn GH Jr. 1993. Phylogenetic hypotheses for the monocotyledons constructed from rbcL sequences. Annals of the Missouri Botanical Garden 80: 607–619. Ray J. 1682. Methodus plantarum nova. Faitborne & Kersey, London. Rudall PJ, Cribb PJ, Cutler DF, Humphries CJ (eds). 1995. Monocotyledons: systematics and evolution. Royal Botanic Gardens, Kew. Wilson KL, Morrison DA (eds). 2000. Monocots: systematics and evolution. CSIRO, Collingwood.

Grove of Livistona mariae subsp. rigida, Joe Creek, Judbara-Gregory National Park, Northern Territory, Australia

Maarten Christenhusz holding a fruit of coco-demer, Lodoicea maldivica, containing the largest seed in the world, Praslin, Seychelles

also provide staple foods, such bananas, dates, yams and taro, but this pales in comparison to food produced by grasses, especially when fodder for grazing animals is included.

ACORALES

MONOCOTS

ACORALES This small order consists of a single family of rhizomatous, sweet-smelling herbs with aromatic oil cells. They are sister to all other monocots.

63. ACORACEAE Sweet-flag family

These plants have leaves that are organised in a fan at the tips of much-branched, functionally vessel-less rhizomes. The leaves are narrow, sword-shaped, unifacial (with an internal morphology that is the same on both surfaces) and without stipules, but the blades are intercalated and sheathing at the base (i.e. equitant). Veins are strictly parallel. Inflorescences are a solitary spadix (without a spathe in contrast to Araceae; see below) borne laterally on a leaf. The conical fingeror tail-like spadix is densely covered with flowers that are partly sunken in the spadix Acorus gramineus, fruits, Mt Tianmu, Anhui, China [63]

axis. The flowers are bisexual and lack bracts. They are trimerous, with the six thin petals clasping the stamen filaments. Six stamens are in two whorls of three, have linear filaments and rounded anthers that sit atop the clasping petals and open via a longitudinal slit. The bi- or trilocular ovary is superior with a sessile stigma. The fruit is a berry with a leathery skin and holds one to five (to nine) oblong to ellipsoidal seeds. Distribution: This is a family comprised of a single genus perhaps native to East and Southeast Asia. It also occurs throughout Europe and North America, but it has been naturalised in Europe and Egypt since ancient times and has become part of the flora in those parts. Phylogeny and evolution: Acorus was for most of its history placed in Araceae, but there is a long list of technical characters that separate them, and it has now been shown that the families are not at all closely related. Acoraceae are isolated and sister Acorus calamus, Ruissalo Botanical Garden, Turku, Finland [63]

to all other monocots and thus placed in their own order Acorales. Macrofossils of Acorus and Acorites are known from the Eocene of North America and Europe. The age of the crown group is estimated to be c. 20 to 50 million years old, whereas the stem lineage is closer to 130 million years old. Genera and species: Acoraceae consist of the single genus Acorus with two species: Acorus calamus and A. gramineus. Uses: Dried rhizomes of Acorus calamus, known as rhizoma calami, have been used for eye and stomach ailments, and were already in use by the ancient Egyptians. Tea from the chopped roots is said to help digestive problems and can remove phlegm from the lungs; it clears congestion and “tranquilises” the mind. Calamus oil is still used in medications today; it contains high levels of β-asarone, which can be effective in the treatment of neurological diseases. Prolonged exposure to this compound is toxic, and this oil is therefore not deemed suitable for human consumption. It should not be used internally without expert supervision. It has been marketed as ‘legal ecstasy’ due to its hallucinogenic effect in large concentrations. Its roots have also been used in perfumes and liqueurs and as a poor-man’s substitute for ginger, cinnamon and nutmeg. Acorus gramineus has been used for similar purposes in East Asia. Etymology: Acorus is derived from ακορών (acoron), the name used by Dioscorides to describe the root of sweet flag, which was initially confused in Europe with the yellow flag iris (Iris pseudacorus); both were used medicinally to treat inflammation of the eyes. The name was derived from the Greek κόρης (koris), pupil.

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ALISMATALES

MONOCOTS

ALISMATALES Families 64 to 77 comprise the order Alismatales, an order that is approximately 125 million years old. The order is composed of predominantly hydrophytes, plants associated with water, although in the large family Araceae, epiphytes, lianas and geophytes have also evolved. The order also includes a number of seagrass families, one of the few vascular plant clades that include submerged marine species. Many small families have been recognised in this order due to their striking and different adaptations to aquatic habitats.

Wolffia arrhiza, Helsinki Botanical Garden, Finland [64]

Gymnostachys anceps, New South Wales, Australia [64]

Lysichiton americanus, Bodnant Gardens, Wales, UK [64]

64. ARACEAE

small leaf-like structures that are poorly differentiated, usually one or several clustered to 18 mm long, surrounded by a bifid scale or not. These structures can be veined or not and can have roots or not. Leaves of other Araceae are usually differentiated into a petiole and blade that can be variable in shape, the primary venation usually pinnate, but often pedate or palmate at the base, or parallel, the secondary venation usually net-veined. The leaf is flat and without a petiole in Gymnostachyoideae. Inflorescences are usually composed of a spadix, a spike in which the small flowers are sessile on or sunken into the axis, which is surrounded by a spathe, a leaf that has a specialised shape that is often colourful, although the spathe is difficult to observe in Gymnostachys, Orontium and Lemnoideae. Lemnoideae have reproductive pouches in

the blade, which are enclosed by a cap, and the bisexual flowers are one or two per blade, with one or two stamens and a bottle-shaped unilocular ovary per flower. The flowers in other Araceae are usually sessile, bisexual or unisexual. When unisexual, the female flowers are usually at the base of the spadix and the male flowers are in the middle or at the tip, sometimes separated from the female flowers by sterile flowers. The reduced petals are free, partly fused or completely fused and are in a single whorl. Stamens are usually in two whorls of two or three, sometimes fewer or more, free or fused. The superior ovary is composed of one to three (or more) fused carpels, with a short or absent style and a wet stigma. The fruit is a berry that is rarely dehiscent, borne on the fleshy spike and rarely with the berries fused into a syncarp, often with oxalate crystals.

Calla-lily family

This varied family consists of terrestrial, epiphytic or aquatic herbs that can be diminutive to enormous, self-supporting, climbing or free-floating. Geophytic species make stem tubers. They can have true or false stems, mature shoots often forming a sympodium. Duckweeds, the free-floating members of Lemnoideae, are reduced to 118

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MONOCOTS

Spathiphyllum wallisii, Guadeloupe [64]

Typhonium giganteum, Royal Botanic Gardens, Kew, UK [64]

Zantedeschia aethiopica naturalised along the coast near San Francisco, California, USA [64]

Distribution: Araceae have a near global occurrence due to the wide distribution of Lemnoideae. The family is absent from permanently dry and frozen regions, and it is most diverse in the tropics.

the broad sense are monophyletic): Gymnostachyoideae – Gymnostachys (1); Orontioideae – Lysichiton (2), Orontium (1) and Symplocarpus (5); Lemnoideae – Lemna (13), Spirodela (4), Wolffia (11) and Wolffiella (10); Aroideae – Aglaodorum (1), Aglaonema (22), Alloschemone (2), Alocasia (80), Ambrosina (1), Amorphophallus (c. 200), Amydrium (5), Anadendrum (12), Anaphyllopsis (3), Anaphyllum (2), Anchomanes (6), Anthurium (c. 940), Anubias (8), Apoballis (12), Aridarum (10), Ariopsis (2), Arisaema (c. 180), Arisarum (3), Arophyton (7), Arum (c. 30), Asterostigma (8), Bakoa (3), Biarum (21), Bognera (1), Bucephalandra (2), Caladium (14), Calla (1), Callopsis (1), Carlephyton (3), Cercestis (10), Chlorospatha (28), Colletogyne (1), Colocasia (8), Croatiella (1), Cryptocoryne (60), Culcasia (27), Cyrtosperma (12), Dieffenbachia (57), Dracontioides (2), Dracontium (24), Dracunculus (2), Eminium (9), Epipremnum (15), Fenestratarum (2), Filarum (1), Furtadoa (2), Galantharum (1), Gearum (1), Gonatopus (5), Gorgonidium (8), Hapaline (8), Helicodiceros (1), Hestia (1), Heteropsis (17), Holochlamys (1), Homalomena (124), Incarum (1), Jasarum (1), Lagenandra (15), Lasia (2), Lasimorpha (1), Leucocasia (1), Lorenzia (1), Mangonia (2), Monstera (40), Montrichardia (2), Nephthytis (6), Ooia (2), Pedicellarum (1), Peltandra (2), Philodendron (c. 490), Phymatarum (1), Pichinia

(1), Pinellia (9), Piptospatha (12), Pistia (1), Podolasia (1), Pothoidium (1), Pothos (56), Protarum (1), Pseudohydrosme (2), Pycnospatha (2), Remusatia (4), Rhaphidophora (c. 100), Rhodospatha (29), Sauromatum (9), Scaphispatha (2), Schismatoglottis (107), Schottariella (1), Scindapsus (35), Spathantheum (2), Spathicarpa (3), Spathiphyllum (50), Stenospermation (50), Steudnera (9), Stylochaeton (20), Synandrospadix (1), Syngonium (34), Taccarum (6), Theriophonum (7), Typhonium (69), Typhonodorum (1), Ulearum (2), Urospatha (11), Vietnamocasia (1), Xanthosoma (75), Zamioculcas (1), Zantedeschia (8), Zomicarpa (2) and Zomicarpella (2).

Phylogeny and evolution: The duckweeds, Lemnoideae (former Lemnaceae), are sister to Aroideae, with Orontioideae and Gymnostachyoideae (with genera always included in Araceae) successively sister to this clade. Calla palustris is difficult to place on morphological and molecular grounds, but nevertheless belongs to Aroideae, in spite of having bisexual flowers. Fossils of Aroideae have been found in Palaeocene and Eocene sediments. Fossils of Limnobiophyllum, interpreted as an intermediate stage between Lemnoideae and Aroideae are known from the Late Cretaceous, but the family is estimated to be of greater age, c. 122 million years old. It was long believed that the duckweeds evolved through Pistia, which indeed has striking similarities in developmental stages to Spirodela. This resemblance is superficial, and it is now known that the two are not closely related. Genera and species: Araceae consist of 118 genera and c. 3,300 species, divided in four subfamilies, with Aroideae sometimes divided into a further five subfamilies (not followed here, given that Aroideae in

Uses: All Araceae have calcium oxalate crystals, and therefore the fruit, leaves and tubers have to be consumed with care. The crystals can often be removed by boiling with a pinch of baking soda or by soaking in cold water overnight. Taro is the edible tuber of several species of Araceae, of which Colocasia esculenta, native to South Asia, is the most commonly and most widely cultivated species. The giant taro or elephant ear, Alocasia macrorrhizos, the domesticated form of which is native to the Philippines, is also edible after cooking the stems for a considerable length of time. Taro is a staple food for several groups of Pacific islanders and is now also cultivated in Plants of the World

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sterile flowers male flowers female flowers

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Calla palustris, Ruissalo Botanical Garden, Turku, Finland [64]

Amorphophallus titanum, Royal Botanic Gardens, Kew, UK [64]

The aroid inflorescence, with female flowers at the base of the spadix, male flowers above and sterile flowers closing the opening of the spathe. Arum maculatum, Lleyn Peninsula, Wales, UK [64]

East Africa. Pulaka, Cyrtosperma merkusii, is another food crop from the Pacific, where it is grown in fresh-water swamps in atolls and is often the only source of starch there. The arrowleaf elephant ear, Xanthosoma sagittifolium, originated in tropical America and is used for various dishes, such as alcapurrias, mondongo, pasteles and sancocho; it is also an important ingredient of Surinamese pom. Leaves are also edible after cooking and called taioba in Brazil. Konjac or konnyaku, Amorphophallus rivieri, is used in Japanese cuisine as (shirataki) noodles. The flavourless jellylike substance is also used to make fruit candy and because konjac has almost no calories, it is used as a diet food. The Swiss-cheese plant, Monstera deliciosa from Central America, is a commonly grown houseplant in temperate regions. It has an edible fruit that looks like a pineapple and has a sweet flavour. Species of Cryptocoryne and Pistia stratiotes are popular aquarium plants. Orontium aquaticum, Calla palustris, Lysichiton americanus, L. camtschatcense and Zantedeschia aethiopica are commonly offered in garden centres as pond plants. Duckweeds have high protein content, and because of their rapid vegetative reproduction they are sometimes grown as a crop for animal fodder. Wolffia globosa is consumed by people in Southeast Asia, and Lemna gibba is eaten as a vegetable in the Levant. Some aquatic species accumulate nutrients

and pollution from water and are used for waste water management. The plants are also composted, releasing nutrients for crop plants. Many species of Araceae are attractive and make good garden or house plants. The most popular are species of Aglaonema, Alocasia, Amorphophallus, Anthurium, Arisaema, Caladium, Colocasia, Dieffenbachia, Epipremnum, Monstera, Philodendron, Sauromatum, Scindapsus, Spathiphyllum, Syngonium, Zamioculcas and Zantedeschia, but other genera may also be on offer from specialist nurseries. Anthurium and Zantedeschia flowers are commonly grown for the cut-f lower industry. A great variety of colour forms of ‘anthuriums’ and ‘calla-lilies’ have been bred for this purpose.

arums frequently feature in news items when they flower in a botanical garden, where they spread their powerful foetid fragrance.

Christenhusz, Fay & Chase

Smallest plants: Wolffia angusta and W. globosa are the smallest species of vascular plants, with the entire plant being 0.8 × 0.4–0.6 × 0.7–1.0 mm. Lemnoideae are usually distributed between fresh water bodies by currents and by sticking to the legs and feathers of water fowl. They mostly spread vegetatively and only rarely reproduce by seeds. The big stink: The titan arum, Amorphophallus titanum, from Indonesia, produces not only enormous umbrella-like leaves, but also a two-metre tall spadix, the largest unbranched inflorescence of any plant. Flowering titan

Kettle traps: Plants with unisexual flowers and complex spathes have a specialised pollination syndrome called the ‘kettle trap mechanism’. In genera including Arisaema, Arisarum, Arum, Cryptocoryne and Philodendron, the female flowers are at the base of the inflated spathe and the male flowers are above, with sterile, down-pointing and hair-like flowers in between or above. The tip of the spadix often produces a scent attracting pollinators, and the spadix can be a few degrees warmer than the surroundings. The pollinators get trapped in the chamber with the female flowers until these are pollinated and the sterile flowers shrivel, during which time the anthers ripen and cover the insects with pollen on their way out of the chamber. Much less is known about the pollination mechanisms of species with bisexual flowers, but in these the flowers must provide other attractants for the insects to remain on the inflorescence. Etymology: Arum is the Roman name for the arum-lily, most likely derived from Greek άρον (aron), also the name for the plant in ancient Greek. Aaron is also the name of the brother of Moses in the Bible, but the name of the plant is not a cognate with the biblical name, which is of Egyptian origin.

ALISMATALES

MONOCOTS

65. TOFIELDIACEAE False-asphodel family

spathe-like bract. The actinomorphic flowers are subtended by bracteoles, which are simple or forked. The six tepals are free or basally fused and usually persistent in fruit. The six or nine stamens are usually free or basally fused, with filaments that are sometimes shorter, but often longer than the petals. The superior ovary has three carpels fused only at the base, each carpel topped with a short style. Fruits are septicidal capsules containing five to numerous seeds.

Tofieldiaceae are perennial rhizomatous herbs. The linear leaves are persistent and mostly basally arranged in a flat fan. The inflorescence has up to four leaves, topped by a raceme or spike, sometimes surrounded by a

Distribution: This North Temperate family occurs mainly in boreal and montane regions, extending south into the southeastern USA, Japan and China, with isolated populations in the northern Andes and Guayana Highlands.

Tofieldia coccinea, Crûg Farm Plants, Wales, UK [65]

Triantha occidentalis subsp. montana, Valley County, Idaho, USA (CD) [65]

Phylogeny and evolution: Tofieldiaceae are dated to have diversified c. 100 million years ago. Pleea is sister to the rest of the family. They were previously included in Nartheciaceae or even in a broad concept of Liliaceae, but these genera are not closely related to either; they form an isolated lineage within Alismatales. Genera and species: This family has four genera and c. 30 species: Harperocallis (14), Pleea (1), Tofieldia (12) and Triantha (4). Etymology: Tofieldia was named for Thomas Tofield of Wilsick (1730–1779), a British botanist and civil engeneer.

Tofieldia calyculata in fruit, private garden, Kingston upon Thames, Surrey, UK [65]

Tofieldia calyculata, private garden, Kingston upon Thames, Surrey, UK [65]

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66. ALISMATACEAE Water-plantain family

This is a family of aquatic, monoecious and dioecious plants that are rooted in the soil under fresh water and have stems that are emergent, submersed or floating. They can be annual but are usually perennial. Stems grow underground or under water (rhizomes) and can have stolons and fibrous roots, often forming a corm. Juice is milky in many species. Aerial stems are fleshy when present. Leaves form a basal rosette, are spirally arranged or are whorled along the stems, usually with a petiole, variable in shape and size, especially between submerged, floating and emergent leaves. Petioles are rounded to triangular, often longer than the blade and sheathing basally. Blades are linear, orbicular, lanceolate or sagittate, the bases usually cordate, sagittate or hastate, sometimes attenuate, the apices blunt, acute or

Alisma plantago-aquatica, Ruissalo, Finland [66]

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MONOCOTS

acuminate, the margins entire and sometimes wavy. Primary veins arch from the base to the apex, diverging where the leaf blade widens, with transverse secondary veins forming a regular net-veined pattern. Inflorescences are erect and emergent or floating, and they are usually umbels that are often in a whorled raceme or panicle. Flowers are regular, with three persistent and green sepals and three coloured delicate petals. There are six, nine or many free stamens, and these have bilocular, basifixed or versatile anthers. The ovary is superior and has three to many carpels that are free or basally fused. The fruit is a dry achene or follicle. The numerous seeds are glandular, ridged and U-shaped. Distribution: This family has a nearly global occurrence, but it is absent from most arid areas. Phylogeny and evolution: Alismataceae evolved during the Late Cretaceous (c. 80 million years ago) and are deeply split into two clades, one corresponding to Alisma, Baldellia, Damasonium and Luronium, the other including all other genera and the former family Limnocharitaceae. Echinodorus is polyphyletic with regard to Sagittaria, hence acceptance of Albidella and Helanthium as

separate genera, but the type species Echinodorus berteroi is morphologically different from the other species, which may result in the majority of Echinodorus requiring a new generic name. Genera and species: This family of 17 genera has c. 115 species: Albidella (1), Alisma (8), Astonia (1), Baldellia (3), Burnatia (1), Butomopsis (1), Caldesia (4), Damasonium (6), Echinodorus (30), Helanthium (3), Hydrocleys (5), Limnocharis (2), Limnophyton (5), Luronium (1), Ranalisma (2), Sagittaria (39) and Wiesneria (3). Uses: Leaves of Limnocharis flava are eaten as a vegetable in tropical Asia. Sagittaria sagittifolia has an edible corm and is cultivated for that purpose in East Asia. Sagittaria latifolia roots were used by native Americans as a food source. Several Echinodorus and Helanthium species are commonly grown as aquarium plants, and species of Alisma, Echinodorus and Sagittaria are grown as pond ornamentals, especially in the tropics and subtropics, where they frequently naturalise in aquaculture. Etymology: Αλισμα (alisma) is an Ancient Greek name for a water plant.

Echinodorus grandiflorus, Helsinki Botanical Garden, Finland [66]

Limnocharis flava, Ruissalo Botanical Garden, Turku, Finland [66]

Hydrocleys nymphoides, Tahiti [66]

Luronium natans, Royal Botanic Gardens, Kew, UK [66]

ALISMATALES

MONOCOTS

67. BUTOMACEAE Flowering-rush family

are superior and composed of six carpels in two whorls that are fused at the base only. Fruits are dry separate follicles that open by central slits and contain numerous ribbed, straight seeds.

68. HYDROCHARITACEAE Frogbit family

Distribution: This family is restricted to temperate Eurasia, but is naturalised in eastern North America.

These freshwater perennials have creeping rhizomes with axillary buds that may develop into bulbils with which they can spread in their aquatic habitat. Leaves are borne on the apex of rhizomes and are somewhat arranged in a plane. Leaves lack petioles and are sheathing basally and linear, triangular in transverse section at the base but flattening towards the apex and up to a metre long. Inflorescences emerge from either side of rhizomes among leaves and are erect and composed of an umbellike complex of cymes, which are subtended by c. three bracts. The actinomorphic flowers are stalked and trimerous. The three sepals are darker pink than the three whitish pink petals, which are all free. The nine pink stamens are in two whorls: the outer of six and the inner of three stamens with basifixed anthers. Ovaries

Phylogeny and evolution: The family used to include Butomopsis, Hydrocleys and Limnocharis, but molecular phylogenetic studies placed these genera firmly in Alismataceae. Fossils attributed to this family are often inconclusive. The family is sister to Hydrocharitaceae. Genera and species: This family includes a single species, Butomus umbellatus. Uses: The starchy rhizomes are eaten in some parts of Russia where they are mixed with grain to make bread or cooked as a vegetable. Etymology: Butomus is derived from ancient Greek βους (bous), an ox, and τομος (tomos), to cut, in reference to leaves that cannot be eaten by cattle because their sap blisters the mouth.

Butomus umbellatus, Ruissalo Botanical Garden, Turku, Finland [67]

These aquatics are annual and perennial, unisexual and bisexual herbs. They have simple or complex branched stems, and the leaves are whorled, spirally arranged or distichous (in a single plane). Leaves are simple, leaf-like or scale-like, and can have thin translucent stipules (not always present). Petioles can be short or elongate, winged or not and are sometimes sheathing at the base or not clearly defined from the leaf base. Leaf blades are submerged or floating, rarely emergent, linear to round or heart-shaped. Venation is often with parallel main veins and numerous cross-veins; the margin is entire or dentate, but sometimes with spines. Inflorescences are sessile or stalked. They develop inside one or two, free or united bracts that are often called a spathe. Inflorescences are composed of one or more monochasia, often with complex arrangements. Flowers are emergent, at the water surface, or submerged (see pollination below). Flowers are bisexual or unisexual, regularly actinomorphic or becoming zygomorphic. Male flowers can become detached before anthesis and float to the female flowers or are persistent on the plant with floating pollen. Female flowers are retained on the mother plant. Bisexual flowers are often selfing without opening (cleistogamous). Perianths are present and showy, evanescent or absent (Najas), and sepals are free, three (or absent). Petals are usually three or fewer, sometimes absent, often large and showy. Stamens are usually formed in six whorls of three, sometimes fewer or partly staminodial and in some cases reduced to a single stamen. Ovaries are inferior, with three to 20 carpels, with as many styles as carpels. Nectaries, if present, are three and formed at the base of the

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ALISMATALES styles. Peduncles are often curved or spiralling, pulling the developing fruit under water. Fruits are berries, regularly or irregularly dehiscing fleshy capsules or achenes (Najas). Seeds are cylindrical or ellipsoidal and have a complex outer structure. Distribution: This widespread family is distributed across North America, the Caribbean, northern South America, southern Brazil, temperate Eurasia, Sub-Saharan Africa, Madagascar, the Indian Ocean, South and Southeast Asia north to Japan, Malesia south

Ottelia ovalifolia, Victoria, Australia (JC) [68]

MONOCOTS

to Australia, New Guinea and throughout the Pacific to French Polynesia and Hawaii. Phylogeny and evolution: Hydrocharitaceae evolved c. 75–65 million years ago, probably originating in Asia. The seagrass clade evolved much more recently, 19 million years ago. Stratiotes dates back to the Mid Eocene with a fossil record of 48 million years, exhibiting little morphological variability. Hydrocharitites and Hydromystria are Miocene and Tertiary fossils similar to extant Hydrocharis dubia. Najas guadalupensis, Payette Lake, Idaho, USA (CD) [68]

Egeria najas, Copenhagen Botanical Garden, Denmark [68]

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Najas has been difficult to place on morphological grounds, due to its extreme reduction and adaptations to aquatic pollination. For this reason, it was often placed in its own family Najadaceae. DNA analyses in the 1990s demonstrated that Najas belongs to Hydrocharitaceae, despite several morphological and anatomical differences. This affiliation was already suggested in the 1930s, although an association with the seagrasses was later more commonly assumed.

Hydrocharis morsus-ranae, private garden, Kingston upon Thames, Surrey, UK [68]

Vallisneria americana, Helsinki Botanical Garden, Finland [68]

ALISMATALES

MONOCOTS

Genera and species: The family consists of c. 16 genera and c. 130 species of fresh water and marine aquatics: Appertiella (1), Blyxa (12), Egeria (3), Elodea (6), Enhalus (1), Halophila (19), Hydrilla (1), Hydrocharis (3), Lagarosiphon (9), Limnobium (2), Najas (36), Nechamandra (1), Ottelia (19), Stratiotes (1), Thalassia (2) and Vallisneria (14). Uses: Leaves and inflorescences of Ottelia cordata are eaten in Southeast and East Asia. Some species are popular pond and aquarium plants, and several of these have escaped into nature: Egeria densa (largeflowered waterweed) from South America in waterways in the Northern Hemisphere and Australasia; Elodea canadensis (Canadian waterweed), probably one of the most infamous invasives, blocking waterways in Europe and later being replaced by an invasion of North American Elodea nuttallii; Hydrilla verticillata creating havoc in the southern USA; frogbit (Hydrocharis morsusranae) from Europe in Canada, where it also has created water management issues; South African Lagarosiphon major becoming a pest in New Zealand; and Asian tapegrass (Vallisneria) creating problems in irrigation canals even within its natural range. Several species of Egeria, Elodea, Hydrocharis, Stratiotes and Vallisneria are still commonly offered in the aquarium and pond plant trade, often as oxygenating plants. Elodea can be used to remove pollutants from bodies of water and as a nutritious fodder for animals. Pollination: There are five major mechanisms of pollination. Insect-pollinated species have large showy flowers and nectaries, with male and female flowers alike. In Blyxa the flowers are insect-pollinated as well, but the flowers do not offer a reward. The female flowers mimic the male flowers but have petaloid styles that resemble the petals of the male flower, whereas the petals of the female flowers are reduced and thread-like. These petaloid structures exude a fluid, and the wetted pollen is transported by carnivorous insects that are attracted by the dead insects in the fluid. In Limnobium, pollen is winddispersed. Male flowers are held above the

females. Hydrilla verticillata has explosive anthers. Female flowers form an inverted bell under water, but open to the air, the inside being hydrophobic and the stigma remaining dry. Only pollen landing directly in the bell is effective in pollination. A number of genera have flowers on the water surface. In these species the male flowers are free-floating or the pollen floats, reaching the female flowers that rest on the water surface. Sepals are water repellent, causing changes in the surface tension of the water, allowing the pollen to be drawn in by the female flowers. In the submerged and marine genera Halophila, Thalassia and Najas, flowers are reduced and pollen released in gelatinous, stringy masses, which are picked up by submerged or superficial female flowers. Etymology: Hydrocharis is derived from the Greek ύδωρ (hydor), water, and χάρης (charis), grace.

69. SCHEUCHZERIACEAE Rannoch-rush family

Scheuchzeria palustris, Kittilä, Finland (HV) [69]

These perennial plants have horizontal rhizomes, clothed in old leaf bases, rooting from the nodes. Stems are erect with alternate leaves. The sheathing base is open and has two delicate protrusions at the tip of the sheath at the base of the linear blade. The blade has several parallel veins. Inflorescences are simple racemes with bracts subtending the pedicels. Flowers are bisexual, actinomorphic, with six tepals all similar, free and lanceolate-elliptic. Six stamens are free and dehisce by longitudinal slits. Ovaries are superior, the carpels three (sometimes six), mostly free (fused at the base only), with sessile stigmas. The fruit is composed of three dehiscing follicles with one or two seeds.

Distribution: This circumarctic inhabits sphagnum bogs.

family

Phylogeny and evolution: They appear morphologically similar to Juncaginaceae and associated families, but despite a shared chemistry (triglochinin) these are not sister groups. Genera and species: This family consists of a single species, Scheuchzeria palustris. Etymology: Scheuchzeria was named in honour of Swiss physician and scholar Johann Jakob Scheuchzer (1672–1733), who was well known for his accounts of the natural phenomena of Switzerland.

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ALISMATALES

MONOCOTS

Aponogeton distachyos, private garden, Kingston upon Thames, Surrey, UK [70]

Triglochin bulbosa, Western Australia [71]

Triglochin bulbosa, Western Australia [71]

Cycnogeton procerum, Royal Botanic Gardens, Kew, UK [71]

70. APONOGETONACEAE

Petals are not differentiated; they can be absent but there are usually two to six. Stamens are usually six (absent in female plants). The ovary is superior, composed of three, free, sessile carpels; the stigma is a ridge on the side of the short-styled carpel, rudimentary in male plants (rarely). The fruit is a follicle with a curved beak, submerged until seeds are released after decay, after which the seeds may float for a short while and sink in a new spot. Seeds have no dormancy.

are considered a delicacy and commonly cultivated as waterblommetjies (‘water flowers’) for this purpose in South Africa. The cooked inflorescences have a meaty flavour. The starchy tubers of several species can be eaten by humans and livestock. Several other species are frequently grown as aquarium or pond ornamentals.

Waterblommetjie family

These perennial fresh water aquatic plants are rarely dioecious and have an underground stem and a starchy rootstock or rhizome. Leaves are spirally arranged in a rosette around the tip of the rhizome and are submerged or floating (rarely emergent). Leaves are usually stalked, and the blade is oval with rounded tips and apices. Venation is parallel, merging at the tip and base of the leaf, with parallel secondary cross-venation. In some species the leaf tissue between the veins is absent, forming a net. Inflorescences are long-stalked, emergent and bracteate, the bracts covering the inflorescence branches in bud. The inflorescence is spicate and can be composed of one to several spikes. Flowers are sessile on the bracteate spikes, more or less spirally arranged or sideways.

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Distribution: A family found in Sub-Saharan Africa, Madagascar, South and Southeast Asia, Malesia, New Guinea and Australia. Phylogeny and evolution: Aponogetonaceae belong to the clade including the seagrasses and Juncaginaceae, in which they are an early-branching lineage. The vegetative morphology is similar to that of Potamogetonaceae, and the group has often been associated with that family.

Etymology: Aponogeton is composed of the name of a Roman spring, the Aquae Aponi, near Padua, Italy (in turn from Greek απονος, aponos, painlessness or heartlessness), and the Greek γειτον (geiton), a neighbour.

71. JUNCAGINACEAE Arrowgrass family

Genera and species: The single genus Aponogeton has 56 species. Uses: Inflorescences and flower buds of Cape pondweed (Aponogeton distachyos)

This is a family of annual and perennial, glabrous herbs with creeping rhizomes. Plants

ALISMATALES

MONOCOTS

can grow terrestrially, in mud or submerged in fresh or brackish water with floating leaves. Roots sometimes bear tubers from which new plants can emerge. Leaves form a rosette at the tip of the rhizome with an open sheathing base and a linear flattened or rounded blade that has one to several veins. Sheaths often remain as a fibrous mantle around the rhizome. Inflorescences lack bracts and are composed of terminal spikes or spike-like racemes. The actinomorphic flowers are bisexual or unisexual, in the latter case usually with female flowers at the base, bisexual flowers in the middle and male flowers at the tip (or males entirely absent). One or six tepals are placed in one or two whorls or are (rarely) absent. Sessile stamens number one, four or six (or are absent); the anthers have two chambers that open with slits. The superior ovary is composed of one, three or six fused carpels that separate upon maturing. Stigmas are sessile or with a short style, rarely the style elongate and filiform in basal flowers (Triglochin scilloides). Fruits are achenes that are sometimes hooked or horned at the tip. Distribution: This family is found across the temperate Northern Hemisphere, in South America, southern Africa, New Guinea, Australia and New Zealand. Phylogeny and evolution: Juncaginaceae are dated to c. 82 million years, diversifying c. 52 million years ago. Australian species of Triglochin are now placed in the genus Cycnogeton. Lilaea scilloides has been merged with Triglochin, from which it differs only in its elongate styles. Tetroncium is sister to the other two genera. Genera and species: A family of three genera with 34 species: Cycnogeton (8), Tetroncium (1) and Triglochin (25). Uses: Rhizomes of Cycnogeton procerum and leaves of Triglochin maritimum are edible. Etymology: Juncago is derived from Latin juncus, a rush, and the suffix -ago meaning ‘a sort of’. Juncago is a later synonym of Triglochin.

72. MAUNDIACEAE Maund’s-arrowgrass family

This family consists of perennial, aquatic, glabrous herbs with 5 mm thick rhizomes and emergent tufts of leaves along their lengths. Leaves are triangular in cross-section and up to 80 cm long, with open sheaths at the base. The sheaths remain when the leaves wither, forming a fibrous mantle around the rhizome. Inflorescences are terminal spikes to 10 cm long, without bracts. Flowers are bisexual, actinomorphic, with two to four, scale-like, fleshy tepals. The two to eight stamens are short and the anthers unilocular, opening with longitudinal slits. The superior ovary is composed of three or four fused carpels that separate upon maturing, each with a spreading beak. Stigmas are sessile and elongate. Fruits are beaked achenes. Maundia triglochinoides, collected by S. T. Blake (20053) in Queensland, Australia (Herbarium Kew)

Distribution: This species occurs in northeastern Australia (coastal New South Wales into southern Queensland), in swamps and shallow fresh water on heavy clay soils. It is listed as vulnerable, highly likely to become endangered, due to draining and filling of swamps and lowering of water tables in the areas where it occurs. Phylogeny and evolution: Even though morphologically similar to Juncaginaceae (and traditionally placed there), Maundia is sister to a clade that unites Potamogetonaceae, Juncaginaceae and the seagrass families (Cymodoceaceae, Posidoniaceae, Ruppiaceae and Zosteraceae). It may be that in the future all of these families can be merged into an expanded Juncaginaceae, but for the time being a conservative approach places Maundia in its own family. Genera and species: This family is composed of the single species Maundia triglochinoides. Etymology: Maundia was named to commemorate Melbourne medical physician John Maund (1823–1858).

73. ZOSTERACEAE Eelgrass family

[72]

This family is composed of perennial, marine, submerged plants. The creeping, branched rhizomes grow in sandy or muddy substrates on the sea floor and in intertidal zones or on submerged rocks. Their leaves are alternate, in one plane (distichous), with an open, sheathing base and linear blade with parallel venation. Leaves lack stomata. Flowers are unisexual or bisexual, arranged on either side of a stem (when unisexual, male and female flowers

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ALISMATALES alternating), with a spadix that is usually enveloped in a modified leaf sheath (spathe). Male flowers consist of only a single stamen. The anther is bilocular with a ridge-like connective. Pollen is filiform. Female flowers are composed of a single ovary with a short style bearing two long stigmas. Pollination takes place under water, the flowers not emergent. The fruit is an achene. Distribution: Zosteraceae occur in temperate and subtropical oceans: North Atlantic, Mediterranean, Pacific, Chilean coast, Indian Ocean around southern and eastern Africa and cool waters around Australia and New Zealand.

MONOCOTS

Phylogeny and evolution: Zosteraceae diversified c. 30 million years ago. There are no known fossils, the fossil genus Archeozostera being misinterpreted. The family is closely related to Potamogetonaceae. A clone of Zostera marina in the Baltic Sea has been estimated to be at least a thousand years old. Genera and species: This family has two genera and 22 species: Phyllospadix (6) and Zostera (16). Uses: Eelgrass, Zostera marina, has been used as packing material in the past. The large sea meadows of Zostera are important breeding

grounds for numerous marine animals, including commercially important fish and shrimp species. Etymology: The name is derived from a Greek belt or girdle called ζωστήρ (zoster), which was mostly worn by men in ancient Greece (from c. 700 to 500 BC). This of course is in reference to the long belt-like leaves of these plants.

74. POTAMOGETONACEAE Pondweed family

Phyllospadix scouleri, Monterey Bay Aquarium, California, USA [73]

These aquatic plants are usually perennials that have a few roots at the basal nodes of the stems. Plants can be free-floating or rooted in substrate. Stems are slender, often dimorphic, the lower ones resembling rhizomes, and can bear tubers, the upper ones erect and floating with the tips developing into winter buds (turions) in many species. Leaves are floating or submerged, alternate or opposite, sometimes in false whorls, sessile Potamogeton tricarinatus, Oodnadatta, South Australia [74]

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Zannichellia palustris, Germany (CF) [74]

ALISMATALES

MONOCOTS

or petiolate, usually with a pronounced ligule that forms a tube-like open sheath around the stem, which may be completely free from the leaf. Leaves can have one vein when linear, but in many species the leaf blades are wider and the veins several, parallel and often connected with regular crossveins. Inflorescences are borne in the leaf axils or terminate a stem. They are long-stalked and often emergent or sessile (in Althenia, Lepilaena and Zannichellia). They can have two to several flowers or be composed of many flowers in a head or spike. When flowers are bisexual, two to four tepals occur in one whorl and surround the usually four stamens, the filaments fused with the sepals and bilocular anthers opening by slits. In species with unisexual flowers, the perianth of the male flowers can be absent or minute and trilobed, with a single stamen and a one to 12-locular anther opening by slits. In perfect flowers, the superior ovary is usually composed of four distinct carpels with a short

style, but in taxa with separate female flowers petals are a small cup-like sheath or three separate segments enclosing the superior ovary with one to eight free carpels. The fruit is usually formed by individual carpels becoming small drupes. Distribution: The family occurs globally but is absent from the great deserts. Phylogeny and evolution: The earliest fossil records of Potamogetonaceae date from the Palaeocene and become more common during the Eocene with a diversification around 25 million years ago. In previous classifications, Potamogeton was placed in a broad assembly of related aquatic plants such as Halodule, Najas, Ruppia and the seagrass families. They are indeed closely related and could be merged into a single family if Juncaginaceae (the name that has priority) is also included. Zosteraceae appear to be closest to Potamogetonaceae, as these families share similar adaptations to

aquatic habitats. Because of their extreme reduction in vegetative and floral morphology, Zannichellia and associated genera Althenia and Lepilaena were previously placed in a separate family Zannichelliaceae. They share coiled cotyledons, which is a good synapomorphy for the family, but is difficult to see with the naked eye. Genera and species: Potamogetonaceae include six genera with 110 species: Althenia (2), Groenlandia (1), Lepilaena (6), Potamogeton (89), Stuckenia (6) and Zannichellia (6). Uses: Apart from occasional use as ornamentals, the family has few uses. It is an important food source for waterfowl and aquatic animals. Etymology: Potamogeton is derived from the Greek ποταμός (potamos), river, and γειτον (geiton), a neighbour.

Potamogeton crispus, Surrey, England, UK [74]

Posidonia coriacea, Perth, Western Australia [75]

Potamogeton illinoensis, Cordley Lake, Michigan, USA [74]

Posidonia australis, Western Australia (KD) [75]

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ALISMATALES

MONOCOTS

Ruppia megacarpa, South Australia [76]

Ruppia megacarpa, habit, South Australia [76]

Amphibolis antarctica, Adelaide, South Australia [77]

75. POSIDONIACEAE

Genera and species: The family has a single genus, Posidonia, with nine species.

or appearing opposite, with a sheath surrounding the stem and a distinct linear, single-veined blade. The blade is apically serrate, entire towards the base. Flowers are axillary, singular or in few-flowered heads, sometimes in terminal spikes, the peduncles elongating to place the flowers near the water surface at anthesis. Flowers are bisexual, without petals, the two stamens sessile and the anthers opening by slits. The superior ovary is composed of usually four free carpels (rarely two or up to 16), each with a style and a sessile, peltate stigma. Fruits are stalked or sessiles drupes.

Tapeweed family

Washed ashore: Most people will encounter these plants washed ashore. Fragments of these plants break off and the waves work them into perfect globes of fibrous material, often found along Mediterranean or Australian shores.

These marine plants are large submerged herbs with creeping rhizomes covered by remnants of old leaf sheaths. Leaves are alternate, in one plane (distichous) with a sheathing base and distinct blade. The blade is linear with three or more vascular bundles and lacks stomata. The terminal inflorescence is a compound spike-like bracteate raceme on a long flattened peduncle. The bisexual flowers lack petals. The three stamens have sessile anthers that open by slits and release filiform pollen. The superior ovary is composed of a single carpel with an ornate stigma. The fruit is a spongy achene. Distribution: This family occurs in the Mediterranean and temperate oceans of southern and western Australia. Phylogeny and evolution: Clones of Posidonia oceanica in the Mediterranean can be several kilometres across and have been estimated to be dozens of millennia old. 130

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Etymology: Posidonia is named for the Greek god, Ποσειδων (Poseidon), one of the 12 Olympian deities of Greek mythology. His realm is the ocean, and this marine genus is thus appropriately named.

76. RUPPIACEAE Tasselweed family

Distribution: This family grows in ponds, saltmarshes and shallow seas worldwide with patchy distributions on all continents. Phylogeny and evolution: The earliest fossils of Ruppiaceae date from the Palaeocene. A dubious fossil of Limnocarpus from the middle Eocene of Denmark may be a Ruppia because of similarities in the endocarp. The family is closely related to and doubtfully distinct from Cymodoceaceae.

This family is composed of usually annual, submerged, aquatic herbs, found mostly in fresh and brackish water. Roots are few and fibrous. Stems are slender, often dimorphic, with the lower parts rhizomatous and the upper part erect and leafy, but not forming winter buds. Leaves are alternate

Genera and species: This family includes the single genus Ruppia with eight species. Etymology: Ruppia was named to commemorate German botanist Heinrich Bernhard Rupp (1688–1719).

PETROSAVIALES

MONOCOTS

77. CYMODOCEACEAE Turtle-grass family

This is a family composed of perennial marine herbs with creeping, branched rhizomes and often erect leafy branches. Leaves are alternate, in one plane (distichous) with a sheathing base and distinct blade. A thin ligule is present between the sheath and blade. The sheath leaves a scar on the stem after being shed. Blades are linear, flat or rounded, and lack stomata. Flowers are

solitary or arranged in cymes. Unisexual flowers lack petals but are enclosed by leaflike bracts. Male flowers are subsessile or with a pedicel and composed of two stamens. Female flowers are composed of two free ovaries, each bearing a long, unbranched or branched, filiform style. Usually only one carpel develops into an indehiscent fruit that in some genera germinates on the mother plant.

from fossils of the Upper Cretaceous of the Netherlands (Thalassocharis) and the Mid Eocene of Florida (Thalassodendron). The crown clade is estimated to have originated 100 million years ago.

Distribution: Cymodoceaceae occur in tropical and warm temperate seas worldwide, particularly around Australia.

Uses: In places where large marine meadows occur, this family is important in providing breeding and feeding grounds for numerous animals including economically important crustaceans and fish.

Phylogeny and evolution: Cymodoceaceae are closely related to Ruppiaceae and Potamogetonaceae, refuting the hypothesis that all seagrasses have an exclusive common ancestor. Cymodocea is possibly not monophyletic. The family is known

Genera and species: This family has five genera and 17 species: Amphibolis (2), Cymodocea (4), Halodule (6), Syringodium (2) and Thalassodendron (3).

Etymology: Cymodocea (Κυμόδοκεα) was one of the nymphs in Virgil’s Aeneid, the daughter of Nereus and receiver of waves.

PETROSAVIALES This is an order with little diversification, consisting of only one family Petrosaviaceae. Age estimates vary, but typically are around 100 million years old.

78. PETROSAVIACEAE Oze-so family

This family is composed of pale yellow, mycoheterotropic (Petrosavia) and perennial green (Japonolirion) herbs. They have slender scaly rhizomes and scale-like or linear, alternate or spirally arranged leaves.

Inf lorescences are terminal corymbs or racemes, usually with more than ten flowers. Actinomorphic bisexual flowers arise from axils of small bracts placed on a pedicel, often subtended by a bracteole or bracteole absent (Japonolirion). Six petals are persistent and free or fused at the base, the inner three larger than the outer. Six stamens are fused to the petal segments, and anthers are ovate and dorsifixed or basifixed. The superior or half inferior ovary is composed of three nearly free carpels, fused at the base and free at least in the top half. The ovary is surrounded by septal nectaries and topped by three short styles. The fruit is a capsule composed of three follicles with several elliptic seeds, winged in Petrosavia.

Japonolirion osense, Royal Botanic Gardens, Kew, UK (RW) [78]

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PETROSAVIALES

MONOCOTS

Distribution: Petrosavia occurs in eastern China, Japan, Taiwan, Southeast Asia, Malaysia and Indonesia, often at high elevations. Japonolirion osense is endemic to serpentine swamps in montane central Honshu and Hokkaido. Phylogeny and evolution: Petrosaviaceae are sister to the rest of monocots except Acorales and Alismatales; they are placed

in Petrosaviales. Japonolirion was placed in its own family as well, but molecular studies have shown it to be closely related to Petrosavia, and the families were merged. The crown group of Petrosaviaceae has been dated to c. 123 million years, although other estimates are closer to 100 million years. Petrosavia was previously associated with Triuridaceae, based on their free carpels and mycoheterotrophic habit. They have also

been placed close to or within Melanthiaceae or Nartheciaceae in the past. Genera and species: There are two genera with four species: Japonolirion (1) and Petrosavia (3). Etymology: Petrosavia is named in honour of Italian botanist Pietro Savi (1811–1871), prefect of the Pisa Botanic Garden.

DIOSCOREALES Families 79 to 81 comprise the order Dioscoreales, a clade that is estimated to have diverged c. 123 million years ago. They all have vascular bundles in rings. Glandular hairs in flowers or inflorescences and winged ovaries are also common.

79. NARTHECIACEAE Bog asphodel family

A family of perennial herbs, these plants have creeping rhizomes tipped with (unifacial)

Aletris pauciflora, Yunnan, China [79]

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linear, distichously or spirally arranged leaves. The flowering stem also often has some smaller linear alternate leaves that clasp the base of the stem. Actinomorphic flowers are arranged in terminal racemes, spikes or corymbs subtended by linear or lanceolate bracts. Each flower has a single linear bracteole attached to the pedicel. The six petals are basally fused and persistent in fruit, with the stamens inserted at the base of the petals. Filaments are thread-like and densely hairy in Narthecium, glabrous in the other genera. Anthers are linear and dorsifixed or basifixed. The superior or half inferior

Aletris pauciflora, Yunnan, China [79]

ovary is composed of three fused carpels with a single columnar style and a capitate stigma. The fruit is a loculicidal capsule that splits in three when ripe, exposing many, small, linear or oblong seeds. Distribution: This family occurs patchily across the Northern Hemisphere (California, eastern and southern USA), Guiana Highlands, Western Europe, Caucasus, China, Korea, Japan, Philippines, Borneo and Sumatra. Narthecium ossifragum and some Aletris species form large populations under suitable conditions.

Narthecium ossifragum, Lake District, England, UK [79]

DIOSCOREALES

MONOCOTS

Phylogeny and evolution: Due to their flowers lacking distinctive characters, they were previously associated with a number of other monocot families, including Tofieldiaceae, Petrosaviaceae, Melanthiaceae and Liliaceae. Molecular studies placed this family as sister to the rest of Dioscoreales. Metanarthecium luteoviride may be sister to the rest of the family, and Aletris, with its spirally arranged leaves, is sister to the remaining genera. The crown group diverged c. 76 million years ago. Genera and species: This family has five genera with 35 species: Aletris (24), Lophiola (1), Metanarthecium (1), Narthecium (7) and Nietneria (2).

Gymnosiphon breviflorus, Costa Rica (PM) [80]

Burmannia bicolor, Colombia (MF) [80]

Uses: The bright orange fruits of Narthecium ossifragum have been used as a dye in the Shetland Islands. Unicorn root (Aletris spp.) contains a diosgenin (an oestrogen mimic) that was an important component of Lydia Pinkham’s women’s tonic, a concoction used in the 19th century to relieve menopausal and menstrual pains. Etymology: A narthecium is a Roman perfume box, which in turn is derived from Greek νάρθεξ (narthex), a giant fennel with enclosing scented leaves, a name used by Theophrastos for Ferula communis (Apiaceae). Narthex is now used as a term in architecture for the entrance hall of Byzantine and early Christian churches.

80. BURMANNIACEAE Bluethreads family

This family consists of annual and perennial herbs that are autotrophic, partially heterotropic (but still green) or more commonly fully mycoheterotropic and then differently coloured (blue, purple or white),

Afrothismia korupensis, Cameroon (CD) [80]

Hexapterella gentianoides, Guyana (PM) [80]

Thismia saulensis, French Guiana (PM) [80]

Gymnosiphon usambaricus, Taita Hills, Kenya [80]

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DIOSCOREALES and lacking chlorophyll. They usually have rhizomes that are densely covered with overlapping scale-like leaves and filiform or coral-like roots. Tubers are sometimes also formed. Leaves are alternate, lacking petioles, simple, usually small and scalelike in fully mycoheterotrophic species, or larger, green and forming a basal rosette in autotrophic or partially heterotrophic species. Inflorescences are terminal, bracteate, one to several-flowered cymes that can be contracted into a head or lax and then usually forking. Bisexual flowers are usually actinomorphic, but zygomorphic in Thismieae. The perianth is composed of a tubular basal part and six tepals arranged in two whorls. The floral tube is usually persistent in fruit and often provided with longitudinal ribs or wings, confluent with the frequently three-winged ovary. Three or six stamens are erect or pendulous with bithecal introrse anthers and a widened connective. The three-branched style has a stigma terminating each branch. The inferior ovary is unilocular with septal nectaries. The fruit is a three-winged capsule that splits longitudinally or transversely by slits or irregularly opening due to a decaying fruit wall. Fruits contain numerous, fine seeds.

MONOCOTS

Crown group Burmannieae are dated to c. 75 million years, with Thismieae dated to c. 68 million years. Genera and species: This is a family of 17 genera and 166 species: Burmannieae – Apteria (1), Burmannia (57), Campylosiphon (2), Cymbocarpa (2), Dictyostega (1), Gymnosiphon (30), Hexapterella (2), Marthella (1) and Miersiella (1); Thismieae – Afrothismia (12), Desmogymnosiphon (1), Geomitra (1), Haplothismia (1), Oxygyne (4), Scaphiophora (2), Thismia (47) and Tiputinia (1). Etymology: Burmannia commemorates the Dutch botanist Johannes Burman (Johannus Burmannius, 1707–1780), a contemporary of Carolus Linnaeus. Burman introduced Linnaeus to George Clifford, resulting in a close collaboration between the two. Burman was a specialist on the flora of Sri Lanka, Ambon and the Cape.

81. DIOSCOREACEAE Yam family

Distribution: Pantropical, extending north to the southeastern USA and Japan and south into temperate eastern Australia and northern New Zealand. Phylogeny and evolution: In some phylogenetic analyses Burmanniaceae are sister to Dioscoreaceae, whereas in others Thismieae are closer to Tacca rather than to Burmannieae. Support for these clades is weak in all analyses. They share morphological characters. Tacca can therefore easily be absorbed into either family, or the two families may be combined in the future. This depends on the data obtained by further studies on Dioscoreales. The alternative is to maintain Burmanniaceae but move Thismieae to Dioscoreaceae. In that case there are few characters that separate the two and they could just as well be united.

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This is a family of herbaceous and woody vines and erect perennial herbs with rhizomes or root tubers. Stems are twining or not, sometimes armed with prickles. Petiolate leaves can be alternate, opposite or wholed, with or without stipules. The petiole is usually sheathing at base and often twining (Dioscorea). Leaf axils sometimes bear bulbils. Blades are simple or palmately compound or palmately lobed with palmate venation, basally with three to 13 main veins or pinnate (in some Tacca species) with the secondary venation reticulate. Inflorescences are usually axillary panicles, racemes, or spikes, sometimes reduced to a single flower, in Tacca an umbel with two

whorls of broad or elongate involucral bracts. Flowers are actinomorphic, usually unisexual (Dioscorea) or bisexual (Stenomeris, Tacca, Trichopus). Male flowers have six tepals in two whorls, fused basally or free. The six stamens sometimes have the inner whorl sterile and staminodial or completely absent, the filaments free, fused to tepals or forming a staminodial column. The connective is often broad or appendiculate. Female flowers are similar to the males, with six, three or no staminodes and an inferior, trilocular, often winged ovary, topped by three free styles, the stigmas petal-like in Tacca. The fruit is a capsule, berry or samara, and seeds often have a membranous wing. Distribution: This is a pantropical family extending into temperate North America, Europe, Asia and Australia. Phylogeny and evolution: On morphological grounds, Trichopodaceae and Taccaceae were merged with Dioscoreaceae, and the genus Dioscorea was taken in its broad sense to include Borderea, Epipetrum, Rajania, Tamus and Testudinaria. Most genera share the winged fruits (but not the former berryfruited genus Tamus), the wings formed from the corolla. They share this character with Burmanniaceae, a family close to (or part of) Dioscoreaceae. In some analyses Tacca appears to be more closely related to Thismieae (Burmanniaceae) rather than to Dioscorea. In other analyses Burmanniaceae are sister to the rest, but support is low. It remains to be seen if Dioscoreaceae will need to be expanded in the future. Tacca diverged from the rest of Dioscoreaceae c. 35 million years ago. Genera and species: A family of four genera and 641 species: Dioscorea (622), Stenomeris (2), Tacca (15) and Trichopus (2). Uses: Tubers of several species of Dioscorea are cultivated as yams. Different species have been brought into cultivation independently in different tropical regions. It has been suggested that yams were first domesticated in West Africa c. 11,000 years ago, but other

DIOSCOREALES

MONOCOTS

Dioscorea communis, male flowers, Box Hill, England, UK [81]

Dioscorea hastata, Western Australia [81]

Dioscorea polystachya, aerial tubers, Helsinki Botanical Garden, Finland [81]

Trichopus zeylanicus by M. Smith from Curtis’s Botanical Magazine vol. 120: plate 7350, 1 (1894)

Tacca palmata, Royal Botanic Garden Edinburgh, Scotland, UK [81]

Tacca integrifolia , Singapore (WA) [81]

species were domesticated about the same time in southern China. Dioscorea alata and D. esculenta are cultigens, not known from the wild but originating in Asia. They are now widely cultivated throughout the tropics where they easily naturalise. Other species cultivated as crops include African D. cayennensis and D. dumetorum, Asian D. bulbifera, D. japonica, D. oppositifolia, D. pentaphylla and D. polystachya, and American D. trifida.

In addition, the large quantities of steroidal saponins found in tubers are used as a fish or arrow poison and for washing clothes and tanning. This has now attracted the attention of the pharmaceutical industry for the synthesis of corticosteroids and hormones. The fruits of Trichopus zeylanicus are used in India to make a ginseng-like rejuvenating drink. Tubers of particular cultivars of Tacca leontopetaloides are a staple food in Pacific and Indian Ocean

islands. Bat-flower, Tacca integrifolia, and elephant foot, Dioscorea elephantipes, are occasionally used as houseplants or tropical garden ornamentals.

[81]

Etymology: Dioscorea is named for the ancient Greek physician, pharmacologist and botanist Pedianus Dioscorides (Διόσκορίδες, c. 40–90 AD), author of De Materia Medica, a medical encyclopedia.

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PANDANALES

MONOCOTS

PANDANALES Families 82 to 86 make up the order Pandanales. This clade diverged c. 114 million years ago and can be recognised by its mostly three-ranked leaves (except for mycoheterotrophic Triuridaceae).

Sciaphila albescens, Guyana (PM) [82]

Triuris hexophthalma, male, Guyana (PM) [82]

Triuris hexophthalma, female, Guyana (PM) [82]

82. TRIURIDACEAE

usually unisexual flowers are actinomorphic or zygomorphic, with three to ten tepals in a single series that is usually fused at the base and sometimes has appendages at the apex. The two to six stamens are inserted at the base of the receptacle and have anthers with two or four thecae that mostly open by transverse slits. The connectives are often reduced into long subulate appendages. Female flowers have six to numerous free carpels with a terminal to subbasal style. Fruits are follicles or achenes opening by slits. In Lacandonia the carpels surround the stamens, a rare situation among angiosperms.

anatomy, Tr iu r idaceae have always been difficult to place among monocots. Molecular evidence now supports an association with Pandanales. Molecular age estimates date this family to c. 90 million years ago, which agrees with similarly dated Cretaceous fossils found in New Jersey. Sciaphila may be paraphyletic, based on molecular analyses.

Threetails family

These mycoheterotrophic, perennial, monoecious and dioecious herbs lack chlorophyll and are usually white, yellow, reddish or purple. Stems are erect, growing from a creeping or erect rhizome covered in scale-like leaves. Above-ground leaves are also strongly reduced, alternate and scalelike. The inflorescence is a few to many flowered, bracteate raceme or subcorymb. The 136

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Distribution: This is a pantropical family, but sparse and patchy in Africa and mainland tropical Asia. Phylogeny and evolution: Because of their mycoheterotrophic habit and reduced

Genera and species: This family has nine genera and c. 55 species: Kihansia (1), Kupea (2), Lacandonia (2), Peltophyllum (2), Sciaphila (37), Seychellaria (4), Soridium (1), Triuridopsis (2) and Triuris (4). Etymology: Triuris is composed of the Greek words τρία (tria), three and ουρά (oura), a tail, in reference to the flowers of that genus that have three long-tipped tepals.

PANDANALES

MONOCOTS

83. VELLOZIACEAE Baboon-tail family

This family includes perennial herbs that are sometimes shrubby with fibrous woody stems without secondary growth. The stems are simple or bifurcating and are covered with persistent leaf sheaths. Roots alternate with the leaves and grow down through the leaf sheaths. The leaves are arranged in threes (tristichous) that can spiral around the stem (spirotristichous) and are often clustered at the tips of branches. Blades are needle-shaped (acerose) or flat and linear, often V-shaped and sheathing at the base, and can be persistent or deciduous from the base along a straight line. What appear to be terminal inflorescences become lateral because of the growth of the axis, with a scape and one to many flowers, a raceme in the latter case, or

Vellozia sp., near Mucugê, Bahía, Brazil [83]

(in Acanthochlamys) a compound head on a scape, surrounded by three leaf-like bracts, each flower subtended by three bractlets. Flowers are usually bisexual (unisexual in Barbaceniopsis), actinomorphic and usually brightly coloured. The six tepals are all similar and in two whorls of three, all brightly coloured and long-persisting, fused at the base in most genera, but fused into a tube in Acanthochlamys. Stamens are six or numerous (but then in six bundles) and have filaments that are free or fused with the petals, sometimes with basal appendages. Anthers are basifixed or dorsifixed, the two thecae opening by longitudinal slits. The trilocular, inferior or half inferior ovary is topped with a slender style and three stigmas. The fruit is a capsule that opens by slits or pores.

have been made, and some placements are still uncertain, which is why seven genera are tentatively accepted here. It is certain that Acanthochlamys is sister to Vellozioideae, and since there are many anatomical differences, recognition at subfamilial level (as Acanthochlamydoideae) seems appropriate. The crown group of this family dates back to c. 115 million years. Velloziaceae currently have a Gondwanan distribution, and the age of the lineage indicates that this could be due to vicariance, although long-distance dispersal cannot be excluded.

Distribution: This family is widespread across South America, particularly Brazil, Sub-Saharan Africa, Madagascar and Yemen, with an isolated genus in China.

Uses: Fibre of Xerophyta equisetoides has some local traditional uses in South Africa. Some species such as Talbotia elegans are valuable ornamental plants, although rarely offered in the commercial trade.

Genera and species: This family has seven genera with c. 285 species: Acanthochlamys (1), Barbacenia (108), Barbaceniopsis (4), Nanuza (3), Talbotia (1), Vellozia (126) and Xerophyta (42).

Phylogeny and evolution: There is still disagreement concerning generic delimitation in this family. A consensus has developed to recognise five genera, but not all combinations

Et ymology: Vellozia is named to commemorate Brazilian botanist José Mariano da Conceição Vellozo (1742–1811).

Acanthochlamys bracteata, Yunnan, China (HS) [83]

Talbotia elegans, Helsinki Botanical Garden, Finland [83]

Xerophyta eglandulosa, Isalo National Park, Madagascar (CD) [83]

Barbacenia sp., near Mucugê, Bahía, Brazil [83]

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PANDANALES

84. STEMONACEAE Baibu family

This is a family of twining and erect, sometimes creeping and shrubby perennial herbs with creeping rhizomes or tuberous roots. Leaves are simple, without stipules, and are alternate, opposite or whorled, usually with a basally sheathing petiole. Blades are ovate with an entire margin, and venation is palmate from the base or arching pinnate with a strongly developed midvein, the primary veins connected by transverse parallel veinlets. Inflorescences are axillary and consist of a single flower or lax, few-flowered cymes,

MONOCOTS

sometimes in false umbels or (compound) racemes. Flowers are actinomorphic and usually bisexual (functionally unisexual and somewhat dimorphic in Pentastemona and Stichoneuron). The four or five tepals are free or fused at the base. The stamens are opposite the tepals with filaments usually fused to them or to each other (Pentastemona). Anthers open by longitudinal slits and often have thecae that protrude into sterile appendages that can be fused at their tips, the connective sometimes with a petal-like appendage. The superior or (semi-) inferior ovary has a single locule topped by a sessile one- to four-lobed stigma. Fruits are berries (Pentastemona) or two- or three-valved capsules that usually open to show seeds that dangle on funicles and are provided with fleshy hairs (in Stemona). They are distributed by ants. Distribution: The family is distributed disjunctly in temperate southern USA, and (sub)tropical Asia: Sri Lanka, Southeast Asia, Malesia, southern Japan, New Guinea and northern Australia.

Stemona tuberosa, Kunming Botanical Gardens, China [84]

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Phylogeny and evolution: Because of the climbing habit of some Stemonaceae and their reticulately veined leaves, this family was traditionally associated with Dioscoreaceae. Molecular evidence shows, however, that they are more closely related to Pandanaceae and Cyclanthaceae (Pandanales), with which they share parietal placentation and absence of a style. Pentastemonaceae are embedded in Stemonaceae. They diverged from Pandanaceae/Cyclanthaceae c. 100 million years ago. Genera and species: Stemonaceae have four genera with 37 species: Croomia (6), Pentastemona (2), Stemona (24) and Stichoneuron (5). Uses: Stemona tuberosa is used in traditional Chinese medicine. Etymology: Stemona is derived from Greek στημών (stemon), a stamen or thread, in reference to the conspicuously elongate thecae of some species.

Croomia japonica, Zhejiang, China (CD) [84]

PANDANALES

MONOCOTS

Carludovica palmata, fruit, Royal Botanic Gardens, Kew, UK [85]

Carludovica drudei, Royal Cyclanthus bipartitus, Trésor Voluntary Botanic Gardens, Kew, UK [85] Reserve, French Guiana (CD) [85]

85. CYCLANTHACEAE Panama hat family

Cyclanthaceae consist of perennial, terrestrial and epiphytic, palm-like herbs and vines, with short-creeping and long, slender and climbing, often somewhat woody stems with only primary growth. Leaves are alternate along the rhizome, either spirally or in one plane (distichous), and have sheathing petioles in most species. Blades are usually split into two parts (bifid), sometimes entire or palmately divided. Inflorescences are axillary or terminal and composed of a stalked spadix subtended by two to 11 leaf-like or petal-like spathes that initially envelop the inflorescence. Unisexual f lowers are densely crowded in spirally arranged groups, each consisting of a female flower surrounded by four male flowers, or (in Cyclanthus) in circles around the spadix, the male and female whorls alternating. Male flowers usually surrounded by perianth lobes,

Asplundia rigida, Guadeloupe [85]

in one or two whorls or lacking; in Cyclanthus male flowers are linear rows of stamens. Stamens are usually numerous, with basifixed anthers and filaments that are usually basally fused. Female flowers are free or partially fused with each other. The four tepals are free or fused and often enlarged in fruit. The superior or semi-inferior ovary is surrounded by four staminodes that are elongate and filiform, protruding from the flowers. The tetracarpellate ovary is unilocular and tipped with four stigmas sometimes placed on styles. In Cyclanthus, female flowers are not discrete but rather fused into circles consisting of a double row of pistils, the locules fused into a single ovary chamber, surrounded by a row of reduced tepals and staminodes on either side. Fruits are usually fused into a fleshy syncarp or free. In Cyclanthus the fruit is a dry, cone-like structure with hollow rings filled with seeds. Distribution: This family occurs only in tropical America. Phylogeny and evolution: The family diverged from their relatives in Pandanales c. 77 million years ago. Fossils from the Eocene in Europe have been identified, demonstrating that the family was once present in the Old World. Cyclanthus is usually placed in its

Asplundia insignis, Guadeloupe [85]

own subfamily on anatomical grounds. Genera and species: This family has 12 genera with c. 230 species: Asplundia (c. 100), Carludovica (4), Chorigyne (7), Cyclanthus (2), Diantho-veus (1), Dicranopygium (54), Evodianthus (1), Ludovia (3), Schultesiophytum (1), Sphaera-denia (52), Stelestylis (4) and Thoracocarpus (1). Uses: In several countries in Central and South America, but mostly in Ecuador, the fibre of Carludovica palmata, called toquilla straw, is woven into hats, baskets and mats, and leaves of several other species are locally collected for roof thatching. Hats were imported to Panama for the foreign Canal workers, who gave the name ‘Panama hats’ to these distinctive, flexible straw hats, even though they did not originate in Panama. Young inf lorescences and fr uits of Asplundia, Carludovica and tucuso (Evodianthus funifer) are sometimes eaten by native peoples. Cyclanthus bipartitus and Carludovica palmata are sometimes grown as garden ornamentals in the tropics. Etymology: Cyclanthus is derived from Greek κύκλος (kyklos), circle and άνθος (anthos), flower, in reference to the disc-shaped inflorescence parts.

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PANDANALES

MONOCOTS

f lowers cannot be distinguished. Male flowers have two to numerous, spicately or umbellately aggregated stamens. Anthers open by longitudinal slits. Female flowers have free to fused carpels with a single ovule per carpel and sessile stigmas, appressed to the apex of the carpel, or with a distinct style that becomes spine-like in fruit. Fruits are fibrous drupes or berries, often fused into larger structures that disintegrate when the fruit is ripe.

86. PANDANACEAE Screwpine family

This family of dioecious woody palm-like trees, shrubs and climbers without secondary growth often forms roots from the leaf axils, which may be fully functioning support, stilt or clasping roots, or early aborted and becoming spine-like. The bark is covered in leaf scars. Leaves are spirally arranged in three (spirotristichous) or four (Sararanga) ranks and sessile, lanceolate or linear, sheathing at the rounded to auriculate base, folded lengthwise and M-shaped in crosssection when mature. In most species, margins and midribs are spiny. The (usually) unisexual inflorescences are terminal or axillary spikes, panicles, false umbels or globose heads, usually subtended by variously coloured spathe-like bracts. They are usually erect at first but become pendent when mature. The unisexual flowers are sessile and densely crowded. They lack a perianth, and flowers are fused in Freycinetia, in which individual Pandanus tectorius female inflorescence, Réunion [86]

Distribution: This family is distributed across the Old World tropics from West Africa, Madagascar, the Mascarenes and the Seychelles to India, southern China, the Philippines, Australia, New Zealand and Polynesia. Phylogeny and evolution: The crown group of Pandanaceae started their diversification c. 51 million years ago. Fossil pollen of Pandaniditis has been found in the Late Cretaceous and Early Tertiary of North America but may instead belong to Araceae. Fossil fruits have usually been proven to belong to Nypa (Arecaceae). Fossils older than the Mid Cretaceous are not known. The biogeography of the family has been discussed in the context of the breakup of Gondwana, but this stretches the age of the group too far back, and their occurrence and diversification on recent oceanic volcanic islands may indicate frequent long-distance dispersal. The large genus Pandanus has been

Pandanus tectorius, male tree, Réunion [86]

divided on the basis of molecular study, but these genera are often difficult to distinguish because characters of female fertile material are needed to identify them. Genera and species: Pandanaceae have five genera and c. 770 species: Benstonea (c. 60), Freycinetia (c. 250), Martellidendron (6), Pandanus (c. 450) and Sararanga (2). Uses: Pandan leaves, Pandanus amaryllifolius and P. tectorius to a lesser extent, are used as a spice in Southeast Asian cuisine, especially in rice and curry dishes or mixed with coconut milk. Fruit pulp of kuansu or buah merah, Pandanus conoideus, is economically important in Papua, and c. 30 cultivars are grown in Indonesia. Seeds of P. odoratus (synonym P. dubius) can be eaten raw. Fruits of the thatch screwpine, P. tectorius, are eaten raw or cooked in Micronesia, where they are an important food source. Pollen is fragrant and used for perfume and to flavour drinks. Kiekie, Freycinetia banksii, has sweet fruits and succulent inflorescence bracts that were considered a delicacy by the Maori of New Zealand. Freycinetia arborea, Pandanus tectorius and P. utilis are used for thatch fibre, basketry, weaving, plaiting and rope. Etymology: Pandanus is derived from pandan, the Malay name for the cooking spice, P. amaryllifolius.

Pandanus multispicatus, female inflorescences, Mahé, Seychelles [86]

Freycinetia scandens, Royal Botanic Gardens, Kew, UK [86]

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Pandanus spiralis, female tree, Northern Territory, Australia [86]

LILIALES

MONOCOTS

LILIALES Families 87 to 96 comprise Liliales, a diverse order that includes many geophytes. They have well-defined non-septal nectaries and extrorse anthers. Liliales have been dated to c. 115 million years ago.

87. CAMPYNEMATACEAE Green-mountainlily family

fertilisation. Stamens are inserted at the base of the tepals, and anthers open by longitudinal slits. The trilocular ovary is inferior and bears free styles. Capsular fruits open by decay of the lateral walls or remain unopened.

88. CORSIACEAE Ghost-flower family

Distribution: This family is restricted to Tasmania and New Caledonia. Phylogeny and evolution: The two genera are united in having green tepals, an inferior ovary and free styles. Together with Corsiaceae they form a sister clade to all other Liliales. The genera diverged c. 73 million years ago.

A family of rhizomatous herbs, these perennials have one or several basally clustered flat, linear leaves that are sheathing at the base and often have three teeth at the leaf tip. Inflorescences are contracted panicles that resemble umbels on a bracteate peduncle. Flowers are actinomorphic, each subtended by bracts, and pedicels each bear a bracteole. The bisexual flowers are generally small and green. The six persistent tepals, sometimes bearing nectaries (Campynemanthe), enlarge after

Genera and species: This family has just two genera and four species: Campynema (1) and Campynemanthe (3). Etymology: Campynema is derived from Greek καμπυλός (kampylos), bent or curved and νήμα (nema), a thread.

This family of mycoheterotrophic, perennial herbs lacks chlorophyll. Shoots emerge from a creaping rhizome (or tuber) and are pale red, purple or salmon, but not green. Three to seven alternate leaves are sessile, clasping the stem, and have three to seven parallel veins. Bisexual, zygomorphic flowers are solitary, terminating the stem. The six tepals form two whorls, the three inner and two outer tepals are narrow and thread-like, filiform, lanceolate or somewhat ovate. The upper

Campynemanthe parva, New Caledonia (JM) [87]

Campynemanthe parva, New Caledonia (JM) [87]

Corsia ornata, New Guinea (TF) [88]

Arachnitis uniflora, Chile (PE) [88]

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LILIALES been placed in Burmanniaceae, but molecular evidence has shown that Corsia does not belong in Dioscoreales, but rather has a relationship to Liliales, but their exact position there has been variable and controversial. Arachnitis was placed close to Thismia in one study, and thus Corsiaceae in the traditional sense may be polyphyletic, but more recent phylogenetic studies place this genus together with Corsia in Liliales sister to Campynemataceae. No data are yet available for Corsiopsis.

89. MELANTHIACEAE

Distribution: This family is found in southern South America, the Falkland Islands, southern China, New Guinea, Melanesia and northern Australia (Queensland).

Genera and species: Corsiaceae have three genera and 27 species: Arachnitis (1), Corsia (25) and Corsiopsis (1).

Phylogeny and evolution: Due to the mycoheterotrophic nature of these plants, they are morphologically much reduced, and it has thus been difficult to infer relationships on the basis of morphological characters. They have

Etymology: Corsia was named to commemorate Marquis Bardo Corsi Salviati (1844–1907), who founded a scientifically important collection of plants in greenhouses on his estate near Florence, Italy.

This family of perennial herbs has erect and creeping rhizomes, bulbs and corms. Leaves are in basal rosettes, all along the stem (Veratrum), sometimes with large basal leaves and smaller cauline leaves, or in a whorl at the stem apex, with or without a petiole. They are sometimes sheathing at the base and can have parallel or reticulate venation. Inflorescences are usually racemes,

Toxicoscordion fremontii, Mt Tamalpais, California, USA [89]

Trillium erectum, North Carolina, USA [89]

Pseudotrillium rivale, Royal Botanic Gardens, Kew, UK [89]

Ypsilandra thibetica, private garden, Kingston upon Thames, Surrey, UK [89]

Paris quadrifolia, Warburg Nature Reserve, Bix Bottom, England, UK [89]

outer tepal is broadly ovate and spathe-like, upright with a basal callus or bent downward covering the flower, with two rows of wartlike structures inside. The six stamens have short filaments, and anthers are dorsifixed. The inferior ovary is tricarpelate and bears free or fused styles with free stigmas. The fruit is a capsule that splits into three or opens at the tip only, exposing the numerous dustlike seeds.

142

MONOCOTS

Christenhusz, Fay & Chase

Wake robin family

Xerophyllum tenax, Josephine Lake, Idaho, USA (CD) [89]

LILIALES

MONOCOTS

sometimes spikes, panicles or false umbels, or (in Paris, Pseudotrillium and Trillium) a single sessile or pedicellate flower in the centre of the leaf whorl. Bisexual, usually actinomorphic flowers (zygomorphic in Chionographis) usually have six tepals that are in two whorls, but in Paris can be up to 18, and sometimes only a single whorl of three is present. Free or basally fused tepals can be green and sepallike or variously coloured and petal-like. Usually six (but up to 24) stamens comprising two (to six) whorls, are persistent and have basifixed or dorsifixed anthers that open by longitudinal slits or valves. The superior (sometimes semi-inferior or inferior) ovary is composed of usually three, rarely up to ten, carpels that are fused for more than a third of their length, sometimes completely so, topped with free or fused styles. Fruits are usually dry capsules that open in threes, berry-like fruits that open like a fleshy capsule exposing seeds that sometimes have a bright red aril and fleshy berries that have many seeds. Distribution: Temperate and boreal Northern Hemisphere extending south into Central America and the Andes south to Peru and in Asia into the Himalayas and Taiwan. Phylogeny and evolution: The crown group of Melanthiaceae may be 54 to 42 million years old. Paridoideae separated c. 16 million years ago. Paris is sometimes subdivided, but recognising the genera Daiswa and Kinugasa renders Paris paraphyletic. Paris, Pseudotrillium and Trillium have in the past been segregated as Trilliaceae, but molecular phylogenetic studies have shown that this clade is closely related to Xerophyllum (i.e. embedded in Melanthiaceae). Genera and species: This family includes 17 genera and 173 species in two subfamilies: Melanthioideae – Amianthium (1), Anticlea (11), Chamaelirium (1), Chionographis (6), Helonias (1), Heloniopsis (6), Melanthium (1), Schoenocaulon (26), Stenanthium (3), Toxicoscordion (8), Veratrum (28), Ypsilandra (6) and Zigadenus (1); Paridoideae – Paris (27), Pseudotrillium (1), Trillium (44) and Xerophyllum (2).

Uses: Species of Anticlea, Helonias, Heloniopsis, Paris, Trillium and Veratrum are grown as unusual garden ornamentals, especially in woodland gardens. Some species of Veratrum have been used medicinally, but the alkaloids they contain are highly poisonous. Largest genome: Paris japonica has the largest genome of any organism found to date, only approached in size by the fern Tmesipteris obliqua (Psilotaceae). All members of Paridoideae have large genomes, and the increases in genome size between Xerophyllum (with a small genome) and Pseudotrillium, and between Pseudotrillium and the rest of Paridoideae, are among the largest known. Etymology: Melanthium is derived from Greek μέλας (melas), dark or black and άνθος (anthos), flower, in reference to the dark petal colour of this species.

90. PETERMANNIACEAE Petermann’s-vine family

style and a trilobed, wet stigma. The fruit is a berry with numerous seeds. Distribution: The family is restricted to the central part of eastern Australia (Queensland and New South Wales), roughly between Brisbane and Sydney, where it grows in temperate rainforests. Phylogeny and evolution: Fossil evidence indicates that the family was more widespread in Australian rainforests during the early Eocene. They previously were associated with Philesiaceae or Smilacaceae, and although morphologically there are many similarities, molecular studies have shown that Petermannia forms an independent lineage closer to Colchicaceae and Alstroemeriaceae. In earlier versions of APG it was included in Colchicaceae, but it is genetically and morphologically distinct from that family (e.g. ovary position, anther fixation etc.). Genera and species: This family consists of a single species, Petermannia cirrhosa. Etymology: Petermannia is named for German botanist Wilhelm Ludwig Petermann (1806–1855), who was director of the herbarium and botanical garden at Leipzig. Petermannia cirrhosa, New South Wales, Australia (JB) [90]

This family consists of perennial, woody vines with underground rhizomes and prickly stems, climbing with tendrils that are opposite the leaves. The alternate leaves have petioles and a simple, entire blade with a reticulate secondary venation and pinnateparallel primary venation. Inflorescences are terminal, opposite a leaf and cymose without bracts. The stalked bisexual flowers are actinomorphic and often pendulous. The six tepals comprise two whorls and have a single vein and nectaries at the base. The six stamens have distinct filiform filaments and basifixed anthers that open with longitudinal slits. The unilocular, inferior ovary is composed of three fused carpels and is topped with an elongate

Plants of the World

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LILIALES

MONOCOTS

Bomarea dulcis, Royal Botanic Gardens, Kew, UK [91]

Luzuriaga radicans, Royal Botanic Gardens, Kew, UK [91]

91. ALSTROEMERIACEAE Inca-lily family

This family includes erect or creeping, mostly perennial herbs and vines, the stems evergreen or not. They are usually terrestrial but sometimes grow epiphytically. Rhizomes usually form thick storage roots, except in the single annual species. Leaves are sessile, evenly spread along the stem or crowded at the tip. Veins are parallel or arching, sometimes with reticulate or parallel cross-venation (tessellate), and leaves mostly twisted at the

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Bomarea multiflora, Ecuador [91]

Alstroemeria aurea, Royal Botanic Gardens, Kew, UK [91]

base and the leaf blade becoming partly or totally inverted (resupinate). Inflorescences are terminal, umbel-like cymes with one to many flowers or one or more flowers in bracteate or axillary cincinni. Bracts can be leaf-like or reduced to scales, sometimes in a false whorl. The bisexual flowers are actinomorphic or zygomorphic. The six (rarely eight in Drymophila) free, petal-like tepals are in two rows that can be similar or different, variously marked and with nectaries at the base, the inner ones usually clawed. Six stamens are free with anthers (seemingly) basifixed, opening by longitudinal slits or terminal pores. The inferior ovary is formed from three fused carpels and is uni- or trilocular, topped by a filiform style. Capsular fruits are usually dry and splitting into threes, sometimes explosively, or a berry. Distribution: This family occurs in tropical and temperate Central and South America,

Greater Antilles, Falkland Islands, southeastern Australia, Tasmania and New Zealand. Phylogeny and evolution: Alstroemeriaceae are sister to Colchicaceae but easily distinguished from that family due to their inverted leaves. The two families diverged from other Liliales c. 58 million years ago. The crown group of Alstroemeriaceae is c. 48 million years old, although much older estimates also exist. Drymophila and Luzuriaga have long been of dubious placement. Previously placed in Liliaceae or Philesiaceae, they were sometimes placed in their own family, but because they are sister to Alstroemeriaceae sensu stricto and share many characters such as the twisted (resupinate) leaves APG included them there. Schickendantzia and Taltalia are embedded in Alstroemeria. Leontochir is part of Bomarea. Schickendantziella, sometimes placed here, is a genus of Amaryllidaceae.

LILIALES

MONOCOTS

Genera and species: This family has four genera and 253 species: Alstroemeria (125), Bomarea (122), Drymophila (2) and Luzuriaga (4). Uses: Chuño is the flour extracted from the roots of Alstroemeria ligtu. Bomarea edulis is cultivated for its edible tubers, and consumption of it is widespread across Latin America. Several other species of Alstroemeria and Bomarea also have edible tubers that are locally consumed in South America. Quilineja (Luzuriaga radicans) produces fibre for rope making. Many cultivars of Alstroemeria are grown for the cut-flower market and make good garden plants. Etymology: Alstroemeria was named to commemorate Swedish naturalist Baron Clas Alströmer (1736–1794), a student and friend of Carolus Linnaeus.

Colchicum autumnale, Pyrenees, France [92]

92. COLCHICACEAE Naked-ladies family

This family comprises perennial, erect and climbing herbs with underground corms, tubers and rhizomes; they rarely have slightly woody stems (Kuntheria). Simple, parallel-arched veined leaves are alternately placed along the stem and often in one plane (distichous), sometimes subopposed or verticillate. Blades are sessile, usually sheathing at base (sometimes amplexicaul

Tripladenia cunninghamii, Australian National Botanic Gardens, Canberra [92]

in Uvularia), and the apex can be simply acute or cirrhose with a climbing tip in vining species. Inflorescences are terminal racemes, cymes, umbels or heads, or the flowers are solitary in leaf axils. The bisexual (sometimes unisexual in Wurmbea) flowers are actinomorphic or slightly zygomorphic. The six (rarely seven to 12) tepals are all equal or somewhat unequal, all petal-like, free or partially fused and usually bear nectaries, often clasping the stamens at the base. The six stamens have dorsifixed anthers that open by longitudinal slits. The superior, trilocular ovary is composed of three fused or partially fused carpels, terminating in free or (partially) fused styles. The fruit is usually a septicidal or loculicidal capsule, rarely (in Disporum) a berry. Seeds are distributed by wind or ants. Distribution: Colchicaceae occur in temperate North America, Europe, North

Gloriosa superba, private garden, Kingston upon Thames, Surrey, UK [92]

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LILIALES

Disporum cantoniense, Royal Botanic Gardens, Kew, UK [92]

MONOCOTS

Wurmbea sp. nov. (Christenhusz 6421), Western Australia [92]

Colchicum hantamense, Royal Botanic Gardens, Kew, UK [92]

Africa, the Middle East to Central Asia and Pakistan, Yemen, tropical East Africa, southern Africa, temperate and tropical Asia, from Sakhalin and Japan south to Burma, Malaysia, Luzon and Java, Australia, Tasmania and New Zealand. The family is remarkably absent from South America. Phylogeny and evolution: The concept of Colchicaceae has varied much in the past, and formerly all species were placed in a broad Liliaceae. A number of genera from the former Uvulariaceae are now included. Also, the Old World members of the genus Disporum, formerly in Convallariaceae, are included here. The New World species of Disporum are now Prosartes (Liliaceae), a good example of parallel evolution. Disporopsis belongs to Asparagaceae. Littonia has been merged with Gloriosa,

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Uvularia perfoliata, Royal Botanic Gardens, Kew, UK [92]

Burchardia congesta, Mt Benia, Western Australia [92]

and Neodregea and Onixotis are part of the Wurmbea clade. Bulbocodium and Merendera are part of Colchicum s.l., and this is in turn embedded in Androcymbium. The result is that all are now treated in Colchicum. Two subfamilies are sometimes recognised, but these do not seem to be monophyletic in their current circumscription. The family diverged c. 64 million years ago. Genera and species: Colchicaceae include 15 genera and c. 285 species: Baeometra (1), Burchardia (6), Camptorrhiza (2), Colchicum (159), Disporum (20), Gloriosa (12), Hexacyrtis (1), Iphigenia (12), Kuntheria (1), Ornithoglossum (9), Sandersonia (1), Schelhammera (2), Tripladenia (1), Uvularia (5) and Wurmbea (c. 50). Uses: Due to the toxic alkaloids in most

species, they are not suitable for human or animal consumption. They cause losses to livestock, and accidental or intentional poisonings of humans have also been reported. Colchicine (obtained from wild or cultivated Colchicum) is used in medicine and phar macology and in botanical laboratories to induce polyploidy in plant cells by disrupting cell division. Several species are popular garden or indoor ornamentals, especially Colchicum (autumn crocus, meadow saffron or naked ladies), Gloriosa superba (flame lily) and Uvularia (bellwort), Disporum and Sandersonia aurantiaca (lantern lily). Etymology: Colchicum is named for the Colchis region along the eastern Black Sea in western Asia, probably equivalent to modernday Georgia.

LILIALES

MONOCOTS

93. PHILESIACEAE Chilean-bellflower family

This is a family of much branched, erect shrubs and vines with the stems growing from short woody (primary) rhizomes. Leaves are spirally arranged or placed in one plane (alternate and distichous) and are usually shortly petiolate and more or less sheathing at the base. The simple leaves have one or several parallel main veins with transverse or reticulate secondary veins. Terminal or axillary inflorescences are one- to three-clustered and formed on short-bracted peduncles. Bisexual flowers are actinomorphic and pendulous. Tepals are in two whorls of three, the outer whorl much shorter and more or less sepal-like in Philesia, similar in size to the inner ones in Lapageria. The inner tepals are large, free and overlap at the edges to form tubes, often with nectaries inside. The six stamens are free or fused basally. The nearly basifixed anthers open by longitudinal slits. The superior ovary has a single locule formed of three fused carpels and is topped by an erect filiform style and capitate stigma. The fruit is a red berry with few to many seeds. Lapageria rosea, Royal Botanic Gardens, Kew, UK [93]

Distribution: Philesiaceae occur the cool, temperate southern beech (Nothofagaceae) forests of central and southern Chile and the Magellan Straits.

Etymology: The origin was not given, but it is most probably derived from Greek φιλεώ ( phileo), to love, because of its attractive flowers.

Phylogeny and evolution: Philesiaceae have in the past been placed near Smilacaceae and used to include Eustrephus (Asparagaceae), Geitonoplesium (Asphodelaceae) and Luzuriaga (Alstroemeriaceae). They were considered a bridging group between Asparagales and Dioscoreales, but this circumscription is polyphyletic and polymorphic, differing in a number of characters such as leaf venation, tepal structure, placentation and pollen and seed morphology. Molecular analyses have shown the correct placement of these genera, and Philesiaceae now consist of only two species that are sister to Ripogonaceae (it has been suggested that the two families should be merged), and together they are sister to Liliaceae and Smilacaceae. This clade separated from their relatives c. 50 million years ago.

94. RIPOGONACEAE Supplejack family

Uses: Copihue, Lapageria rosea, is the national flower of Chile and a popular, unusual garden ornamental in temperate humid climates. Fruits of Lapageria are edible. Philesia is also sometimes cultivated, but much less commonly so.

These woody, evergreen shrubs and vines have shoots emerging from a stout horizonal rhizome that can be swollen at the base, forming a woody tuber. The long-twining stems are occasionally spiny and lack tendrils but are covered with sheathing scalelike leaves. When these non-leafy stems reach the light, leafy stems emerge that are branched and spreading. Leaves are mostly opposite, sometimes alternate, usually arranged in a plane (distichous) and often have a well-developed drip-tip. Margins are entire, and leaves are usually petiolate and not sheathing at the base. Inflorescences are terminal or axillary racemes, panicles or spikes, the flowers stalked or sessile with bracts. The bisexual, actinomorphic flowers have six free, petal-like, spreading tepals in two whorls. The six stamens have free

Ripogonum discolor, Queensland, Australia (JC) [94]

Ripogonum scandens, fruit, New Zealand (JC) [94]

Genera and species: This family has two genera, each with a single species: Lapageria rosea and Philesia magellanica. An intergeneric hybrid exists.

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LILIALES

MONOCOTS

Smilax guianensis, showing the two petiolar tendrils, Guadeloupe [95]

Smilax aspera, Sicily, Italy [95]

Smilax megalantha, Royal Botanic Gardens, Kew, UK [95]

filaments and greenish, basifixed anthers opening by longitudinal slits. The superior, trilocular ovary is topped by a single style or the stigma sessile on the ovary. The fruit is a berry.

95. SMILACACEAE Catbrier family

Distribution: This Australasian family is found in New Guinea, eastern Australia and New Zealand. Phylogeny and evolution: Fossil Ripogonum is known from the Miocene in New Zealand and the Eocene in Tasmania. Genera and species: The single genus Ripogonum consists of six species. Uses: Stems of Ripogonum have been sometimes used for basketry and building, and the young shoots can been eaten like asparagus. Etymology: Ripogonum is derived from Greek ριπος (ripos), wickerwork, and γόνυ (gony), a knee or node, referring to the long nodose shoots. It is sometimes spelled ‘Rhipogonum’, an orthographic variant. 148

Christenhusz, Fay & Chase

This family of perennial, vines, shrubs and herbs has erect or spiny stems that are usually prickly. Stems sprout from short fibrous, woody, sometimes tuberous rhizomes. Leaves are alternate, opposite or whorled, petiolate, the petiole mostly bearing a pair of tendrils. The leathery leaves are simple and prominently three- to seven-veined with reticulate secondary venation. The umbellate (rarely racemose or spicate) inflorescences are axillary or terminate a branch. Flowers are actinomorphic and unisexual, the male and female flowers borne on different plants. The six tepals are free or united into a perianth tube and are in two

whorls. Male flowers have three to 12 stamens in two or three whorls, the filaments free or fused and the anthers basifixed and opening by longitudinal slits. Female flowers often have staminodes, the superior ovary is trilocular and composed of three carpels (rarely formed of a single carpel), tipped with three spreading styles. Fruits are black, purple or red berries. Distribution: The family is pantropical, extending into the temperate zones north to boreal North America, Mediterranean Europe, the Russian Far East and south into temperate eastern Australia. Phylogeny and evolution: The family is closely related to and was sometimes included in Liliaceae. They differ mainly in leaf characteristics and in being dioecious. Heterosmilax, which was recognised on the basis of its fused tepals and basally fused stamens (free in Smilax s.s.), has now been merged with Smilax. Mediterranean Smilax aspera appears to be sister to the rest of the genus that otherwise comprises a New World and an Old World clade. Many late Cretaceous,

LILIALES

MONOCOTS

Eocene and Miocene fossils of Smilax are known, but a revision of this material is needed to confirm the identification. Divergence in Smilax started c. 40 million years ago. Genera and species: This family consists of the single genus, Smilax, with c. 260 species.

the classical Greek name for Smilax aspera, but oddly it became the modern Greek word for yew (Taxus).

96. LILIACEAE Lily family

Uses: Sarsaparilla (Smilax aristolochiifolia) gives the flavour to root beer and confectionery but was traditionally used as an antisiphilitic. Smilax megacarpa fruits are consumed on Java, where they are made into jam, and S. china is used medicinally to treat gout. The young shoots of many species are eaten in Asia. Etymology: The name is derived from Greek mythology, about love between the mortal man Κρόκος (Krokos) and the nymph Σμίλαξ (Smilax). This dismayed the gods, and the young man was punished and turned into a saffron flower (hence Crocus sativus, Iridaceae) and the poor nymph into a prickly vine. Smilax also gave her name to Fritillaria imperialis, Royal Botanic Gardens, Kew, UK [96]

This family of perennial herbs grows from underground bulbs and creeping rhizomes. The erect, usually leafy, stems sometimes produce bulbils in leaf axils or on the inflorescence. Leaves are alternate or appearing opposite or verticillate, sometimes in a basal rosette or crowded at the apex of the stem. Blades are linear to oval, usually with an acute tip, cirrhose and climbing in some

Fritillaria, sometimes sheathing at the base or amplexicaul (Streptopus). Venation is parallel (in bulbous genera) or reticulate (in some rhizomatous genera). Inflorescences are often single flowered, but can be arranged in thyrses, racemes or umbels, which are often bracteate. The bisexual flowers are usually actinomorphic, the perianth composed of six free tepals in two whorls, which are usually similar, but sometimes whorls are differentiated. Tepals can vary greatly in colour, often having blotches, spots, streaks, checkering or other marks and bearing nectar glands. Sometimes (e.g. Tricyrtis) tepals are pouched or shortly spurred. The six (rarely three) stamens have free filaments that are usually thin and filiform, but can also be flattened, thickened, often glabrous, but sometimes covered in hairs. The dorsally fixed or (pseudo)basifixed anthers open by longitudinal slits. Ovaries are superior, usually trilocular and lacking septal nectaries. A style is present or absent (e.g. in most Tulipa species, in which the stigma is sessile). Fruits are capsules opening by three locules or fleshy berries.

[96]

Lilium candidum, Bucharest Botanical Garden, Romania [96]

Gagea minima, Finland [96]

Erythronium albidum, Royal Botanic Gardens, Kew, UK [96]

Tulipa suaveolens, Keukenhof, the Netherlands

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LILIALES Distribution: This is mostly a North Temperate family that extends south to North Africa, India, subtropical China, Taiwan and Luzon. It is absent from the Southern Hemisphere. Phylogeny and evolution: Many authors circumscribed Liliaceae broadly in the past, but the members that were previously included can now be found spread across families in Alismatales, Asparagales, Dioscoreales, Liliales, Pandanales and Commelinales. Some authors segregated as separate families the genera with either reticulate venation or secondary growth, but this left obviously unrelated taxa in Liliaceae, and placed unrelated taxa in separate families without their close relatives. Iridaceae and Orchidaceae were never included in Liliaceae s.l., but nearly every other member of the above-mentioned orders was at some point in time. Dahlgren et al. (1968) published the first major classification to break with this tradition, and APG restructured these circumscriptions. The crown group of Liliaceae is estimated Scoliopus bigelovii, Royal Botanic Gardens, Kew, UK [96]

Tricyrtis hirta, Jardin des Plantes, Paris [96]

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to have diverged c. 53 million years ago. Reticulately veined, rhizomatous, berryfruited genera are thought to be plesiomorphic in Liliales, and the large-flowered, bulbous, linear-leaved, capsular-fruited genera are more derived. Some generic reorganisation has also occurred. For instance, Lloydia has been merged with Gagea, Nomocharis is embedded in Lilium, and Amana is kept separate from Tulipa. The New World species of Disporum are not related to the Asian members of this genus, and are now placed in the genus Prosartes in Liliaceae. Asian Disporum keep this name and are placed in Colchicaceae. Genera and species: In its current circumscription, this family includes 15 genera with c. 700 species: Amana (4), Calochortus (73), Cardiocrinum (3), Clintonia (5), Erythronium (32), Fritillaria (c. 140), Gagea (204), Lilium (c. 120), Medeola (1), Notholirion (4), Prosartes (6), Scoliopus (2), Streptopus (10), Tricyrtis (20) and Tulipa (c. 75). Calochortus tolmiei, Sea Ranch, California, USA [96]

Fritillaria meleagris, private garden, Kingston upon Thames, Surrey, UK [96]

Uses: Bulbs of Liliaceae are used as food (Amana, Lilium, Tulipa) or for drugs, especially Fritillaria, known in China as ‘beimu’. Good quality starch can be obtained from Cardiocrinum cordatum and Erythronium japonicum. The flower bulb industry of the Netherlands (and elsewhere) thrives on bulbs of especially tulips (Tulipa) and lilies (Lilium) and to a lesser extent Fritillaria. Tulipa and Lilium are also important in the cut-flower industry. Other genera such as Calochortus (mariposa lily), Cardiocrinum (giant lily), Clintonia (beard lily), Erythronium (dog’s tooth violet) and Tricyrtis (poor-man’s orchid) are popular garden plants in the temperate zones. Etymology: Lilium is the classical Latin name for lily, already in use in Roman times. This name was in turn derived from the Greek λείριων (leirion), a name used for pure white lilies like Lilium candidum. That in turn may have been derived from an extinct eastern Mediterranean language that gave the word hleri in Coptic and hrrt in Egyptian, meaning flower. Cardiocrinum giganteum, Royal Botanic Gardens, Kew, UK [96]

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ASPARAGALES Families 97 to 110 comprise the order Asparagales. This order started to diverge c. 119 million years ago, is highly diverse and encompasses the great majority of monocot species.

97. ORCHIDACEAE Orchid family

This is the largest family of vascular plants. Orchids are generally terrestrial or epiphytic herbs, but there are also vines, bamboolike and even shrubby species, although none is truly woody. They can be green and autotrophic or have lost chlorophyll and become mycoheterotrophic, but many green species also maintain mycorrhizal relationships. Plants are rarely entirely underground (Rhizanthella). Some have slightly woody rhizomes, corms or tubers (no true bulbs are known). Stems are often swollen into pseudobulbs. Leaves are alternate, clasping at base, pointed or irregularly emarginate at the tip, and have an entire margin and parallel (rarely reticulate) venation. Blades are sometimes plicately folded, and leaves are rarely opposite, whorled, scalelike or absent. Inflorescences are racemes, panicles or flowers. The inflorescence is sometimes epiphyllous (emerging from a leaf). Flowers are usually bisexual and zygomorphic, although there are orchids with nearly radial symmetry. The three sepals are free or fused at the base and petal-like or green and sepal-like. The three petals are unequal, one of the three is usually modified into a labellum or lip, a showy ornamented petal that can function as a landing platform, or formed as a pouch to guide pollinating insects. The one to three stamens are at least

fused at the base to the style, although often they are fused with the style into a column. Pollen is usually bound tightly into a few (mostly two to six) masses called pollinia. In most cases, when a pollinator visits an orchid flower, it picks up (and later deposits) all of the pollen in that flower. The inferior ovary is composed of three fused carpels, the entire ovary often twisted, and turning the flower upside-down (resupinate). The style is usually fused with the stamens into a simple or variously ornamented column. Fruits are capsules opening by lateral slits with numerous minute seeds, rarely a fleshy berry. In most cases seeds have no stored food, and when germinating they grow into a protocorm which lives off a fungal connection, allowing the plant to become established. Distribution: Orchids are cosmopolitan, with the greatest diversity in the tropics. Phylogeny and evolution: Orchidaceae evolved during the Late Cretaceous, c. 76–105 million years ago, much earlier than originally thought due to their lack of fossil record. This age makes them one of the 15 oldest families of angiosperms. Herbs generally do not fossilise well, and pollen, which from most plants is generally well preserved, is so highly modified in orchids that it would be difficult to recognise in the fossil record. All five orchid subfamilies evolved before the end of the Cretaceous, which means that orchids and dinosaurs overlapped in existence. This was confirmed by the finding of a c. 15–20 million year old fossil bee (Proplebeia dominicana) embedded in Hispaniolan amber. On its back the bee carried pollinia of an extinct orchid, Meliorchis caribea, belonging to Goodyerinae. Using this fossil in a molecular clock analysis, we can see that extant orchid diversity dates back at least

76–84 million years. Considering the great diversity of animals that pollinate orchids nowadays, one can only wonder if any orchids evolved flowers adapted to be pollinated by dinosaurs before the latter went extinct at the Cretaceous-Tertiary boundary (c. 65 million years ago), after which, admittedly, orchids diversified. How to recognise an orchid: The single character that truly unites all Orchidaceae is the formation of a protocorm. This is an undifferentiated ball of cells that grows from the germinating seeds (that in nearly all species have no stored food). This protocorm connects with hyphae (filaments) of particular species of fungi that are permitted to enter the tissue of the protocorm. The great majority of orchid species then go on to develop normal photosynthetic tissues and become less reliant on their fungal connections. This form of endomycorrhizal relationship appears to be completely to the benefit of the orchid because the plants at this early stage have nothing to offer their fungal host. Later on, orchids may provide their fungal associates with carbon, but some orchids remain dependent on a fungal association. These orchids are not photosynthetic and switch their host to a fungal species that forms ectomycorrhizal connections, which occur commonly in many other groups of flowering plants. Normally, ectomycorrhizal species do exchange carbon for minerals and water from the fungi, but obviously if the orchid species is not photosynthetic then there is nothing to be exchanged; it has been demonstrated that carbon produced by nearby trees ends up in the achlorophyllous orchid, with the ectomycorrhizal fungus acting as the intermediate. This complicated life-history strategy is the hallmark of orchids, and the orchid protocorm is the only truly unique

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ASPARAGALES characteristic (synapomorphy) shared by all orchid species. In more general terms, most botanists recognise an orchid by the presence of a structure in the centre of an orchid flower called the column (or gynostemium), which is the product of fusion of the androecium (reduced to a single fertile anther) and gynoecium (three-parted as in most monocots). The column, however, is not universal; it is missing in subfamily Apostasioideae. In Apostasioideae and Cypripedioideae two or three anthers are present. Pollination: Orchids are well known for their elaborate pollination mechanisms that have evolved to achieve cross-pollination. These have fascinated numerous scientists including Charles Darwin, who studied pollination of orchids in detail and was so enthralled by orchids that his first book after the publication of The origin of species was entirely dedicated to orchids. Most orchids produce pollen in tight bundles (two to six), called pollina. These are often attached to ancillary structures that together are called a pollinarium, which attach the pollinia to the body of the pollinator, usually in a position that makes it difficult for the animal to remove the pollen. Few orchids offer their pollinators a reward, but most orchids look as if they contain a reward; some even produce long spurs although they are devoid of nectar. Rates of visitation are notoriously low, and higher rates of pollination are possible if nectar is artificially added to their spurs. However, this increases the rate of pollination by pollen from flowers on the same plant, and it appears that out-crossing is so advantageous for most orchids that low rates of fruit set are the general rule. Having acknowledged this, the combination of delivery of whole pollinaria with a single visit and a correspondingly large number of ovules in the ovary means that from a single visit a massive number of seeds are produced. Many orchids produce large inflorescences with hundreds of flowers (e.g. some species of Dendrobium, Epidendrum and Oncidium), which seems highly wasteful energetically, but production of mature ovules ready for fertilisation is delayed until pollination takes

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place, thus reducing energy input. Orchid seeds are fungal parasites, and most orchid flowers are likewise pollinator parasites and do not offer a reward for pollination. Being such a large family with highly modified floral structures due to pollinator specialisations, floral diversity is enormous. Orchids make use of nearly every species of pollinator used by other angiosperms, and some orchid pollination mechanisms are truly bizarre. Some orchids mimic other flowers that provide rewards. For instance some South American Gomesa species mimic flowers of Malpighiaceae that produce oil collected by euglossine bees. Gomesa is visited by these bees even though it does not produce these oils, but this is because the mimic is nearly perfect, including the absorbance spectrum. Others are even more specialised mimics, including Australian hammer orchid (Drakaea) species that have a hinged lip that resembles a female thynnid wasp. These wasps have flightless females that are picked up by the males; they mate in flight and females are dropped off to lay eggs. The flower even produces pheromone-like substances similar to those that female wasps produce to lure the males. When the males try to pick up the Drakaea lip, they swing into the pollinia which become stuck to the head of the male wasp. When they fall into the trap a second time the orchid is pollinated. Pollination by all sorts of animals is known in orchids, varying from small insects to birds, lizards and small mammals. There is even a species pollinated by a cricket, a group of animals that are only known otherwise to eat flowers, not to pollinate them. Genera and species: Orchidaceae are the largest family of vascular plants with c. 26,460 species in 750 genera, and the family is divided into five subfamilies (Apostasioideae, Vanilloideae, Cypripedioideae, Epidendroideae and Orchidoideae) that are discussed separately below. The largest genera are Bulbophyllum (1,867), Dendrobium (1,509), Epidendrum (1,413), Lepanthes (1,085), Stelis (879), Habenaria (835), Maxillaria (658), Masdevallia (589), Pleurothallis (552) and Liparis (426).

Uses: A great number of orchid species have minor economic usages, but by far the economically most important species is Vanilla planifolia, indigenous to Mexico and now widely cultivated in the tropics. In the absence of a pollinator outside its natural range, the flowers of vanilla are hand pollinated to produce pods. Vanilla is produced commercially from the fermentation of the nearly mature seed capsules, which contain vanillin. Major production areas for commercial vanilla (Vanilla planifolia) are Madagascar and Réunion (e.g. Bourbon vanilla). Tahitian vanilla (V. ×tahitensis) and West Indian vanilla (V. pompona) are minor spice crops as well, both valuable. Additionally, Leptotes bicolor is grown in Paraguay and Brazil for its vanillin-rich seed pods. An ice-cream-like dessert, dondurma, and a cinnamon-flavoured hot drink, salep, is made from orchid tubers in the eastern Mediterranean, especially from Orchis mascula and other species of tribe Orchideae; it is so popular in some areas that extraction from nature of orchid tubers is endangering many species, at least locally. Chikanda is a bread made from the tubers of wild Disa, Habenaria and Satyrium species in East Africa, mainly Zambia. It is so widely consumed there that it is now causing issues in orchid conservation for the adjacent countries. Many orchids are used in the perfume and cosmetics industry, and sometimes flowers are used to flavour drinks. Jumellea fragrans is used to flavour rum in Réunion, threatening this endemic species with extinction. Some species of Dendrobium are so abundant in India that they are used as cattle-fodder. Other species of Dendrobium and Gastrodia (an obligate mycoheterotroph) are widely used in China and other Asian countries in traditional herbal medicine. Orchids are popular ornamental plants, with hybrids of Cattleya, Cymbidium, Oncidium, Phalaenopsis, Paphiopedilum and Vanda being the most widely grown, but most genera can be found in specialist collections. There are currently over 100,000 cultivars (usually hybrids) in the trade, many of them not officially named. Cultivars of Cymbidium and many other genera make attractive, longlasting cut flowers.

ASPARAGALES

MONOCOTS

Apostasia odorata, Vietnam (JL) [97a]

Vanilla pompona subsp. grandiflora, Helsinki Botanical Garden, Finland (PH) [97b]

Vanilla planifolia by M. Smith from Pogonia rosea, Sarah Duke Botanical Garden, Durham, Curtis’s Botanical Magazine vol. North Carolina, USA [97b] 171, 7167 (1891) [97b]

Paphiopedilum lawrenceanum, Ruissalo Botanical Garden, Turku, Finland [97c]

Paphiopedilum venustum, private garden, Irvine, California, USA [97c]

Phragmipedium kovachii, Ecuagenera Cypripedium calceolus, private garden, Cypripedium reginae, private garden, – Orchids from Ecuador, Ecuador [97c] Kingston upon Thames, Surrey, UK Kingston upon Thames, Surrey, UK [97c]

[97c]

Cypripedium fargesii, Royal Horticultural Society alpine show, London, UK [97c]

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ASPARAGALES Etymology: Orchis is derived from the Greek όρχις (orkhis), meaning testicle, in reference to the shape of the root tubers of many Mediterranean species of subfamily Orchidoideae. They are still used as an aphrodisiac in the eastern Mediterranean.

97a. ORCHIDACEAE SUBFAMILY APOSTASIOIDEAE Apostasioids This subfamily consists of terrestrial, rhizomatous herbs, with broad, often plicate leaves. The inflorescences are usually erect racemes or spikes. The flowers differ from standard orchid flowers by having two or three stamens with powdery pollen that is not bound into pollinia. The stamens and pistils are only partially fused into a column. The inferior ovary becomes a dry capsule in fruit. Distribution: This subfamily is restricted to tropical Asia and northern Australia. Phylogeny and evolution: This pair of small genera has often been cited as primitive members of the family, and their floral traits are suggestive of this. However, relative to their chromosome numbers and vegetative features, they are highly derived. In terms of habit, these rhizomatous plants with broad, often plicate leaves are more similar to the related families in Asparagales than to Orchidaceae, especially Hypoxidaceae, with which they are often confused in the vegetative state. Apostasioideae are nevertheless well supported as sister to the rest of orchids and also have a protocorm, a unique character of orchids. Some authors have placed these two genera in their own family, Apostasiaceae, but given their phylogenetic position there is nothing gained by this placement. Genera and species: There are two genera with 14 species: Apostasia (6) and Neuwiedia (8). Etymology: Apostacy, a revolt or departure, is derived from Greek απόστασης (apostasis), distance, refering to the unusual f loral structure (for an orchid, at least) of these plants, which was the basis for some to

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‘distance’ them from orchids and to argue for separate family status.

97b. ORCHIDACEAE SUBFAMILY VANILLOIDEAE Vanillas Many Vanilloideae are vines, some without chlorophyll, but several are terrestrial herbs. All have typical orchid flowers, in that they have a lip and column, the latter with a single anther. Flowers persist on the tip of the ovary long after flowering, and there is often a flange of tissue at the apex of the ovary that persists even longer. Most members of this subfamily also have micro-seeds typical of most orchids, but some genera, such as Vanilla, have larger crustose seeds that resemble those of Hypoxidaceae. This development is presumably a response to the seed dispersal strategy of these genera, which is based on consumption by animals of the highly fragrant capsules (vanillin is released by fermentation while the fruit is attached to the plant); the seeds have to survive the gut of the animal and thus need more protection than the dust-like, thin-walled seeds of most other winddispersed orchids. Distribution: This family is pantropical with a few genera in eastern North America and Japan. Phylogeny and evolution: Vanilloideae are positioned as sister to all other orchids except Apostasioideae, leaving us with the impression that reduction from three to one anther happened more than once in the evolutionary history of Orchidaceae. They are the most recently recognised of the orchid subfamilies, although the family Vanillaceae was already described. Their possession of a single anther led most taxonomists to consider them members of the same subfamily (Epidendroideae) as most tropical epiphytic orchids. Genera and species: Vanilloideae include 14 genera and 245 species: Cleistes (64), Cleistesiopsis (2), Clematepistephium (1), Cyrtosia (5), Duckeella (3), Epistephium (21), Eriaxis (1), Erythrorchis (2), Galeola (6), Isotria (2), Lecanorchis (20), Pogonia (5), Pseudovanilla (8) and Vanilla (105).

Etymology: Vanilla is the diminutive of the Spanish ‘vaina’, a pod, in reference to the production of vanilla from the seed capsule.

97c. ORCHIDACEAE SUBFAMILY CYPRIPEDIOIDEAE Ladies’ slippers These herbaceous perennial, terrestial or epilithic plants, sometimes tall and reed-like (Selenipedium), have a highly constrained floral morphology that makes them unmistakable. They have a pouched lip that functions to trap pollinators inside, only permitting their exit via two openings on either side of the column, which has two anthers that partially block these exits, thereby bringing the pollinator (bees, flies or beetles) into contact with their pollen masses, which are greasy and mostly not formed into tight pollinia (but there are exceptions). The combination of this particular lip structure and two anthers (with a third sterile staminode) is the hallmark of Cypripedioideae. Fruits of Selenipedium are fleshy and ferment to release small amounts of vanillin, not enough for human exploitation, but enough to earn one species the common name of vainilla chica or little vanilla. Other Cypripedioideae have capsular, dry fruits. Distribution: This subfamily occurs in the Northern Hemisphere, tropical America, tropical Asia. Phylogeny and evolution: Cypripedioideae are sister to the remaining two subfamilies, O rch idoidea e a nd Epide nd roidea e. Collectively, the genera have a distribution that probably reflects their origin before the end of the Cretaceous and a Gondwanan association, although they are absent from Africa, presumably due to extinction during the dry periods experienced by this continent during its earlier history. Genera and species: This subfamily includes five genera with 169 species: Cypripedium (51), Mexipedium (1), Paphiopedilum (86), Phragmipedium (26) and Selenipedium (5). Etymology: Cypripedium is derived from Cypris (the Lady of Cyprus) an alternative

ASPARAGALES

MONOCOTS

name for the Greek goddess Aphrodite, and Greek πεδίλων (pedilon), a sandal.

97d. ORCHIDACEAE SUBFAMILY EPIDENDROIDEAE Epidendroids This is by far the largest subfamily of orchids and is nearly impossible to characterise due to the diversity of its members. They include herbs with underground tubers, vines, achlorophyllous fungal holoparasites and a huge number of epiphytic species, some of which have swollen stems (pseudobulbs), and others that are seasonally or habitually leafless (and photosynthesis takes place in their green, exposed roots). Florally, they all have the typical orchid flower, but some have mealy pollen that is not transferred as a unit, whereas others have complicated pollen-bearing ancillary structures, making these the most advanced pollen transfer mechanisms known in flowering plants. No other prominent features are useful for recognising this subfamily; they are distinguished from the second-largest subfamily, Orchidoideae, by their fibrous, often evergreen leaves. Distribution: This cosmopolitan subfamily is most diverse and numerous in the wet tropics at moderate to high elevations. Phylogeny and evolution: Orchidoideae and Epidendroideae are sister taxa; this last major split in the family also took place before the end of the Cretaceous. Unlike many families of flowering plants, the major groups of this subfamily, divided into tribes and subtribes, are restricted to hemispheres and continents, perhaps reflecting patterns of mycorrhizal relationships, particularly during the early stages of germination and establishment. Unfortunately, we still know relatively little about patterns of fungal association among tropical epiphytic species. Genera and species: Currently there are 520 accepted genera and c. 21,100 species: Acampe (8), Acanthephippium (13), Acianthera (118), Acineta (17), Acriopsis (9), Acrolophia (7), Acrorchis (1), Adamantinia

(1), Adenoncos (17), Adrorhizon (1), Aerangis (58), Aeranthes (43), Aerides (25), Aetheorhyncha (1), Aganisia (4), Aglossorrhyncha (13), Agrostophyllum (100), Alamania (1), Alatiliparis (5), Ambrella (1), Amesiella (3), Anathallis (152), Ancistrochilus (2), Ancistrorhynchus (17), Andinia (13), Angraecopsis (22), Angraecum (221), Anguloa (9), Ania (11), Ansellia (1), Anthogonium (9), Aphyllorchis (22), Aplectrum (1), Appendicula (146), Arachnis (14), Arethusa (1), Arpophyllum (3), Artorima (1), Arundina (2), Ascidieria (8), Aspasia (7), Auxopus (4), Barbosella (19), Barkeria (17), Basiphyllaea (7), Batemannia (5), Beclardia (2), Benzingia (9), Biermannia (11), Bifrenaria (21), Bletia (33), Bletilla (5), Bogoria (4), Bolusiella (6), Brachionidium (75), Brachypeza (10), Bracisepalum (2), Braemia (1), Brassavola (22), Brassia (64), Bromheadia (30), Broughtonia (6), Bryobium (8), Bulbophyllum (1867), Bulleyia (1), Calanthe (216), Callostylis (5), Calopogon (5), Caluera (3), Calymmanthera (5), Calypso (1), Calyptrochilum (2), Campanulorchis (5), Campylocentrum (65), Capanemia (9), Cardiochilos (1), Catasetum (176), Cattleya (112), Cattleyella (1), Caucaea (9), Caularthron (4), Centroglossa (5), Cephalanthera (19), Cephalantheropsis (4), Ceratocentron (1), Ceratostylis (147), Chamaeanthus (3), Chamelophyton (1), Changnienia (1), Chaubardia (3), Chaubardiella (8), Chauliodon (1), Cheiradenia (1), Chelonistele (13), Chiloschista (20), Chondrorhyncha (7), Chondroscaphe (14), Chroniochilus (4), Chrysoglossum (4), Chysis (10), Chytroglossa (3), Cirrhaea (7), Cischweinfia (11), Claderia (2), Cleisocentron (6), Cleisomeria (2), Cleisostoma (88), Cleisostomopsis (2), Clowesia (7), Cochleanthes (4), Coelia (5), Coeliopsis (1), Coelogyne (200), Collabium (14), Comparettia (78), Conchidium (10), Constantia (6), Corallorhiza (11), Coryanthes (59), Corymborkis (6), Cottonia (1), Cremastra (4), Crepidium (260), Cribbia (4), Crossoglossa (26), Crossoliparis (1), Cryptarrhena (3), Cryptochilus (5), Cryptopus (4), Cryptopylos (1), Cuitlauzina (7), Cyanaeorchis (3), Cycnoches (34), Cymbidiella (3), Cymbidium (71), Cypholoron (2), Cyrtochiloides (3), Cyrtochilum (137),

Cyrtopodium (47), Cyrtorchis (18), Dactylostalix (1), Daiotyla (4), Danxiaorchis (1), Deceptor (1), Dendrobium (1509), Dendrochilum (278), Dendrophylax (14), Devogelia (1), Diaphananthe (33), Diceratostele (1), Dichaea (118), Dickasonia (1), Didymoplexiella (8), Didymoplexis (17), Dienia (6), Diglyphosa (3), Dilochia (8), Dilochiopsis (1), Dilomilis (5), Dimerandra (8), Dimorphorchis (5), Dinema (1), Dinklageella (4), Diodonopsis (5), Diplocentrum (2), Diploprora (2), Dipodium (25), Distylodon (1), Domingoa (4), Draconanthes (2), Dracula (127), Dresslerella (13), Dressleria (11), Dryadella (54), Dryadorchis (5), Drymoanthus (4), Dunstervillea (1), Dyakia (1), Earina (7), Echinorhyncha (5), Echinosepala (11), Eclecticus (1), Eggelingia (3), Eleorchis (1), Elleanthus (111), Eloyella (10), Embreea (2), Encyclia (165), Entomophobia (1), Ephippianthus (2), Epiblastus (22), Epidendrum (1413), Epilyna (2), Epipactis (49), Epipogium (3), Erasanthe (1), Eria (237), Eriodes (1), Eriopsis (5), Erycina (7), Eulophia (200), Eulophiella (5), Euryblema (2), Eurychone (2), Fernandezia (51), Frondaria (1), Galeandra (38), Galeottia (12), Gastrochilus (56), Gastrodia (60), Gastrorchis (8), Geesinkorchis (4), Geodorum (12), Glomera (131), Gomesa (119), Gongora (74), Govenia (24), Grammangis (2), Grammatophyllum (12), Grandiphyllum (7), Graphorkis (4), Grobya (5), Grosourdya (11), Guanchezia (1), Guarianthe (4), Gunnarella (9), Gynoglottis (1), Hagsatera (2), Hammarbya (1), Hancockia (1), Hederorkis (2), Helleriella (2), Hexalectris (10), Hintonella (1), Hippeophyllum (10), Hoehneella (2), Hofmeisterella (2), Holcoglossum (14), Homalopetalum (8), Horichia (1), Horvatia (1), Houlletia (9), Huntleya (14), Hymenorchis (12), Imerinaea (1), Ionopsis (6), Ipsea (3), Isabelia (3), Ischnogyne (1), Isochilus (13), Ixyophora (5), Jacquiniella (12), Jejewoodia (6), Jumellea (59), Kefersteinia (70), Kegeliella (4), Koellensteinia (17), Kraenzlinella (9), Lacaena (2), Laelia (23), Lemurella (4), Lemurorchis (1), Leochilus (12), Lepanthes (1085), Lepanthopsis (43), Leptotes (9), Limodorum (3), Liparis (426), Listrostachys (1), Lockhartia (28), Loefgrenianthus (1), Lueckelia (1), Lueddemannia (3), Luisia (39), Lycaste (32), Lycomormium (5),

Plants of the World

155

ASPARAGALES

MONOCOTS

Cymbidium devonianum, Royal Horticultural Society Garden, Wisley, UK [97d]

Dendrobium nobile, Helsinki Botanical Garden, Finland [97d]

Bulbophyllum thaiorum, New York Botanical Garden, USA [97d]

Phaius tankervilleae, Royal Botanic Gardens, Kew, UK [97d]

Arundina graminifolia, Singapore Botanical Garden [97d]

Vanda tricolor var. suavis, Santa Barbara Orchid Estate, California, USA [97d] Calypso bulbosa, Mt Tamalpais, California, USA [97d]

Dracula verticulosa, Ecuagenera – Orchids from Ecuador, Ecuador [97d]

Pleurothallis gargantua, Ecuagenera – Orchids from Ecuador, Ecuador [97d]

Restrepia guttulata, Ecuagenera – Orchids from Ecuador, Ecuador [97d]

Oncidium reflexum, Mexico [97d]

Cattleya sincorana, Mucugê, Bahía, Brazil [97d]

156

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Epipactis palustris, Royal Botanic Gardens, Kew, UK [97d]

Porroglossum muscosum, Ecuagenera Brassavola nodosa, New York – Orchids from Ecuador, Ecuador [97d] Botanical Garden, USA [97d]

Cleisostoma birmanicum, New York Botanical Garden, USA [97d]

Kefersteinia trullata, Ecuagenera – Orchids from Ecuador, Ecuador [97d]

ASPARAGALES

MONOCOTS

Ophrys speculum, Sicily, Italy [97e] Disa uniflora, Hampton Court Flower Show, UK [97e] Aa colombiana, Ecuador [97e]

Anacamptis papilionacea, Sicily, Italy [97e]

Habenaria medusa, University of Wisconsin, Madison, USA [97e]

Caladenia flava, near Perth, Western Australia [97e]

Thelymitra variegata, Geraldton, Western Australia [97e]

Orchis militaris, Homefield Wood, England, UK [97e]

Drakaea glyptodon, near Albany, Western Australia [97e]

Pterostylis aspera, Geraldton, Western Australia [97e]

Rhizanthella gardneri, Western Australia (KD) [97e]

Plants of the World

157

ASPARAGALES Macradenia (11), Macroclinium (42), Macropodanthus (8), Malaxis (182), Margelliantha (6), Masdevallia (589), Maxillaria (658), Mediocalcar (17), Meiracyllium (2), Microcoelia (30), Microepidendrum (1), Micropera (21), Microsaccus (12), Miltonia (12), Miltoniopsis (5), Mobilabium (1), Monophyllorchis (1), Mormodes (80), Mycaranthes (36), Myoxanthus (48), Myrmecophila (10), Mystacidium (10), Nabaluia (3), Nemaconia (6), Neobathiea (5), Neoclemensia (1), Neocogniauxia (2), Neogardneria (1), Neogyna (1), Neomoorea (1), Neottia (64), Nephelaphyllum (11), Nephrangis (2), Nervilia (67), Nidema (2), Nohawilliamsia (1), Notheria (15), Notylia (56), Notyliopsis (2), Oberonia (323), Oberonioides (2), Octarrhena (52), Octomeria (159), Oeceoclades (38), Oeonia (5), Oeoniella (2), Oestlundia (4), Oliveriana (6), Omoea (2), Oncidium (311), Ophioglossella (1), Oreorchis (16), Orestias (4), Orleanesia (9), Ornithocephalus (55), Ossiculum (1), Otochilus (5), Otoglossum (13), Otostylis (4), Oxystophyllum (36), Pabstia (5), Pabstiella (29), Pachystoma (3), Palmorchis (21), Panisea (11), Paphinia (16), Papilionanthe (11), Papillilabium (1), Paradisanthus (4), Paralophia (2), Paraphalaenopsis (4), Pelatantheria (8), Pennilabium (15), Peristeranthus (1), Peristeria (13), Pescatoria (23), Phaius (45), Phalaenopsis (70), Phloeophila (11), Pholidota (39), Phragmorchis (1), Phreatia (211), Phymatidium (10), Pilophyllum (1), Pinalia (105), Platyrhiza (1), Platystele (101), Plectorrhiza (3), Plectrelminthus (1), Plectrophora (10), Pleione (21), Pleurothallis (552), Pleurothallopsis (18), Plocoglottis (41), Poaephyllum (6), Podangis (1), Podochilus (62), Pogoniopsis (2), Polycycnis (17), Polyotidium (1), Polystachya (234), Pomatocalpa (25), Ponera (2), Porpax (13), Porphyroglottis (1), Porroglossum (43), Porrorhachis (2), Promenaea (18), Prosthechea (117), Pseuderia (20), Pseudolaelia (18), Psilochilus (7), Psychilis (14), Psychopsiella (1), Psychopsis (4), Pteroceras (27), Pterostemma (3), Pygmaeorchis (2), Quekettia (4), Quisqueya (4), Rangaeris (6), Rauhiella (3), Renanthera (20), Restrepia (53), Restrepiella (2), Rhaesteria (1), Rhinerrhiza (1), Rhinerrhizopsis (3), Rhipidoglossum (35), Rhynchogyna (3),

158

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MONOCOTS

Rhyncholaelia (2), Rhynchostele (17), Rhynchostylis (3), Ridleyella (1), Risleya (1), Robiquetia (45), Rodriguezia (48), Rossioglossum (9), Rudolfiella (6), Saccolabiopsis (14), Saccolabium (5), Sanderella (2), Santotomasia (1), Sarcanthopsis (5), Sarcochilus (25), Sarcoglyphis (12), Sarcophyton (3), Sarcostoma (5), Saundersia (2), Scaphosepalum (46), Scaphyglottis (69), Schistotylus (1), Schlimia (7), Schoenorchis (25), Schunkea (1), Scuticaria (11), Seegeriella (2), Seidenfadenia (1), Seidenfadeniella (2), Sertifera (7), Sievekingia (16), Singchia (1), Sirhookera (2) Smithsonia (3), Smitinandia (3), Sobennikoffia (4), Sobralia (149), Solenangis (8), Solenidium (3), Soterosanthus (1), Spathoglottis (48), Specklinia (135), Sphyrarhynchus (1), Spongiola (1), Stanhopea (61), Stelis (879), Stenia (22), Stenotyla (9), Stereochilus (7), Stereosandra (1), Stichorkis (8), Stolzia (15), Suarezia (1), Sudamerlycaste (42), Summerhayesia (2), Sutrina (2), Systeloglossum (5), Taeniophyllum (185), Taeniorrhiza (1), Tainia (23), Tamayorkis (1), Taprobanea (1), Teagueia (13), Telipogon (205), Tetramicra (14), Teuscheria (7), Thaia (1), Thecopus (2), Thecostele (1), Thelasis (26), Thrixspermum (161), Thunia (5), Thuniopsis (1), Thysanoglossa (3), Tipularia (7), Tolumnia (27), Tomzanonia (1), Trachoma (14), Trevoria (5), Triceratorhynchus (1), Trichocentrum (70), Trichoceros (10), Trichoglottis (69), Trichopilia (44), Trichosalpinx (111), Trichotosia (78), Tridactyle (47), Triphora (18), Trisetella (23), Trizeuxis (1), Tropidia (31), Tuberolabium (11), Uleiorchis (2), Uncifera (6), Vanda (73), Vandopsis (4), Vargasiella (1), Vasqueziella (1), Vitekorchis (4), Warczewiczella (11), Warmingia (4), Warrea (3), Warreella (2), Warreopsis (4), Wullschlaegelia (2), Xerorchis (2), Xylobium (30), Yoania (4), Ypsilopus (5), Yunorchis (1), Zelenkoa (1), Zootrophion (22), Zygopetalum (14), Zygosepalum (8) and Zygostates (22).

Phylogeny and evolution: As indicated above, Orchidoideae are sister to Epidendroideae and parallel their patterns of distribution, except that this subfamily has specialised to a greater degree in temperate zones, both northern and southern. Previous taxonomists recognised subfamily Spiranthoideae as distinct from Orchidoideae, based largely on the replacement of root tubers found predominantly in the latter with the fat roots of the former, but molecular phylogenetic studies demonstrated that Spiranthoideae are embedded in Orchidoideae, leading to recognition of an expanded Orchidoideae.

Etymology: Epidendrum is composed of the Greek prefix επί (epi) meaning ‘on’ and δένδρων (dendron), a tree, referring to the epiphytic habit of many species of Epidendrum, the single genus into which Linnaeus originally placed all tropical epiphytic orchids.

Genera and species: Currently there are 217 accepted genera and 4,965 species: Aa (25), Aceratorchis (1), Achlydosa (1), Acianthus (20), Adenochilus (2), Aenhenrya (1), Altensteinia (7), Amitostigma (27), Amitostigma (27), Anacamptis (11), Androcorys (10), Anoectochilus (43), Aporostylis (1), Aracamunia (1), Arthrochilus

97e. ORCHIDACEAE SUBFAMILY ORCHIDOIDEAE Orchidoids The habits of this subfamily are less diverse than those of their sister clade, Epidendroideae. The majority of Orchidoideae are deciduous or soft-leaved herbs (without fibres) with root tubers or fat elongate roots. Like Epidendroideae, these plants have typical orchid flowers adapted for pollination by a wide variety of animals. Pollen is in general not bound into tight masses, and in some cases multiple visits have been recorded before most pollen is removed from a flower. The majority of the species use deceit-pollination without reward to the pollinator, and in both the North and South Temperate zones genera have become adapted for pollination by male insects that mistake the flowers for female members of their species; floral scents of such species also mimic the sex pheromones of female insects. Fruits are generally dry, many-seeded capsules. Distribution: This cosmopolitan group has centres of diversity in Australia, South Africa and the Mediterranean.

ASPARAGALES

MONOCOTS

(15), Aspidogyne (45), Aulosepalum (7), Bartholina (2), Baskervilla (10), Beloglottis (7), Benthamia (29), Bhutanthera (5), Bidoupia (1), Bipinnula (11), Bonatea (13), Brachycorythis (36), Brachystele (21), Brownleea (8), Buchtienia (3), Burnettia (1), Caladenia (267), Caleana (1), Calochilus (27), Centrostigma (3), Ceratandra (6), Chamaegastrodia (3), Chamorchis (1), Cheirostylis (53), Chiloglottis (23), Chloraea (52), Coccineorchis (7), Codonorchis (1), Coilochilus (1), Corybas (132), Corycium (15), Cotylolabium (1), Cranichis (53), Cryptostylis (23), Cyanicula (10), Cybebus (1), Cyclopogon (83), Cynorkis (156), Cyrtostylis (5), Cystorchis (21), Dactylorhiza (40), Danhatchia (1), Degranvillea (1), Deiregyne (18), Dichromanthus (4), Diplomeris (3), Disa (182), Discyphus (1), Disperis (78), Diuris (71), Dossinia (1), Dracomonticola (1), Drakaea (10), Eltroplectris (13), Elythranthera (2), Epiblema (1), Ericksonella (1), Eriochilus (9), Erythrodes (26), Eurycentrum (7), Eurystyles (20), Evotella (1), Fuertesiella (1), Funkiella (27), Galearis (10), Galeoglossum (3), Galeottiella (6), Gavilea (15), Gennaria (1), Genoplesium (47), Glossodia (2), Gomphichis (24), Gonatostylis (2), Goodyera (98), Greenwoodiella (3), Gymnadenia (23), Habenaria (835), Halleorchis (1), Hapalorchis (10), Helonoma (4), Hemipilia (13), Herminium (19), Herpysma (1), Hetaeria (29), Himantoglossum (11), Holothrix (45), Hsenhsua (1), Huttonaea (5), Hylophila (7), Kionophyton (4), Kipandiorchis (1), Kreodanthus (14), Kuhlhasseltia (9), Lankesterella (11), Lepidogyne (1), Leporella (1), Leptoceras (1), Ligeophila (12), Ludisia (1), Lyperanthus (2), Lyroglossa (2), Macodes (11), Manniella (2), Megalorchis

(1), Megastylis (7), Mesadenella (7), Mesadenus (7), Microchilus (137), Microtis (19), Myrmechis (17), Myrosmodes (12), Neobolusia (3), Neotinea (4), Nothostele (2), Odontochilus (25), Odontorrhynchus (6), Oligophyton (1), Ophrys (34), Orchipedum (3), Orchis (21), Orthoceras (2), Pachites (2), Pachyplectron (3), Papuaea (1), Paracaleana (13), Pecteilis (8), Pelexia (77), Peristylus (103), Pheladenia (1), Physoceras (12), Physogyne (3), Platanthera (136), Platycoryne (19), Platylepis (17), Platythelys (13), Ponerorchis (55), Ponthieva (66), Porolabium (1), Porphyrostachys (2), Praecoxanthus (1), Prasophyllum (131), Prescottia (26), Pseudocentrum (7), Pseudogoodyera (1), Pseudorchis (1), Pterichis (20), Pteroglossa (11), Pterostylis (211), Pterygodium (19), Pyrorchis (2), Quechua (1), Rhamphorhynchus (1), Rhizanthella (3), Rhomboda (22), Rimacola (1), Roeperocharis (5), Sacoila (7), Sarcoglottis (48), Satyrium (86), Sauroglossum (11), Schiedeella (22), Schizochilus (11), Schuitemania (1), Serapias (13), Shizhenia (1), Silvorchis (3), Sirindhornia (3), Skeptrostachys (13), Solenocentrum (4), Sotoa (1), Spiculaea (1), Spiranthes (34), Stalkya (1), Stenoglottis (7), Stenoptera (7), Stenorrhynchos (5), Stephanothelys (5), Steveniella (1), Stigmatodactylus (10), Svenkoeltzia (3), Thelymitra (110), Thelyschista (1), Thulinia (1), Townsonia (2), Traunsteinera (2), Tsaiorchis (1), Tylostigma (8), Veyretella (2), Veyretia (11), Vrydagzynea (43), Waireia (1) and Zeuxine (74). Etymology: Greek όρχις (orkhis) means testicle, refering to the shape of the root tubers of many Mediterranean species of subfamily Orchidoideae.

Borya sphaerocephala, Geraldton, Western Australia [98]

98. BORYACEAE Pincushion-lily family

This is a family of perennial shrubby herbs forming tufts, clumps or cushions. The woody stems are branched and form fibrous, wiry, sometimes stilt-like roots. Sessile linear leaves are placed spirally around the stem, are sheathing at the base and often have a sharp tip. Venation is parallel. Inflorescences are often dense, terminal spikes or racemes, surrounded by involucral bracts, often condensed to appear capitate. White flowers are bisexual and actinomorphic, with two whorls of three similar, free or basally fused tepals. The six stamens are free or fused with the tepals, and the basifixed anthers open by longitudinal slits. The superior ovary is trilocular and bears septal nectaries that resemble external nectar slits on the ovary wall. The ovary is topped with a single style and a capitate stigma. The fruit is a capsule that splits in three. Distribution: This family is endemic to Australia. Borya are resurrection plants found in exposed xeric environments where they have a unique adaptation to desiccation, at which stage they turn rusty reddish-orange

Borya sphaerocephala, Perth, Western Australia [98]

Plants of the World

159

ASPARAGALES only to become green and start growing again when rain occurs. By contrast, Alania is found in constantly moist habitats. Phylogeny and evolution: The crown group of Boryaceae is dated to be 54 million years old. The family was previously associated with Anthericaceae or Asparagaceae, but molecular analyses place Boryaceae as sister to a clade that includes Asteliaceae and Hypoxidaceae. Genera and species: This family has two genera with a dozen species: Alania (1) and Borya (11). Etymology: The Australian genus Borya was named to commemorate French naturalist Jean Baptiste Bory de Saint-Vincent (1778–1846), who embarked on an expedition to Australia in 1798, but disembarked in Mauritius and explored the Mascarene Islands instead.

99. BLANDFORDIACEAE Christmas-bells family

MONOCOTS

These perennial herbs grow in tufts from short tuber-like rhizomes. The alternate sessile leaves are placed in a fan (distichous) and sheath the rhizome at the base. Blades are linear with many parallel veins, secondary crossveins and a prominent midvein. Inflorescences grow terminally from the rhizome and are racemes that in some species can grow up to 1.5 m tall. The bisexual flowers are actinomorphic to slightly zygomorphic and pedicels are enveloped by a scale-like bract. The petal-like tepals are fused to form a bell-shaped tube with six broad lobes. The six stamens are attached to the lower part of the perianth tube, and anthers are dorsifixed and open by latrorse slits. A trilocular superior ovary tapers below to form a stalk (gynophore), and septal nectaries are present as external grooves. The ovary is tipped with a single erect style and a three-grooved stigma. The fruit is a capsule that splits in three.

160

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Genera and species: The single genus Blandfordia consists of four species. Uses: Some species are cultivated as ornamentals and for cut flowers. Etymology: Blandfordia was named in honour of George Spencer-Churchill, the fifth Duke of Marlborough and Marquis of Blandford (1766–1840), a collector of books and antiquities.

100. ASTELIACEAE Pineapple-grass family

Distribution: This family is restricted to eastern Australia and Tasmania. Phylogeny and evolution: Blandfordia was traditionally associated with Hemerocallis, Kniphofia, Phormium or even Dasypogonaceae. Molecular analyses placed the genus in an isolated position among Asteliaceae, Boryaceae and Hypoxidaceae. Morphological similarities between Boryaceae

Blandfordia nobilis, Australia (KD) [99]

and Blandfordiaceae are evident, which may be plesiomorphic for the larger clade.

Blandfordia cunninghamii by W. Fitch from Curtis’s Botanical Magazine vol. 94: plate 5734 (1868) [99]

These perennial short-stemmed tufted herbs usually grow terrestrially but are sometimes found as epiphytes. Plants are either male or female (except in Milligania), and they are characterised by the peculiar indumentum of scales. Leaves are spirally arranged, usually three-ranked, linear to ensiform and deeply keeled with a broad, closed sheathing base. Leaves are densely parallel veined and usually densely hairy (the hairs branched) at least when young. Inflorescences are terminal compound panicles composed of reduced racemes, each raceme with a spathe-like bract, the bracteoles usually on short pedicels. The unisexual, actinomorphic flowers are usually tri- sometimes pentamerous, the tepals usually six in two equal whorls, free or fused at the base to form a tube. In male flowers, six stamens are free or fused to the perianth tube, with dorsifixed to basifixed anthers that open by longitudinal slits. Female flowers have six staminodes surrounding the superior ovary that is composed of three to seven carpels forming one to three locules.

ASPARAGALES

MONOCOTS

Slit-like septal nectaries are present on the ovary around the short and thick style between the stigmatic surfaces. The fruit is a berry or a capsule. Distribution: This mostly Southern Hemisphere family has a patchy distribution, Asteliaceae can be found in southern South America, the Falkland Islands, the Mascarene Islands, New Guinea, southern Australia and New Zealand and from New Caledonia to Polynesia and Hawaii. Phylogeny and evolution: Fossils of Astelia are known from the Upper Eocene of New Zealand. The group has been dated to be c. 92 million years old. Collospermum is embedded in Astelia and should be merged with that genus. The position of Milligania is uncertain, but may together with Neoastelia form the

sister to Astelia. Long-distance dispersal must be common in this family, given the broad distribution and occurrence on recently formed oceanic islands. They share branched hairs with Hypoxidaceae and Lanariaceae and mucilage canals with Hypoxidaceae, the former perhaps a synapomorphy of the hypoxid clade minus Blandfordiaceae.

101. LANARIACEAE Lambtails family

Genera and species: This family has three genera and 37 species; Astelia (31), Milligania (5) and Neoastelia (1). Uses: Fruits of Astelia nervosa are edible, and fibre of A. grandis is used for handicrafts in New Zealand. Etymology: Astelia is composed of the Greek prefix α- (a-), not, and στήλη (steli), a column, in reference to the lack of a stem or trunk.

Astelia hemichrysa, infructescence, Réunion [100]

These are perennial herbs with short erect rhizomes that produce roots in the lower part and have spirally arranged leaves around the tip. Glabrous, linear leaves are sessile, have a closed sheath at the base and are parallel veined. Inflorescences are dense corymbose panicles on a long-bracted scape, and all parts of the inflorescence are densely woolly, with branched hairs. Bisexual, actinomorphic flowers are covered with dense woolly hairs outside and are pale mauve or purple inside. Six equal tepals are fused into a tube for the lower half, and six anthers are fused to this tube. The inner three stamens are shorter than the outer three, and anthers are dorsifixed and open by longitudinal slits. The inferior ovary is trilocular, has septal nectaries and is topped Lanaria lanata, South Africa (JA) [101]

Astelia neocaledonica, Royal Botanic Gardens, Melbourne, Australia [100]

Lanaria lanata, South Africa (JA) [101]

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ASPARAGALES with a narrow filiform style. The fruit is a capsule with a single seed.

MONOCOTS

102. HYPOXIDACEAE Stargrass family

Distribution: The family is restricted to the southern coast of the Cape Province of South Africa, from Somerset Bay to Albany, where it grows in fynbos vegetation. Phylogeny and evolution: Previously Lanaria was associated with Haemodoraceae, which it resembles in inflorescence morphology and hairy flowers. It had also been thought to have a relationship with Lophiola (Nartheciaceae), but this was refuted on the basis of differences in leaf morphology, anatomy and molecular phylogenetics. Lanaria is sister to Hypoxidaceae, with which it shares several characters but lacks mucilage canals, which are present in both Asteliaceae and Hypoxidaceae.

Etymology: Lanaria is derived from Latin, lana, wool, in reference to the woolly cottonlike appearance of the plant.

This family of perennial herbs forms a tuberous rhizome or corm that is covered with membranous, fibrous remains of former leaf sheaths. Leaves are all basal, alternately arranged in three ranks at the apex of the rhizome. They can be sessile or petiolate (Curculigo) and sheathing at the base. Blades are linear to lanceolate and hairy, the hairs branched, often along margins, or glabrous, usually V-shaped in cross-section or plicate and palm-like (in Curculigo). Inflorescences are spikes, corymbs or false umbels that are usually on hairy scapes arising from the leaf axils. The bisexual flowers emerge from axils of bracts that can be small or large and leafy.

Curculigo seychellensis, Mahé, Seychelles [102]

Hypoxis setosa, Royal Horticultural Society Garden, Wisley, UK [102]

Genera and species: The family consists of the single species Lanaria lanata.

Curculigo capitulata, Gunung Datuk, Malaysia [102]

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Smaller cleistogamous flowers are sometimes present. The six (rarely four) tepals form two equal whorls and are petal-like, often yellow, white or pink, sometimes red or maroon. They are free or basally fused into a tube and are more or less persistent in fruit. Usually six (sometimes three) stamens are inserted at the base of perianth segments and have short filaments and basifixed or dorsifixed anthers that open by longitudinal slits. The inferior ovary is trilocular, topped with a short style and three stigmas that can have free lobes or grooves along the style. The fruit usually is a capsule crowned by the persistent perianth that opens irregularly, often by circular slits, sometimes the fruit is a fleshy berry. Distribution: This pantropical family extends into temperate North America, southern Japan and Australia but is absent from Europe and the Middle East. It is especially diverse in temperate South Africa. Phylogeny and evolution: The crown group Hypoxidaceae have been dated to c. 78 million years ago. In older classifications, Hypoxidaceae were included in Amaryllidaceae. Hypoxis baurii, Royal Botanic Gardens, Kew, UK [102]

ASPARAGALES

MONOCOTS

Etymology: Hypoxis is an ancient Greek name for plants with somewhat acid leaves, from υπό (hypo), below and οξύς (oxys), sharp; later used for this genus by Linnaeus.

These giant, tufted, evergreen perennials make short underground stems from which the numerous long, spirally arranged, linear to lanceolate leaves grow. Their roots are thick and fleshy, and the leaf bases are filled with starch and form a bulb-like structure; leaf tips have a tubular brown tip that withers away to leave a tuft of thread-like fibres. Inf lorescences are large terminal

slightly or densely contracted thyrses that can grow up to five and a half metres tall and terminate a scape that at the base is covered with leaf-like bracts; towards the apex flowers and inflorescence branches are subtended by bright red bracts. The large bisexual flowers are on short pedicels, each bearing a prophyll. The six petallike tepals are united at the base in a tube that encloses the ovary, forming a nectar cup. The six stamens are fused with the perianth tube for about half their length; anthers are connected to the filament in a tube formed by the connective, the thecae opening by longitudinal slits. The inferior ovary is trilocular and has well-developed septal nectaries that open around the base of the style and fill the nectar cup. The simple slender style has a blunt triangular stigma. The fruit is a capsule that splits in three, exposing laterally winged seeds.

Doryanthes palmeri, Adelaide Botanic Garden, South Australia [103]

Doryanthes excelsa, Australian National Botanic Gardens, Canberra [103]

Doryanthes excelsa, Australian National Botanic Gardens, Canberra [103]

On the basis of morphology, a relationship to Orchidaceae has been suggested. On the basis of recent molecular studies, the family is redefined to include only four genera, with Curculigo including Hypoxidia and Molineria, Hypoxis including Rhodohypoxis, and Pauridia including Spiloxene and most Australian species of Hypoxis. Genera and species: Hypoxidaceae include four genera and c. 160 species: Curculigo (25), Empodium (7), Hypoxis (97) and Pauridia (c. 30). Uses: Curculigo capitulata and Hypoxis baurii (and cultivars) are grown as ornamentals.

Doryanthes palmeri, with Maarten Christenhusz for scale, Adelaide Botanic Garden, South Australia [103]

103. DORYANTHACEAE Gymea-lily family

Ixiolirion tataricum, Royal Botanic Gardens, Kew, UK [104]

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ASPARAGALES Distribution: This family is restricted to eastern Australia, where it is found between Jervis Bay and Burkedin River, in rocky gullies near the coast. Phylogeny and evolution: Doryanthes was thought to be closely related to Phormium (Asphodelaceae), but it is distant from that genus, instead forming a clade with Ixioliriaceae and Tecophilaeaceae and is now placed in its own family. Fossils 75–70 million years old assigned to Doryanthaceae or Iridaceae have been found in Late Cretaceous rocks of eastern Siberia. The split between Doryanthaceae and Iridaceae has been estimated by molecular clock to be at least c. 82 million year old, but another estimated age for this lineage is 107 million years. Genera and species: The single genus Doryanthes consists of two species: D. excelsa and D. palmeri. Uses: Both species are sometimes cultivated as ornamentals in Australia, and are an important food source for birds, mainly honeyeaters. Etymology: Doryanthes is composed of the Greek δόρυ (dory), a spear, and άνθος (anthos), a flower.

104. IXIOLIRIACEAE Tartar-lily family

MONOCOTS

bear a prophyll. The bisexual actinomorphic flowers have six petal-like tepals in two equal whorls, which are free nearly to the base. The outer tepals end in a prominent tip just below the apex. The six stamens are inserted at the base of the tepals and have basifixed anthers that open by longitudinal slits. The inferior ovary is trilocular and tipped with a slender style that is split in three at the dry stigmatic area. The fruit is a dry capsule that splits at the top revealing the numerous black seeds. Distribution: This family is found in semiarid lands of southwestern and inner Asia, from Egypt and the Levant to Central Asia. Phylogeny and evolution: In the past Ixiolirion was placed in Amaryllidaceae, but the position in that family has always been uncomfortable due to its corm, leafy scapes, blue flowers, lack of alkaloids and presence of simultaneous sporogenesis, all rare in Amaryllidaceae. Molecular analyses placed Ixiolirion with Doryanthes and Tecophilaeaceae in proximity to Iridaceae in Asparagales, supporting this placement as a separate family, although it could potentially be merged with Tecophilaeaceae, with which it shares many characters. Genera and species: The single genus Ixiolirion consists of four species. Uses: Ixiolirion tataricum is sometimes available in the ornamental bulb trade. Etymology: Ixiolirion is derived from the Greek ιξιά (ixia), bird lime, and λείριων (leirion), a lily.

105. TECOPHILAEACEAE Chilean-crocus family

These are perennial herbs with tunicate corms that renew from the base plate. Stems are erect, leafy and unbranched. Flat linear leaves alternate along the stem are sheathing at the base with a cylindrical apex. Inflorescences are thyrses or false umbels with few to many blue, violet or white flowers on pedicels that

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These perennial herbs grow from a globose or ellipsoid underground corm, from which leaves and stems emerge. Leaves are basal or (in Walleria) alternate along the stem, and are usually sheathing at the base, but not always. The simple leaves are sessile and linear-lanceolate or (in Cyanastrum and Kabuyea) lanceolate and ovate with a distinct petiole, and sometimes with a cordate base. Venation is parallel or parallel-arching in broader leaves with transverse secondary veins, the midvein often pronounced. Flowers are formed in panicles or solitarily in the leaf axils, often with bracts or breacteoles on the pedicel or peduncle. The bisexual flowers are actinomorphic or zygomorphic, and the six petal-like tepals are fused at the base and arranged in two whorls. The corona is sometimes adorned with minute appendages between the stamens. The six stamens arise at the mouth of the perianth tube, and some of these are sometimes staminodial. Stamens are regular in actinomorphic flowers, facing one side of the flower (often upwards) in zygomorphic flowers. The fertile anthers are basifixed or nearly so and open by pores. The ovary is semi-inferior or nearly superior and is composed of three carpels forming three locules. The fruit is a capsule that splits into three, exposing the black or brown seeds. Distribution: This disjunct family is found in California, Chile and Sub-Saharan Africa. Phylogeny and evolution: The family was previously associated with Haemodoraceae, but the presence of simultaneous microsporogenesis, phytomelan in the seed coat and the absence of cell-wall ferulates place this family in Asparagales. Molecular evidence shows that it is related to Ixioliriaceae, Doryanthaceae and Iridaceae. The crown group of Tecophilaeaceae are dated to be 77–87 million years old. Their biogeography is complex, and colonisation of the family in what are now Mediterranean ecosystems must have happened before this type of climate was present in these areas. The South American genera Conanthera and Zephyra appear to be sister to the rest of the family, although in some studies Tecophilaea takes this position.

ASPARAGALES

MONOCOTS

Conanthera bifolia, Royal Botanic Gardens, Kew, UK [105]

Odontostomum hartwegii, Royal Botanic Gardens, Kew, UK [105]

Tecophilaea cyanocrocus, Royal Botanic Gardens, Kew, UK

Genera and species: This small family of nine genera has 27 species: Conanthera (5), Cyanastrum (3), Cyanella (9), Eremiolirion (1), Kabuyea (1), Odontostomum (1), Tecophilaea (2), Walleria (3) and Zephyra (2).

This family includes mostly perennial herbs, rarely shrubs (Klattia, Nivenia, Witsenia) or annuals that are usually chlorophyllous and autotrophic apart from Geosiris, which is a mycoheterotrophic herb lacking chlorophyll. Plants are evergreen or seasonal, sometimes clump-forming, with rhizomes, corms or bulbs, rarely with a woody caudex or with an indistinct rootstock. Leaves are formed basally or along the stem, sheathing at the base with an open or closed sheath, often in one plane (distichous) with the bases usually overlapping (equitant), the blades flattened, often unifacial and oriented with their edge towards the stem, or terete, rarely with false petioles and broadened blades. Veins are parallel, with or without a distinct midvein, the surface flat, ribbed or plicate. Leaves are scale-like in Geosiris. Flowering stems are usually aerial (subterranean at anthesis in Romulea) and are simple or branched, round or flattened and angular or winged. Inflorescences are umbellate cymes (rhipidia), arranged in panicles, spikes or randomly clustered on terminal or lateral branches. Rhipidia comprise pedicellate or nearly sessile flowers subtended by a bract (or spathe) that encloses the flower until anthesis, sometimes reduced to a spike of sessile flowers subtended by opposite bracts or reduced to single flowers. The bisexual actinomorphic or zygomorphic flowers are

Uses: Apart from occasionally being grown as ornamental plants (especially Cyanastrum cordifolium and Tecophilaea cyanocrocus), this family has no economic importance. Tecophilaea cyanocrocus was previously thought to be extinct in the wild, but new populations have since been found. Collecting from the wild is still prohibited. Etymology: Tecophilaea was named by Italian botanist Luigi Aloysius Colla (1766–1848) for his daughter, botanical artist Tecophila Colla-Billotti.

106. IRIDACEAE Iris family

[105]

Cyanella lutea, Royal Botanic Gardens, Kew, UK [105]

composed of two whorls or three petal-like tepals, the inner whorl sometimes suppressed, when zygomorphic the flower often two-lipped, the upper tepal often enlarged and hooded. Tepals are free or fused into a tube, often bearing nectaries and nectar guides. Stamens are usually three (two in Diplarrena) and inserted at the base of the outer tepals or inside the tube. They can be symmetrically arranged in actinomorphic flowers or are unilateral and curved in zygomorphic ones. Filaments are free or completely fused, the anthers are basifixed to centrifixed with the two thecae usually opening by longitudinal slits. The inferior (superior only in Isophysis) ovary is trilocular and topped by a single filiform style that is branched or lobed in three at the tip. The fruit is a capsule that generally splits in three, rarely indehiscent, exposing the seeds that can be simple, usually dry and brown, but sometimes winged or covered with a red aril. Distribution: This is a cosmopolitan family that is patchy in tropical Asia and absent from true deserts. Phylogeny and evolution: Iridaceae diverged from the remaining Asparagales c. 80 million years ago in the Late Cretaceous. Isophysis (now restricted to Tasmania) may have diverged from the rest of the family c. 66

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ASPARAGALES

166

MONOCOTS

Diplarrena moraea, private garden, Kingston upon Thames, Surrey, UK [106]

Tigridia pavonina, private garden, Kingston upon Thames, Surrey, UK [106]

Sisyrinchium bellum, California, USA [106]

Gladiolus carmineus, Royal Botanic Gardens, Kew, UK [106]

Crocus sativus, Royal Botanic Gardens, Kew, UK [106]

Iris pseudacorus, Kew Green, Richmond, Surrey, UK [106]

Moraea sisyrinchium, Ronda, Spain [106]

Lapeirousia oreogena, Royal Botanic Gardens, Kew, UK [106]

Orthrosanthus laxus, Mt Benia, Western Australia [106]

Christenhusz, Fay & Chase

ASPARAGALES

MONOCOTS

million years ago. Most other early-branching lineages are also of Australasian origin, and it is therefore hypothesised that Iridaceae has an Australian-Antarctic origin from where it dispersed across the globe. Their current centres of diversity are in the Andes and Southern Africa, where they diversified from the Eocene onwards. Iridaceae are currently divided into seven subfamilies (Aristeoideae, Crocoideae, Geosiridoideae, Iridoideae, Isophysidoideae, Nivenioideae and Patersonioideae) based on morphological characters and the results of molecular analyses. Isophysidoideae comprise Isophysis tasmanica with a superior ovary; it is sister to the rest of Iridaceae. Patersonioideae are only found in Australia and often lack the inner whorls of tepals. Shrubby Nivenioideae are sister to the cormous Crocoideae, but sometimes include Patersonioideae as a section. Recent molecular studies have resulted in the reorganisation of a number of genera: e.g. Schizostylis is now placed in Hesperantha; Belamcanda, Juno, Pardanthus and Hermodactylus are merged with Iris; Acidanthera is part of Gladiolus, Anomatheca belongs to Freesia, and Homeria and Gynandriris have been merged with Moraea. Genera and species: Iridaceae include 66 genera and c. 2,244 species, in seven subfamilies. Isophysidoideae – Isophysis (1); Iridoideae – Alophia (5), Bobartia (15), Calydorea (22), Cipura (8), Cobana (1), Cypella (33), Dietes (6), Diplarrena (2), Eleutherine (4), Ennealophus (5), Ferraria (18), Gelasine (7), Herbertia (9), Hesperoxiphion (5), Iris (281), Lethia (1), Libertia (16), Mastigostyla (27), Moraea (206), Nemastylis (6), Neomarica (28), Olsynium (17), Orthrosanthus (9), Phalocallis (1), Pseudiris (1), Pseudotrimezia (18), Sisyrinchium (200), Solenomelus (2), Tapeinia (1), Tigridia (54) and Trimezia (28); Patersonioideae – Patersonia (25); Geosiridoideae – Geosiris (2); Aristeoideae – Aristea (56); Nivenioideae – Klattia (3), Nivenia (11) and Witsenia (1); Crocoideae – Babiana (92), Chasmanthe (3), Crocosmia (8), Crocus (104), Cyanixia (1), Devia (1), Dierama (43), Duthiastrum (1), Freesia (16), Geissorhiza

(97), Gladiolus (276), Hesperantha (84), Ixia (79), Lapeirousia (41), Melasphaerula (1), Micranthus (3), Pillansia (1), Radinosiphon (2), Romulea (111), Savannosiphon (1), Sparaxis (15), Syringodea (8), Thereianthus (11), Tritonia (28), Tritoniopsis (23), Watsonia (52), Xenoscapa (3) and Zygotritonia (4). Uses: Stigmas of Crocus sativus are the source of the expensive spice, saffron. They are grown on a commercial scale in Spain, Morocco, India and Turkey. Corms of Crocus, Lapeirousia and Moraea fugax are locally eaten. Starch of orris root, Iris pallida, was formerly used medicinally, for perfumes and to flavour gin. Many genera are grown as ornamentals, especially Babiana, Crocosmia, Crocus, Dierama, Freesia, Gladiolus, Hesperantha, Iris, Ixia, Libertia, Moraea, Neomarica, Sisyrinchium, Sparaxis, Tigridia, Tritonia and Watsonia. For the cut-f lower industry, Freesia, Gladiolus, Iris and Ixia are important. Many species escape from cultivation and cause considerable trouble as invasives, crowding out native plants. Etymology: Iris (Ίρις) is the ancient Greek goddess of the rainbow, in reference to the multitude of colours found among the nearly 300 species.

107. XERONEMATACEAE Poor-Knights-lily family

This family includes robust stemless, rhizomatous, tufted, evergreen perennial herbs. Rhizomes are short and upright, but hidden by the fibrous remains of old leaves. Iris-like, ensiform (unifacial) leaves are alternate, in a plane and overlapping (distichous), rigidly upright and sheathing at the base. Inflorescences are terminal, spikelike racemes that grow horizontally with the flowers pointing upwards. The peduncle has several sheathing bracts, and pedicels are subtended by floral bracts. Six red tepals are free, more or less equal in size, erect at first but reflexing and twisting downwards and persisting with the stamen filaments until the capsule ripens. Six stamens are longer than the tepals and have dorsifixed anthers opening by a longitudinal slit. The superior ovary is more or less stipitate (on a gynophore), constricted

Xeronema callistemon, Royal Botanic Gardens, Kew, UK [107]

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ASPARAGALES in the middle and trilocular. The style is also longer than the tepals and terete with a narrow stigma. The fruit is a triangular capsule that is broader below than above the equatorial constriction. It splits into three, but the valves usually only separate widely in the upper part. Distribution: This family is restricted to central New Caledonia and Poor Knights and Taranga Islands in northern New Zealand. Phylogeny and evolution: The genus has been variously placed in Asphodelaceae or Hemerocallidaceae (now a subfamily of Asphodelaceae), but molecular evidence has shown that it is the sister to the combined Amaryllidaceae, Asparagaceae and Asphodelaceae. The age of the stem group of Xeronemataceae is estimated to be c. 70 to 100 million years old. Genera and species: Xeronema includes two species: X. callistemon off New Zealand and X. moorei in New Caledonia. Etymology: Xeronema is derived from Greek ξηρός (xeros), dry, and νήμα (nema), a thread, in reference to the persistent filaments of the flowers.

108. ASPHODELACEAE Daylily family

A diverse family of clump-forming or rhizomatous perennial herbs, shrubs, vines or pachycaul branched or unbranched trees with anomalous secondary growth. Roots are usually succulent and sometimes thickened and tuberous, occasionally stilt-like. The leaves are held in a plane (distichous) or spirally arranged, herbaceous or succulent, or brittle, flat, V-shaped, swollen or terete,

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MONOCOTS

with parallel venation that may be obscure in succulent species; some have thickened resiniferous bases (Xanthorrhoea). Inflorescences are terminal variously branched panicles, racemes and cylindrical spikes or reduced to a single flower, rarely an umbel (Tricoryne), the peduncle is leafless or with (sometimes colourful) bracts. The bisexual flowers are actinomorphic or zygomorphic and composed of six free tepals that can be slightly fused into a tube at base, the whorls equal or the inner slightly larger than the outer whorl. The six stamens are free or some fused basally with the tepals. Anthers are basifixed or dorsifixed and open by pores or slits. The superior or semi-inferior ovary is uni- or trilocular and has a simple style at the apex, rarely gynobasic (Tricoryne), terminated by a small capitate stigma or a tuft of hairs. The fruit is a berry, nutlet or capsule that splits into three, exposing black seeds. Distribution: This widespread family is found in western South America, southern and central Europe, North Africa, Turkey east to temperate and tropical East Asia and south throughout the Middle East, Sub-Saharan Africa, Madagascar, Malesia, New Guinea, Australia, New Zealand and throughout the Pacific. Phylogeny and evolution: The crown group of Asphodelaceae is dated to c. 90 million years. Fossil pollen of Phormium is known from the Upper Miocene of New Zealand. Excremis and Pasithea represent independent migrations of the phormioid clade to South America. The three subfamilies have often been treated as independent families, but they share a number of characters. The family circumscription has been controversial and a three-family concept is preferred by some. However, if this is done, the daylily family is to be expanded with a number of Southern Hemisphere genera (e.g. former families Johnsoniaceae and Phormiaceae) making it morphologically just as diverse as a broader Asphodelaceae. Recognising Aloaceae, as has also often been advocated, would make Asphodelaceae non-monophyletic. The

large genus Aloë is not monophyletic in its traditional sense, Astroloba, Chortolirion, Gasteria and Haworthia being embedded, which resulted in the merger of Chortolirion with Aloë and the segregation of Aloidendron, Aloiampelos, Aristaloë, Gonialoë, Haworthiopsis, Kumara and Tulista. Many intergeneric hybrids are known, which makes one wonder if it is not better to treat Aloë in a broader sense and include Gasteria and Haworthia. Geitonoplesium has resupinate leaves, which resulted in an association of that genus with Alstroemeriaceae, but it is placed here on the basis of other characters. Many species produce copious nectar and are pollinated by birds, driving differences in flower structure among closely related taxa. Many know this family as Xanthorrhoeaceae as that name was used in APG III because this name had nomenclatural priority, but the name Asphodelaceae has been conserved by committee and is now the one to be applied to this family. Genera and species: A family of 39 genera with c. 1,200 species, subdivided into three subfamilies: Asphodeloideae – Aloë (536), Aloiampelos (10), Aloidendron (6), Aristaloë (1), Asphodeline (17), Asphodelus (17), Astroloba (7), Bulbine (77), Bulbinella (25), Eremurus (59), Gasteria (20), Gonialoë (3), Haworthia (149), Haworthiopsis (3), Kniphofia (71), Kumara (2), Trachyandra (57) and Tulista (2); Xanthorrhoeoideae – Xanthorrhoea (28); Hemerocallidoideae – Agrostocrinum (2), Arnocrinum (3), Caesia (13), Chamaescilla (3), Corynotheca (6), Dianella (41), Excremis (1), Geitonoplesium (1), Hemerocallis (19), Hensmania (3), Herpolirion (1), Hodgsoniola (1), Johnsonia (5), Pasithea (1), Phormium (2), Simethis (1), Stawellia (2), Stypandra (2), Thelionema (3) and Tricoryne (8). Uses: Dried leaf sap of aloes is used in a wide range of cosmetics, medicines and drinks. Aloë vera is cultivated for this purpose on a commercial scale, especially in the Caribbean, whereas A. ferox is mostly harvested from the wild in Africa. Young shoots and flower buds of daylilies, Hemerocallis, are often eaten

ASPARAGALES

MONOCOTS

Phormium cookianum, Royal Botanic Gardens, Kew, UK [108]

Geitonoplesium cymosum, National Botanic Gardens, Glasnevin, Ireland [108]

Hemerocallis fulva, Helsinki Botanical Garden, Finland [108]

Aloë macra, Réunion [108]

Kniphofia hirsuta, private garden, Kingston upon Thames, Surrey, UK [108]

Xanthorrhoea preissii, Mt Benia, Western Australia [108]

Asphodeline lutea, Greece [108]

Chamaescilla versicolor, Mt Benia, Western Australia [108]

Dianella revoluta, Royal Botanic Gardens, Kew, UK [108]

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ASPARAGALES

MONOCOTS

in Asia, although not all species are edible. Many cultivars and hybrids exist, and these are popular garden plants. New Zealand flax, Phormium tenax, was used to make clothes and ropes and planted commercially for this purpose, becoming problematic on oceanic islands like St Helena, where it was grown to supply twine to the UK post office. It is now mostly grown as an ornamental. Dianella leaves were used for similar purposes. Fruits of Dianella ensifolia are sometimes eaten and have been used as a cloth dye. Grass trees (Xanthorrhoea) were formerly harvested for their resin, which was made into varnish, lacquer and wood stains. Many are popular ornamental plants especially species of Aloë, Asphodeline, Bulbine, Eremurus, Gasteria, Haworthia and Kniphofia. Etymology: Asphodelus is a Latinised form of the Greek plant name ασφόδελος (asfodelos), a plant associated with the afterlife, and it was often associated in mythology with Persephone, who was depicted wearing a garland of asphodels as a reference to the underworld. Tulbaghia ludwigiana, Royal Botanic Gardens, Kew, UK [109]

Crinum purpurascens, Royal Botanic Gardens, Kew, UK [109]

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109. AMARYLLIDACEAE Onion family

This family includes perennial herbs with f leshy rhizomes or bulbs, enveloped by sheathing dry leaves or leaf bases. They usually grow terrestrially, but aquatics and rarely epiphytes also occur. The alternate, distichously arranged leaves are usually linear and filiform, sometimes lanceolate to ovate, flat, angular, round or swollen and hollow (fistular), sessile or petiolate, with sheaths closed at the base and parallel venation. Inflorescences are umbels (reduced helicoid cymes), sometimes reduced to a single flower, or forming a spike (Allium spicatum),

subtended by one or two (rarely more) spathelike bracts that cover the inflorescence in bud, sometimes with inner bracteoles. Bisexual flowers are actinomorphic or zygomorphic. There are usually six, sometimes three or five tepals, fused at least at the base, sometimes forming a tube. Scales or appendices between tepals and stamens sometimes form a corona (some Allium, Tulbaghia and some Leucocoryne). Outgrowth of the perigone may sometimes also form a corona (Narcissus). Functional stamens are usually six, sometimes two or three and the missing ones then often staminodial, or 18 or more (Gethyllis), the stamens adnate to the style in Strumaria. Filaments are inserted on the tepals, free from each other or fused, often with appendages or forming a corona (e.g. Hymenocallis, Hieronymiella). Anthers are dorsifixed or basifixed and open by longitudinal slits or by pores in Galanthus and Leucojum. The trilocular ovary is superior (Agapanthoideae and most Allioideae), semi-inferior (some Allium) or inferior (Amaryllidoideae) with three nectaries and a style at the the base

Amaryllis belladonna, Beth Chatto Agapanthus inapertus, private garden, Kingston upon Thames, Surrey, UK [109] Gardens, England, UK [109]

Speea humilis, Royal Botanic Gardens, Kew, UK [109]

Leucocoryne pauciflora, Royal Botanic Gardens, Kew, UK [109]

Proiphys cunninghamii, Australian National Botanic Gardens, Canberra [109]

Allium callimischon subsp. haematostictum, Royal Botanic Gardens, Kew, UK [109]

ASPARAGALES

MONOCOTS

(Allium) or at the apex of the ovary. The fruit is a capsule that splits in three, rarely indehiscent or a berry. Distribution: This is a cosmopolitan family. Agapanthoideae are restricted to South Africa. Allioideae are mostly found in the Northern Hemisphere, southern Africa and South America. Amaryllidoideae have their greatest phylogenetic diversity in southern Africa, from where lineages moved northwards through Africa into Eurasia and thence to the Americas and Australasia. Phylogeny and evolution: Amaryllidaceae are thought to have evolved c. 87 million years ago. South African Agapanthus is sister to the rest of the family and evolved c. 50 million years ago. Even though this genus has been placed in its own family, it shares the umbellate inflorescence that is covered by bracts in bud, representative for all members of this family, including ones with singleflowered inflorescences. Allioideae diversified in the Northern Hemisphere, from where southern Africa (Tulbaghieae) and South America (Gilliesieae) lineages originated. The bilaterally symmetrical flowers of Gilliesia, Miersia and Solaria are thought to mimic female insects. In Amaryllidoideae, the American species are sister to the Eurasian members, which suggests a so-called Boreotropical hypothesis, one in which the family entered the Americas through the boreal zones when these regions were much warmer than they are today. Similar as they may be in appearance, the South African Amaryllis is not directly related to South American Hippeastrum (which bears amaryllis as its common name). Generic delimitation needs some work especially among American Amaryllidoideae; genera in Hippeastreae are poorly circumscribed and require reclassification. To maintain Galanthus as separate from Leucojum, Acis was reinstated. Nectaroscordum belongs to Allium. This family was formerly known as Alliaceae (e.g. APG II), which had nomenclatural priority, but the name was rejected

by committee in favour of Amaryllidaceae, which is now the name to be used. Genera and species: This family includes 77 genera and c. 2,140 species, divided in three subfamilies: Amaryllidoideae (62 genera, c. 1,000 species) – Acis (9), Amaryllis (2), Ammocharis (7), Apodolirion (6), Boophone (2), Brunsvigia (9), Caliphruria (4), Calostemma (3), Cearanthes (1), Chlidanthus (4), Clinanthus (22), Clivia (6), Crinum (105), Crossyne (2), Cryptostephanus (3), Cyrtanthus (56), Eithea (1), Eucharis (17), Eucrosia (8), Eustephia (6), Galanthus (20), Gethyllis (30), Griffinia (21), Habranthus (83), Haemanthus (22), Hannonia (1), Hessea (13), Hieronymiella (8), Hippeastrum (91), Hymenocallis (65), Ismene (10), Lapiedra (1), Leptochiton (2), Leucojum (2), Lycoris (22), Mathieua (1), Namaquanula (2), Narcissus (c. 30), Nerine (25), Pamianthe (3), Pancratium (21), Paramongaia (2), Phaedranassa (9), Phycella (5), Placea (6), Plagiolirion (1), Proiphys (4), Pyrolirion (8), Rauhia (4), Rhodophiala (27), Scadoxus (9), Sprekelia (2), Stenomesson (16), Sternbergia (8), Strumaria (27), Tocantinia (1), Traubia (1), Ungernia (10), Urceolina (7), Vagaria (2), Worsleya (1) and Zephyranthes (87); Allioideae (14 genera, 1,134 species) – Allium (920), Gilliesia (7), Ipheion (3), Latace (2), Leucocoryne (48), Miersia (1), Nothoscordum (89), Prototulbaghia (1), Schickendantziella (1), Solaria (6), Speea (2), Trichlora (4), Tristagma (17) and Tulbaghia (27); Agapanthoideae – Agapanthus (7). Uses: Members of Allium has been used as vegetables or condiments since the Bronze Age. The onion, Allium cepa, is the most widely cultivated species. It is a cultigen, not known in the wild, but probably has its origin in Central Asia. Onions were such an important food that bulbs have been used as currency. A common variety of the onion is the shallot (A. cepa var. aggregatum, previously known as A. ascalonicum), which has a milder flavour and is often pickled. Leek (A. ampeloprasum) is another ancient vegetable and, with the daffodil (Narcissus pseudonarcissus), the symbol of Wales. Elephant garlic is probably close to the

wild ancestor of the leek. Another important crop is garlic (A. sativum), also with a 7,000year history of cultivation; it is important for food and human health. The first recorded cultivated onions, garlic and leeks were grown in Ancient Egypt. Other minor crops are chives (A. schoenoprasum), Welsh onion (A. fistulosum), garlic chives (A. tuberosum), Persian shallot (A. stipitatum) and scallion (A. chinense). Society garlic (Tulbaghia violacea) is also eaten and has the benefit of not making your breath smell as does garlic. Onion-scented Nothoscordum species are also sometimes used to flavour food. Species of Amaryllidoideae are poisonous but have been used in shamanic tradition to induce hallucinations and visions. Many genera are important in the bulb trade: Amaryllis, Allium, Crinum, Cyrtanthus, Galanthus, Hippeastrum, Hymenocallis, Ipheion, Ismene, Leucocoryne, Leucojum, Lycoris, Narcissus, Nerine, Sprekelia, Sternbergia and Zephyranthes. Agapanthus and Tulbaghia are popular garden perennials, and Clivia, Haemanthus, Eucharis and Scadoxus are frequent houseplants. Amaryllis (Hippeastrum) and narcissi (Narcissus) are important in the cut-flower industry. Some species of Nothoscordum are noxious weeds. Etymology: Amaryllis is the name of a shepherdess in Virgil’s Eclogues. It was probably derived from the Greek αμαρυσσό (amarysso), to sparkle.

110. ASPARAGACEAE Hyacinth family

This varied family includes perennial, annual, polycarpic or monocarpic, terrestrial and rarely epiphytic herbs, shrubs, trees and vines. Stems can form rhizomes, erect woody trunks

Plants of the World

171

ASPARAGALES

Yucca brevifolia, Joshua Tree National Park, California, USA [110]

MONOCOTS

Aphyllanthes monspeliensis, Spain [110]

Agave parryi, New Mexico (DZ) [110]

Asparagus plocamoides, Helsinki Botanical Garden, Finland [110]

Lomandra insularis, New Caledonia [110]

Dracaena reflexa var. angustifolia, Seychelles [110]

Thysanotus patersonii, Flinders Ranges, South Australia [110]

Theropogon pallidus, Royal Polygonatum odoratum, Finland [110] Botanic Gardens, Kew, UK [110]

Milla biflora, Royal Botanic Gardens, Kew, UK [110]

Bessera elegans, private garden Kingston upon Thames, Surrey, UK [110]

172

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Ledebouria cooperi, private garden Kingston upon Thames, Surrey, UK [110]

Ornithogalum nutans, Royal Eucomis comosa, private garden Botanic Gardens, Kew, UK [110] Kingston upon Thames, Surrey, UK [110]

ASPARAGALES

MONOCOTS

(with secondary growth), tunicate bulbs, corms and tubers, or stems can be scandent. Roots are often fleshy but can be fibrous in some species. Alternate, sometimes opposite or verticillate, leaves are clumped, in a rosette or spread along the stem and are usually spirally arranged, sometimes in a plane (distichous), or singular, sometimes lacking in the adult stage (Bowiea, Drimia), sessile or petiolate, herbacous or succulent or rigid and fibrous, sometimes the leaves reduced to scales (Aphyllanthes, Asparagus). Blades are linear, lanceolate, elliptic or ovate and the venation is parallel, sometimes with parallel or reticulate cross-veins, with the bases often clasping the stem, sheathing or not. Bases sometimes form a pseudostem, and leaf tips are occasionally cirrhose. In Asparagoideae and Nolinoideae, stems are sometimes broadened and resemble leaves (phylloclades; e.g. in Asparagus, Danaë, Ruscus and Semele). Flowering stems are bracteate, often scapose, sometimes underground or absent, the inflorescences terminal or axillary panicles, spikes and racemes, sometimes reduced to a single flower, sometimes umbel-like (Brodiaeoideae, Leucocrinum, some Dracaena), fascicled or capitate (Aphyllanthes), the bracts usually many and not enclosing buds. The bisexual, sometimes unisexual, flowers are actinomorphic or zygomorphic, usually subtended by one to several bracts. The six (sometimes four or five) tepals are petallike and free or united into a long or short tube or cup. The six (sometimes three, four, eight, ten or 12) stamens have free filaments that are sometimes fused with the perianth at the base. Stamens are rudimentary in female flowers of dioecious species, and staminodial when a whorl of stamens is missing. Anthers are dorsifixed or basifixed and open by longitudinal slits. The superior to inferior ovary is composed of three fused carpels, unior trilocular, rarely stalked (with a gynophore in Chlorophytum and some Brodiaeoideae) and topped by a single style with a capitate, trilobed or three-branched stigma, or the three styles free (Yucca). The fruit is usually a loculicidal, sometimes septicidal, capsule or

a berry, sometimes a three-winged samara or a three-ribbed nut. Distribution: Asparagaceae have a nearly cosmopolitan distribution but are absent from the coldest regions in the Arctic and Antarctic and have not been found in eastern Amazonia or the Sahara and Gobi Deserts. Aphyllanthoideae are restricted to the western Mediterranean. Agavoideae occur in the Americas from southern Canada to Patagonia, throughout central and southern Europe, Sub-Saharan Africa, South and East Asia and Malesia. Brodiaeoideae are restricted to western North America, Mexico and Guatemala. Scilloideae occur in western South America, Europe, North and Sub-Saharan Africa, Madagascar, western, central and southern Asia, China, Korea and Japan. Lomandroideae are restricted to Southeast Asia and Australasia. Asparagoideae are widespread across the Old World and have an isolated population in Mexico (Hemiphylacus). Nolinoideae are spread across North America, Mesoamerica, Europe, North, West and Central Asia, Sub-Saharan Africa and northern Australia. Phylogeny and evolution: Circumscription of this group of plants has varied greatly in the past. Most subfamilies mentioned below were known at some point as separate families: Agavaceae, Anemarrhenaceae, Anthericaceae, Behniaceae, Herreriaceae and Hostaceae (now all Agavoideae), Convallariaceae, Dracaenaceae, Eriospermaceae, Nolinaceae and Ruscaceae (now all Nolinoideae), Hyacinthaceae (now Scilloideae) and Themidaceae (now Brodiaeoideae). The seven subfamilies together form a well-supported clade. Asparagoideae and Lomandroideae are sister to the rest. The placement of Aphyllanthes has been problematic. Although here placed in Nolinoideae, Eriospermum has also been found as sister to Asparagoideae. Classification of Nolinoideae needs further study. Brodiaeoideae are similar to Allioideae of Amaryllidaceae (where they were

previously placed), but differ from those in having corms and three or more bracts that do not completely enclose the developing umbellate inflorescence. Fossils of Protoyucca are known from the Eocene of Nevada and are remarkably similar to modern Yucca. Nolinoideae-like fossils (Palaeocene Maianthemophyllum and Eocene Soleredera) are known from Canada. Fossil Paracordyline is known from Oligocene and Eocene deposits in Australia and Kerguelen. Pollen of Cordyline is known from the Miocene onwards. Divergence within Asparagaceae (crown group) is dated to have begun c. 89 million years ago. New World Oziroë diverged from the remaining Scilloideae (Old World) c. 28 million years ago. The tree-like habit frequent in Asparagaceae has evolved independently several times in Agavoideae (Yucca), Lomandroideae (Cordyline) and Nolinoideae (Beaucarnea, Dasylirion, Dracaena, Nolina). Some genera will need to be recircumscribed, pending further study. Beaucarnea is embedded in Nolina, and Sansevieria and Dracaena have also been united, but these changes are found to be somewhat controversial, especially in horticultural circles, and have not yet been adopted by the World Checklist of Selected Plant Families, which is why those are maintained here. Polianthes and Manfreda are merged with Agave, Smilacina has been merged with Maianthemum and Urginea is now part of Drimia. The genera Scilla and Muscari were also found to be polyphyletic, resulting in the acceptance of Alrawia, Fessia, Hyacinthella, Hyacinthoides, Leopoldia, Mer willa, Prospero, Pseudomuscari, Pseudoprospero, Resnova, Schizocarphus and Zagrosia in addition to Scilla, Muscari, Bellevalia, Puschkinia and Hyacinthus. Chionodoxa is part of Scilla sensu stricto. Nevertheless, generic delimitation in Scilloideae is still not yet well studied and is likely to undergo further change. Genera and species: A large family with 118 genera and c. 3,220 species divided in

Plants of the World

173

ASPARAGALES seven subfamilies: Aphyllanthoideae – only Aphyllanthes (1); Agavoideae – Agave (252), Anemarrhena (1), Anthericum (7), Behnia (1), Beschorneria (8), Camassia (6), Chlorogalum (5), Chlorophytum (193), Clara (3), Diamena (1), Diora (1), Diuranthera (4), Echeandia (80), Eremocrinum (1), Furcraea (23), Hagenbachia (6), Hastingsia (4), Herreria (8), Herreriopsis (1), Hesperaloë (8), Hesperocallis (1), Hesperoyucca (2), Hosta (23), Leucocrinum (1), Paradisea (2), Schoenolirion (3), Trihesperus (2) and Yucca (49); Brodiaeoideae – Androstephium (2), Bessera (3), Bloomeria (3), Brodiaea (17), Dandya (4), Dichelostemma (5), Jaimehintonia (1), Milla (10), Muilla (3), Petronymphe (1), Triteleia (15) and Triteleiopsis (1); Scilloideae – Albuca (160), Alrawia (2), Barnardia (2), Bellevalia (65), Bowiea (1), Brimeura (3), Daubenya (8), Dipcadi (41), Drimia (106), Drimiopsis (14), Eucomis (11), Fessia (11), Fusifilum (15), Hyacinthella (17), Hyacinthoides (11), Hyacinthus (3), Lachenalia (132), Ledebouria (55), Leopoldia (12), Massonia (14), Merwilla (3), Muscari (44), Namophila (1), Ornithogalum (185), Oziroë (5), Prospero (15), Pseudogaltonia (1), Pseudomuscari (7), Pseudoprospero (1), Puschkinia (2), Resnova (5), Schizocarphus (1), Scilla (81), Spetaea (1), Veltheimia (2) and Zagrosia (1); Asparagoideae – Asparagus (212) and Hemiphylacus (5); Lomandroideae – Acanthocarpus (7), Arthropodium (9), Chamaexeros (4), Cordyline (24), Dichopogon (5), Eustrephus (1), Laxmannia (14), Lomandra (51), Murchisonia (2), Romnalda (3), Sowerbaea (5), Thysanotus (50), Trichopetalum (2) and Xerolirion (1); Nolinoideae – Aspidistra (101), Beaucarnea (11), Comospermum (1), Convallaria (3), Danaë (1), Dasylirion (22), Disporopsis (7), Dracaena (112), Eriospermum (112), Heteropolygonatum (6), Liriope (6), Maianthemum (38), Nolina (28), Ophiopogon (67), Peliosanthes (23), Polygonatum (65), Reineckea (1), Rohdea (17), Ruscus (6), Sansevieria (69), Semele (3), Speirantha (1), Theropogon (1) and Tupistra (21).

174

Christenhusz, Fay & Chase

MONOCOTS

Uses: Garden asparagus, sometimes colloquially called sparrowgrass, (Asparagus officinalis) has been eaten since classical times and was depicted on an Egyptian frieze from 3000 BC. Other Asparagus species are harvested from the wild and eaten in spring in the Mediterranean and Asia. Prussian or Bath asparagus, young Ornithogalum pyrenaicum shoots, are also consumed sometimes. Muscari comosum bulbs are pickled in Greece. Quamash, Camassia was eaten by native North American tribes. Agave species have been used extensively by North American native peoples. Species like henequen (A. fourcroydes) and sisal (A. sisalana) are cultivated for their fibre, sometimes on a large scale. Agave tequilana is used to make alcoholic beverages like tequila. Mescal is a similar beverage made from A. americana, and sotol is made of roasted tubers of Dasylirion. The syrup made from A. tequilana and A. salmiana is marketed as agave nectar, which is a sweetener high in fructose. Furcraea foetida is also cultivated for its fibre that is similar to sisal. Plants are prolific, and some produce young plants in the inflorescence, making them problematic invasives. In the Pacific and Australasia, fibre of Cordyline and Lomandra is used extensively. Cordyline berries are used in Polynesian tattoo ink. Roots, shoots, leaves and inflorescences of species of Leucocrinum and Chlorophytum are eaten by indigenous peoples in North America and Africa, respectively. Flowers and buds of Convallaria keiskei and Tupistra nutans, young shoots and fruits of various Maianthemum species and rhizomes of some Polygonatum, Ophiopogon and Liriope species are eaten in Asia, and Cordyline was eaten in New Zealand and Indonesia. Leaves of Hosta are cooked and eaten in Korea and Japan. Bunches of Ruscus aculeatus (butcher’s broom) were cut to make brushes for cleaning the floor of butcher’s shops. Many species are popular garden or namentals and houseplants: e.g. Anemarrhena (zhi mu), Anthericum (St Bernard’s lily), Aspidistra (cast-iron plant),

Beschorneria, Beaucarnea (ponytail palm), Chlorophytum (spider plant), Convallaria (lily of the valley), Cordyline (cabbage tree), Dasylirion (desert spoon), Dracaena (dragon tree), Hosta (plantain lily), Ledebouria (silver squill), Liriope (monkey grass), Lomandra (mat-rush), Maianthemum (may lily), Ophiopogon (lilyturf), Polygonatum (Solomon’s seal), Sansevieria (mother-inlaw’s tongue) and Yucca (yuccapalm), and a good number are grown on a large scale for the f lower bulb industry: Camassia (quamash), Bellevalia, Bessera (coral drops), Brimeura (amethyst hyacinth), Brodiaea (cluster lily), Dichelostemma (blue dicks), Eucomis (pineapple lily), Hyacinthoides (bluebells), Hyacinthus (hyacinth), Lachenalia (Cape cowslip), Muscari (grape hyacinth), Ornithogalum (star-of-Bethlehem), Puschkinia (striped squill), Scilla (squill), Triteleia (triplet lily) and Veltheimia (elephant’s eye). Some species are used frequently in the cut-flower industry, e.g. Agave polianthes (tuberose), Convallaria majalis (lily-of-the-valley), Hyacinthus orientalis (hyacinth), Ornithogalum arabicum, O. dubium, O. saundersiae and O. thyrsoides, often because of their strong fragrances that are also used in the perfume industry. Stems of Danaë racemosa and the asparagus fern (Asparagus setaceus) are much used by florists. Dracaena reflexa var. angustifolia, usually sold as ‘D. marginata’, takes second place (after Ficus benjamini, Moraceae) as the most common indoor foliage plant in Europe and North America. Longest name: Ornithogalum adseptentrionesvergentulum, a species from the South African karroo, has the longest plant species epithet with 38 letters, yet it is a tiny plant species, flowering at just under 3 cm tall. A label with its name has to be taller than the plant. Etymology: Asparagus is the classical Latin name for the spring vegetable, which is derived from Ancient Persian asparag, a shoot or sprout.

ARECALES

COMMELINIDS The remainder of monocots comprise what are known as the “commelinids”, a clade of monocots with cell-wall ferulates (that fluoresce), Strelitzia-type epicuticular wax and an endosperm with abundant starch, although Arecaceae lack the last (most likely a secondary loss). Some authors have recognised this clade formally, often as the superorder Commelinanae, but they are not formally named in APG IV (2016).

ARECALES This order is composed of two families, Arecaceae and Dasypogonaceae, the latter unplaced until recently. The two families form trees in a similar manner and form with reasonable support a clade. It is noteworthy that the area where Dasypogonaceae are found in Western Australia lacks native palms, even though the climate is perfectly suitable for Arecaceae.

111. DASYPOGONACEAE Saviour-grass family

This is a family of shrubby and arborescent perennials. Stems are woody, creeping or erect, and when erect are covered in roots growing down from the crown to the base (Kingia). Leaves are spirally arranged, usually furrowed on the upper side, and can be short and broad or elongate and grasslike. Flowers can be solitary or in dense globular heads. The bisexual flowers have six Dasypogon obliquifolius, Mt Benia, Western Australia [111]

persistent petals in two similar whorls, which are variably fused or free. The six stamens are usually attached to the petal base, and anthers are basifixed or dorsifixed with two thecae that open by two slits or pores. The superior ovary is tri- or unilocular, the single style with a capitate or trilobed stigma. Fruits are indehiscent, rarely explosive capsules, enclosed by persistent petals. Distribution: Southwestern Australia houses nearly all the diversity of this family. A single species of Calectasia occurs disjunctly in western Victoria and adjacent South Australia. Phylogeny and evolution: The family is estimated to have originated c. 100 million years ago, but they have no fossil record. They are distantly related to Arecales. Members of the family were previously associated Calectasia grandiflora, Perth, Western Australia [111]

with other tree-forming monocots, but this was based on superficial resemblance: Xanthorrhoea for example, has (anomalous) secondary growth, whereas all growth in Dasypogonaceae, as in palms, is primary, even when woody as in Kingia. Genera and species: This small family includes four genera and c. 17 species: Baxteria (1), Calectasia (11), Dasypogon (3) and Kingia (2). Uses: Although species have ornamental potential, they grow slowly and the seeds are difficult to germinate. Leaf bases and other parts of Baxteria smell of rotten flesh, and this may be connected with attraction of pollinators. Etymology: Dasypogon is composed of Greek δασυς (dasys), hairy, and πωγων (pogon), beard.

Kingia australis, Mt Benia, Western Australia [111]

Plants of the World

175

ARECALES

Lodoicea maldivica, female tree, Praslin, Seychelles [112]

Lodoicea maldivica, male inflorescence, Praslin, Seychelles [112]

Nypa fruticans, Singapore (KH) [112]

Nypa fruticans, inflorescence, Singapore (KH) [112]

Chamaerops humilis, Royal Botanic Gardens, Kew, UK [112]

176

MONOCOTS

Christenhusz, Fay & Chase

Cyrtostachys renda, Singapore Botanical Garden [112]

Washingtonia filifera, Andreas Canyon, Palm Springs, California, USA [112]

Bismarckia nobilis, cultivated in Guayaquil, Ecuador [112]

Calamus longipinna, Royal Botanic Gardens, Kew, UK [112]

Ceroxylon quindiuense, San Francisco Botanical Garden, USA [112]

ARECALES

MONOCOTS

112. ARECACEAE Palm family

Palms include trees and shrubs with single or clustered, slender to thick, climbing, creeping and erect subterranean to tall, usually unbranched (rarely forking), woody stems that are armed with spines or not and often have clear leaf scars on the bark. They do not produce secondary growth, and although woody their trunks are formed solely through primary growth. Trunks are sometimes supported by stilt roots, and sometimes roots are modified into spines. Leaves are usually arranged in a spiral at the tip of the stem, sometimes in one plane (distichous) or three-ranked (tristichous). Petioles are sheathing, with the sheath initially tubular, often splitting later, sometimes with a ligule-like appendage. Above the sheath, petioles are armed with spines or not, often with a hastula (a small outgrowth between the petiole and the base of the leaf blade). Blades are entire and pinnately or palmately veined and costapalmately or pinnately divided, often formed by elongation of the rachis ripping the blades into leaflets, sometimes bifid (split in twos). Leaves are plicately folded in bud, the segments V- or Λ-shaped in cross-section. In some species, apices of the leaves are modified into a climbing whip, with the final leaflets replaced by spines. Inflorescences are solitary or multiple, compound or simple, axillary spikes, surrounded by one or more large bracts (prophylls), and peduncular bracts are absent to many. Flowers are unisexual or bisexual, similar or dimorphic, solitary or in cincinni. Sepals are usually three (sometimes two or more than three), free or variously fused. Petals are usually as many as sepals, free or fused. The usually six stamens (sometimes three or more numerous, up to c. 1,000) have erect filaments that are free or variously fused

with each other or with petals. Staminodes are strongly reduced and tooth-like or well developed, rarely absent. Ovaries are superior and have (1–)3(–4) fused carpels, with or without styles, the stigmas erect or curved backwards. Pistillodes can be present or absent in male flowers and are variable in shape and size when present. Fruits are usually singleseeded (sometimes with two to ten seeds), ranging from small to large (then often called ‘coconuts’), the skin smooth, hairy, prickly, warty or covered in overlapping scales, the mesocarp dry, fibrous or fleshy and the endocarp thin or woody, with one to three or without pores. Even though it was often previously assumed that all palms are windpollinated, it has been shown that numerous species are visited frequently and pollinated by insects, especially beetles, but also by bees, ants and flies. Distribution: Palms are pantropical and most diverse in montane rainforests in the tropics, but also in desert areas. They also extend into subtropical and temperate regions of North and South America, southern Europe, temperate Asia and New Zealand. Phylogeny and evolution: Palms were previously often associated with other ‘palmlike’ monocots such as Cyclanthaceae and Pandanaceae, but DNA analyses have shown them to be members of the commelinid clade, closest to Dasypogonaceae in the Arecales. They are divided into five subfamilies. Nypoideae have the oldest fossil record. Coryphoideae are considered the least specialised of palms and consist mostly of palmate and costapalmate genera. Calamoideae are distinguished by their scaled fruits that resemble pangolin skin. Some genera have been reorganised recently, notably Calamus includes the genera Daemonorops, Ceratolobus and Pogonotium. Genera and species: This is a family of 195 genera in five subfamilies and c. 2,400 species: Calamoideae – Calamus (c. 500), Eleiodoxa (1), Eremospatha (11), Eugeissona (6), Korthalsia (27), Laccosperma (6), Lepidocaryum (1), Mauritia (2), Mauritiella (4),

Metroxylon (7), Myrialepis (1), Oncocalamus (5), Pigafetta (2), Plectocomia (15), Plectocomiopsis (6), Raphia (20), Retispatha (1) and Salacca (22); Nypoideae – Nypa (1); Coryphoideae – Acoelorrhaphe (1), Arenga (24), Bismarckia (1), Borassodendron (2), Borassus (5), Brahea (11), Caryota (14), Chamaerops (1), Chelyocarpus (4), Chuniophoenix (3), Coccothrinax (51), Colpothrinax (3), Copernicia (22), Corypha (5), Cryosophila (10), Guihaia (2), Hemithrinax (3), Hyphaene (8), Itaya (1), Johannesteijsmannia (4), Kerriodoxa (1), Lanonia (8), Latania (3), Leucothrinax (1), Licuala (162), Livistona (28), Lodoicea (1), Maxburretia (3), Medemia (1), Nannorrhops (1), Phoenix (14), Pholidocarpus (6), Pritchardia (28), Rhapidophyllum (1), Rhapis (11), Sabal (14), Sabinaria (1), Saribus (9), Satranala (1), Schippia (1), Serenoa (1), Tahina (1), Thrinax (3), Trachycarpus (9), Trithrinax (4), Wallichia (8), Washingtonia (2) and Zombia (1); Ceroxyloideae – Ammandra (1), Aphandra (1), Ceroxylon (12), Juania (1), Oraniopsis (1), Phytelephas (6), Pseudophoenix (4) and Ravenea (20); Arecoideae – Acanthophoenix (3), Acrocomia (8), Actinokentia (2), Actinorhytis (1), Adonidia (2), Aiphanes (29), Allagoptera (5), Archontophoenix (6), Areca (45), Asterogyne (5), Astrocaryum (37), Attalea (76), Bactris (77), Balaka (9), Barcella (1), Basselinia (14), Beccariophoenix (2), Bentinckia (2), Brassiophoenix (2), Burretiokentia (5), Butia (22), Calyptrocalyx (26), Calyptrogyne (17), Calyptronoma (3), Carpentaria (1), Carpoxylon (1), Chamaedorea (106), Chambeyronia (2), Clinosperma (4), Clinostigma (11), Cocos (1), Cyphokentia (2), Cyphophoenix (4), Cyphosperma (5), Cyrtostachys (7), Deckenia (1), Desmoncus (24), Dictyocaryum (3), Dictyosperma (1), Dransfieldia (1), Drymophloeus (3), Dypsis (162), Elaeis (2), Euterpe (7), Gaussia (5), Geonoma (68), Hedyscepe (1), Heterospathe (41), Howea (2), Hydriastele (49), Hyophorbe (5), Hyospathe (4), Iguanura (32), Iriartea (1), Iriartella (2), Jailoloa (1), Jubaea (1), Jubaeopsis (1), Kentiopsis (4), Laccospadix (1), Lemurophoenix (1), Leopoldinia (2), Lepidorrhachis (1), Linospadix (7), Loxococcus (1), Lytocaryum (4), Manicaria (2), Manjekia (1), Marojejya (2), Masoala (2), Nenga (5),

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ARECALES Neonicholsonia (1), Neoveitchia (2), Nephrosperma (1), Normanbya (1), Oenocarpus (9), Oncosperma (5), Orania (28), Parajubaea (3), Pinanga (138), Pelagodoxa (1), Pholidostachys (8), Phoenicophorium (1), Physokentia (7), Podococcus (2), Ponapea (4), Prestoea (10), Ptychococcus (2), Ptychosperma (30), Reinhardtia (6), Rhopaloblaste (6), Rhopalostylis (2), Roscheria (1), Roystonea (10), Satakentia (1), Sclerosperma (3), Socratea (5), Solfia (1), Sommieria (1), Syagrus (53), Synechanthus (2), Tectiphiala (1), Veitchia (11), Verschaffeltia (1), Voanioala (1), Welfia (1), Wallaceodoxa (1), Wendlandiella (1), Wettinia (21) and Wodyetia (1). Uses: Palms are among the most useful of plant families. Man evolved in tropical ecosystems, and palms were certainly a major factor in allowing Homo sapiens to have easy access to plant products to be used for food, shelter, clothes, tools, ropes, drugs and weapons. Even though there was a great diversity of plants available to mankind, palms had a primary role and have been a dominant factor in the development of human culture ever since. Fruit: there are numerous species of which fruit or seed is consumed, especially coconut, Cocos nucifera, date, Phoenix dactylifera, and snake palm, Salacca zalacca, being currently of major economic importance. Other species with edible fruits cultivated as minor crops are seashore palm (Allagoptera arenaria), tucum (Astrocaryum vulgare), peach palm (Bactris gasipaes), lontar palm (Borassus flabellifer), Guadalupe palm (Brahea edulis), jelly palms (Butia capitata, B. yatay), lawyer vines (Calamus spp.), carnauba palm (Copernicia prunifera), triangle palm (Dypsis decaryi), acai (Euterpe oleracea, E. precatoria), doum palm (Hyphaene thebaica), Chilean wine palm (Jubaea chilensis), coco-de-mer (Lodoicea maldivica), moriche palm (Mauritia flexuosa), mangrove palm (Nypa fruticans), bacaba (Oenocarpus bacaba), palmetto (Sabal spp.), queen palm (Syagrus romanzoffiana) and many other species. Coconut was named in the 16th century for resemblance of the seed with its three holes to a skull, ‘coco’ meaning head in Spanish.

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Of Indo-Pacific origin, coconuts were valued throughout that region and soon spread to most tropical islands and continents. Because the seeds float, they rapidly colonised new shores, and even wash ashore occasionally on beaches in Europe. Coconuts were domesticated in early times and selected for meatiness of seeds and thinner shells. They were already known in the American Pacific before European discovery of the continents; however, they were only introduced to the Caribbean after the 1500s. Seeds yield oil used in food. The meaty part of coconuts is eaten fresh or desiccated, used as a spice in cooking or for production of coconut milk. Coconut can also be dried, fried, added to granola and other breakfast cereals and chocolate bars etc. The embryonic fluid is tapped, and because coconut water is isotonic, it is a refreshing drink. Date palms (Phoenix dactylifera) have been cultivated since ancient times throughout the Middle East and North Africa. Dates have a high sugar content and are a valued commodity in desert climates, where they form an easily transportable staple food for desert caravans. They are not known in the wild and may be of hybrid origin. Beverages: palm wine or toddy is a fermented drink made from the sap of a variety of palm species, especially Borassus spp., Caryota urens, Cocos nucifera, Elaeis guineensis, Hyphaene coriacea, Jubaea chilensis, Phoenix sylvestris and Raphia spp. Vegetable: terminal buds of some palms are harvested and eaten as ‘cabbage’ or ‘palm heart’, but this is not sustainable in most cases and has resulted in many species being threatened. Palm cabbage is known to be or has been harvested from Acanthophoenix, Bactris, Cocos, Deckenia, Euterpe, Geonoma, Prestoea, Roystonea and Sabal, of which clonal palms like Euterpe edulis and Prestoea acuminata should be preferred, because if carried out in moderation harvesting does not kill the palm. Canned palm heart is often harvested from plantations of Bactris gasipaes, grown for this purpose in South America. Oil: African oil palm, Elaeis guineensis, is of major commercial value and used in many processed food products and as a biofuel.

It has replaced vast stands of rainforest. Products containing palm oil are therefore ecologically unsound, and consumers should try to avoid them. American oil palm (E. oleifera), Acrocomia and, of course, coconut (Cocos nucifera) are also used for oil, the last now heralded as a superfood. Wax: several palm species yield a valuable wax. Carnauba wax, harvested from the leaf coating of Copernicia prunifera, is used to produce a glossy finish in car wax, shoe polish, dental floss, candy, furniture and floor wax, paper coating, cosmetics etc. Wax of Andean wax palm (Ceroxylon quindiuense) is formed at the base of the leaf sheaths around the trunk apex. It was threatened with extinction because the wax was used to make candles, but it is now rarely used for this purpose. Starch: sago is starch extracted from the spongy pith of several palm species, most notably Metroxylon sagu, M. amicarum and M. salomonense. It is the major staple food for tribes in New Guinea and the Moluku Islands. It is also cultivated and exported as sago flour. Rotten trunks of palms may house large palm weevil larvae (Rhynchophorus palmarum), which are considered a delicacy called ‘suri’ by local people in Amazonia. Drugs: seeds of betel nut (Areca catechu) are chewed with leaves or inflorescences of Piper betle (Piperaceae), lime and other ingredients. They contain an alkaloid, arecaine, that acts as a mild narcotic. Fruits are also used as a source of tannin for dyeing and in traditional medicine. Betel nut is commonly cultivated in the Old World tropics, sometimes in larger plantations but usually on a smaller scale. Actinorhytis calapparia is chewed as a betel substitute. Saw palmetto (Serenoa repens) is used in the treatment of prostate problems. Handicrafts: many species are used for their fibre, especially in handicrafts, rope, mats and loin cloths. Economically, the most important is rattan, mostly made from Calamus rotang and other climbing species (often called lawyer vines because they often have fish-hook spines that snag clothing and follow you around). Rattan makes climbing stems of over 10 m long, which are flexible and have an even thickness. They are therefore useful for woven-furniture making, basketry

COMMELINALES

MONOCOTS

and even umbrellas. Raphia produces raffia fibre, which is used for textiles, mats, ropes, shoes, natural string and garden ties that are often used for wrapping grafts. Giant leaves of Attalea were used for roofing by Mayan tribes, and many species of this and other genera are still commonly used for thatch. Dransfieldia is used to make harpoons. Seeds of Attalea maripa have been found in 9,000-year old archaeological sites in Colombia. Phytelephas produces ‘vegetable ivory’, hard seeds that can be carved into beautiful ornaments. Seeds of some Calamus species are polished and crafted into prayer beads. Ornamental: palms are gracious plants and have an association with the tropics, where they are commonly planted as ornamentals. Tropical beaches are usually adorned with coconut palms, and royal palms (Roystonea regia) are an all-time favourite for lining avenues. Several species are common as house plants in the temperate zones, especially species of Howea, Dypsis, Rhapis and Chamaedorea. In mild temperate

gardens, cold-tolerant species such as Brahea armata, Butia capitata, Chamaerops humilis, Jubaea chilensis, Livistona chinensis, Nannorhops ritchieana, Parajubaea torallyi, Phoenix canariensis, Rhapidophyllum hystrix, Sabal palmetto, Syagrus romanzoffiana, Trachycarpus fortunei, T. takil, T. wagnerianus, Washingtonia filifera and W. robusta are planted to provide a ‘tropical’ look to the garden. Records: Arecaceae excel in size. Lodoicea maldivica, the coco-de-mer, produces the largest seeds of any angiosperm, weighing up to 30 kg. Corypha umbraculifera produces the largest inflorescence of any plant, which can be 7.5 m long and bear around 10 million flowers. With climbing stems of c. 200 m long, Calamus manan produces the longest stems of any angiosperm, and because palms can be hundreds of years old and have no secondary thickening, they also have the oldest vessels of any vascular plants. Seeds of some species can be viable for hundreds of years,

and germination experiments on the extinct Easter Island palm (Paschalococos) are currently been trialled. A seed of a date palm (Phoenix dactylifera) found in a 2,000-yearold tomb in Israel has been germinated and is growing well. With 19 letters, Johannesteijsmannia is one of the longest plant genus names, although the record appears to be Pseudoblepharispermum (Asteraceae). The largest leaf of any plant is also grown by a palm: Raphia regalis has leaves measuring 25.11 × 3.00 m, the largest in length, albeit divided into hundreds of leaflets. This record may be contested by the vining leaves of Salpichlaena volubilis (Aspleniaceae), as they have indeterminate growth and climb into the canopy of the forest, but actual measurements of those climbing leaves are wanting and therefore Raphia regalis remains the world record holder. Etymology: Areca is a Latinised form of the common name for betel nuts in Malabar, India.

COMMELINALES Families 113 to 117 form the order Commelinales. This order diversified c. 110 million years ago.

113. HANGUANACEAE Susum family

These are terrestrial and aquatic, perennial, unisexual herbs that, when young, are covered with branched hairs. Stems are erect, but form creeping or floating stolons. Leaves are spirally arranged with clasping, opensheathing bases that gradually form a petiole topped by a simple, lanceolate to linear leaf blade. Venation is parallel with a prominent

midvein and many tertiary crossveins, and blades are rolled up in bud. Inflorescences are terminal and composed of spikelets organised in a panicle or in whorls. The sessile flowers are radially symmetrical and have six bractlike petals in two whorls, the outer ones small and basally connate, the inner ones larger and hooded. Male flowers have six stamens that have thin filaments, broadened at the base, and anthers with two thecae opening by longitudinal slits. Stamens surround a disc-like structure that envelops a rudimentary ovary. Female flowers have six staminodes, often minute, that surround a superior trilocular ovary with a sessile stigma. Fruits are berries with one or three bowl-shaped seeds. Distribution: This family occurs from Sri Lanka to Southeast Asia through Malesia and the Philippines. It extends into Micronesia

up to Palau and south into New Guinea and northern Australia. Hanguana grows in humid forests in swamps and lakes and along rivers. It may form dominant stands, suppressing other vegetation, and may even form floating islands. Phylogeny and evolution: Relationships of Hanguana were considered puzzling in the past. They share some morphological similarities with Asteliaceae and Zingiberales, but molecular evidence placed Hanguana firmly in Commelinales. Hanguana was previously associated with Flagellariaceae, but it was shown to differ from Poales in many anatomical characters. Its position within Commelinales is still uncertain, but it may be sister to Commelinaceae.

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COMMELINALES

Hanguana anthelminthica, flower detail, Singapore Botanical Garden [113]

Hanguana anthelminthica, Singapore Botanical Garden [113]

Hanguana rubinea, Singapore (MN) [113]

Genera and species: The single genus Hanguana consists of c. 13 species, possibly more.

it that lacks ligules or stipules. The blade is simple, often somewhat fleshy and has an entire margin and parallel venation. Flowers are formed on cymose branches (cincinni), which are then part of panicles or thyrses, axillary or terminally. Cincinni are often enclosed by a leafy bract (spathe). Flowers are radially or laterally symmetical and usually bisexual, although separate male or female flowers also occur, but usually in combination with bisexual flowers. The three sepals are free or fused and equal or unequal in size and can be sepaloid or petaloid. The three petals are all of the same size or one reduced and are free or basally fused and sometimes clawed. The six stamens occur in two whorls, all fertile or two, three or four staminodial and reduced or lacking. Filaments are glabrous or hairy with beaded or simple hairs, the anthers opening by slits, rarely by pores. The superior ovary is bi- or trilocular with a terminal style. Fruits are dehiscent capsules, fleshy indehiscent capsules or berries.

stomata and pollen anatomy rather than flower structure. Subfamily Cartonematoideae are sister to the remainder of the family and have previously been placed in a separate family, based mainly on anatomical characters. Commelinaceae are most likely sister to Hanguanaceae, from which they diverged c. 89 million years ago; together this pair is sister to the remainder of families in the order.

Uses: The aquatic Hanguana anthelminthica has edible stems, although it is rarely consumed. Etymology: ‘Hanguan kassintu’ is the local name used in the Sunda Islands (Indonesia) for this genus, which was Latinised to name the genus.

114. COMMELINACEAE Spiderwort family

Perennial and annual, usually terrestrial, herbs make up this family. They have fleshy stems that often root from the swollen nodes and may have tuberous roots, especially in temperate species. Leaves are placed alternately along stems, sometimes clustered into false whorls; they are sheathing at the base with the closed sheath enveloping part of the internode above 180

MONOCOTS

Christenhusz, Fay & Chase

Distribution: They occur pantropically, extending into temperate North America and East Asia. Phylogeny and evolution: Previously the family was subdivided into two groups with zygomorphic and actinomorphic flowers. This division still stands in subfamily Commelinoideae, although it is based on

Genera and species: This family has 41 genera and 731 species in two subfamilies: Cartonematoideae – Cartonema (7), Triceratella (1); Commelinoideae – Aëtheolirion (1), Amischotolype (23), Aneilema (65), Anthericopsis (1), Belosynapsis (6), Buforrestia (3), Callisia (20), Cochliostema (2), Coleotrype (10), Commelina (198), Cyanotis (49), Dichorisandra (47), Dictyospermum (5), Elasis (1), Floscopa (21), Geogenanthus (3), Gibasis (14), Gibasoides (1), Matudanthus (1), Murdannia (55), Palisota (25), Plowmanianthus (5), Pollia (19), Polyspatha (3), Porandra (3), Pseudoparis (3), Rhopalephora (4), Sauvallea (1), Siderasis (1), Spatholirion (6), Stanfieldiella (4), Streptolirion (1), Tapheocarpa (1), Thyrsanthemum (3), Tinantia (13), Tradescantia (74), Tricarpelema (8), Tripogandra (22) and Weldenia (1). Uses: Commelina communis is sometimes eaten as a vegetable in Asia. Flowers of this species yield a blue juice, which is used in

COMMELINALES

MONOCOTS

Aneilema aequinoctiale, Royal Botanic Gardens, Kew, UK [114]

Tradescantia hirsuticaulis, Sarah Duke Botanical Garden, Durham, North Carolinas, USA [114]

Commelina diffusa, Picinguaba, Brazil [114]

Cyanotis somaliensis, National Botanic Gardens of Ireland, Glasnevins [114]

Weldenia candida, Royal Botanic Gardens, Kew, UK [114]

Dichorisandra thyrsiflora, Royal Botanic Garden, Sydney, Australia [114]

Japan as a dye for colouring origami paper. It was commonly used as an ink for woodblock prints in 18th and 19th century Japan, but the blue colour is unstable in sunlight. A number of species are grown as ornamentals, as garden and house plants. Even though flowers are ephemeral, lasting only part of the day, they flower over long periods of time, making them valuable ornamentals.

Commelinus senior, who was a successful bookseller and newspaper publisher but did not botanise.

base and linear or sword-like blade, sometimes rounded and then with internal segments. Flowers are formed in axils of spathe-like bracts that are arranged in terminal, simple or compound spikes. The bisexual flowers are zygomorphic, the petals four, free or fused at the base, the outer two larger than the inner two, the upper one often two or three pointed at the tip, a result of fusion of two or three petals. The single stamen is placed above the lower petal and has a broad filament and an anther with two thecae opening by slits. The superior ovary is often somewhat fused with the base of the perianth and filament and has three unequal carpels, the median one often smaller than the lateral two. The fruit is a capsule that splits in threes, or a berry.

Etymology: Commelina is named for Dutch botanists Jan Commelijn (Johannus Commelinus, 1629–1692), founder of the hortus botanicus in Amsterdam, and his nephew Caspar Commelijn (1667–1734), who followed in his uncle’s footsteps. Linnaeus named the genus for the two botanists in the family because of the two well-developed petals of some species of Commelina, leaving its reduced petal to refer to Jan’s brother Casparus

115. PHILYDRACEAE Frogsmouth family

This is a family of perennial herbs with rhizomes and corms. Leaves are all crowded at the base of the plant or some along the flowering stem; they occur in a plane, with a sheathing

Distribution: They occur in Southeast Asia, New Guinea and Australia, with Philydrum Plants of the World

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COMMELINALES

MONOCOTS

Helmholtzia glaberrima, Australian National Botanic Gardens, Canberra [115]

Philydrella pygmaea, Perth, Western Australia

Philydrum lanuginosum, Helsinki Botanical Gardens, Finland [115]

Heteranthera reniformis, Santa Catarina, Brazil [116]

Eichhornia crassipes, Réunion [116]

Pontederia cordata, Chen Shan Botanical Garden, Shanghai, China [116]

lanuginosum extending north to southern Japan and south to Victoria (Australia).

116. PONTEDERIACEAE

herbs. Stems often have rhizomes or stolons. Leaves are alternate and arranged in a plane (distichous), sometimes whorled (in Hydrothrix). They are usually sheathing at base, the stipules large and enveloping the stem (ochrea) or reduced and forming a small ligule. Blades are simple and linear without a petiole or differentiated into petiole and blade. Petioles are sometimes inflated, forming floaters (Eichhornia). Blades are simple, linear to round or heart-shaped, the venation, following the leaf shape and confluent at tip and base. Flowers are solitary, paired or composed of bracteate panicles, spikes or false

[115]

Water-hyacinth family

Phylogeny and evolution: Philydraceae are most closely related to Pontederiaceae plus Haemodoraceae. Crown group Philydraceae are estimated to be c. 33–47 million years old. Genera and species: Philydraceae have three genera with six species: Helmholtzia (3), Philydrella (2) and Philydrum (1). Etymology: Philydrum is derived from Greek φίλος (philos), friend, and υδώρ (hydor), water.

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This is a family of perennial and annual, aquatic, submerged, floating and emergent

COMMELINALES

MONOCOTS

umbels. The bisexual flowers are zygomorphic (sometimes nearly actinomorphic) and have six, sometimes three or four, nearly free petals. The lower median tepal often has a yellow spot. The six, four, three or one stamens are all alike or of two kinds. Filaments are fused with the tepal tube and variously shaped, glabrous or hairy. Anthers have two thecae that are basifixed or appearing dorsifixed, opening by slits or pores. The superior ovary is trilocular but sometimes only one locule is fertile. The single style is usually simple and elongate. The small stigma is entire and round or trilobed. The fruit is a capsule that splits into three or a nutlet with a single seed.

Phylogeny and evolution: Fossil Pontederiaceae are known from Eocene deposits in North America and India. They are sister to Haemodoraceae, the pair then sister to Philydraceae. Diversification in the family is estimated to have occurred c. 40 million years ago. Eichhornia is polyphyletic, with E. meyeri probably sister to the entire family, although this needs further study. Genera and species: This family has six genera and 33 species: Eichhornia (6), Heteranthera (12), Hydrothrix (1), Monochoria (7), Pontederia (6) and Scholleropsis (1).

Distribution: This is a pantropical family, with Pontederia extending into temperate North America. Monochoria and Scholleropsis are Palaeotropical, whereas the others are found in the New World. Eichhornia crassipes has naturalised worldwide in warmer areas, causing havoc in waterways.

Uses: Monochoria leaves are sometimes eaten in Southeast Asia. The water hyacinth, Eichhornia crassipes, has many uses as animal fodder, mulch, compost, paper making and biofuel and for water cleaning. The species is a great pest in warm aquatic systems worldwide. Sometimes pickerelweed (Pontederia) is grown as a pond ornamental.

Anigozanthos manglesii, Perth, Western Australia [117]

Conostylis aculeata subsp. breviflora, Mt Benia, Western Australia [117]

Some tropical submerged species are offered in the aquarium trade but not frequently so. Etymology: Pontederia is named to commemorate Italian botanist Guilio Pontedera (1688–1757), who was a professor in Padua University and director of its botanical garden (the oldest surviving botanical garden). Even though he rejected Linnaeus’s sexual system of classification, Linnaeus named this genus for him.

117. HAEMODORACEAE Kangaroo-paw family

Wachendorfia thyrsiflora, private garden Kingston upon Thames, Surrey, UK [117]

Blancoa canescens, Mt Benia, Western Australia [117]

Haemodorum simplex, Mt Benia, Western Australia [117]

Macropidia fuliginosa, Mt Benia, Western Australia [117]

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COMMELINALES This is a family of perennial herbs with rhizomes, corms and bulbs that are usually reddish or orange. Leaves are basal or located along the flowering stem, alternately placed in one plane (distichous). The base is sheathing (like an Iris), and the blades are sword-like, unifacial, flat and lanceolate or tubular and hollow, sometimes plicately folded. Inflorescences are bracteate and racemose, often comprising a panicle, corymb or head of flowers. The bisexual flowers are actinomorphic or zygomorphic and have two whorls of three simple, basally fused petals, or the two whorls are fused into a single tube with six lobes, the tube splitting on one side when the flower opens (as in Anigozanthos). The three or six stamens are free or fused with the petal tube. Staminodia are sometimes present. Filaments are flattened, and anthers have two thecae opening by longitudinal slits. The superior ovary is tri- (sometimes uni-) locular and

MONOCOTS

bears a terminal style that is round or flattened on one side. The fruit is a capsule that splits in threes or opens by apical pores. Distribution: The greatest diversity in this family occurs in Western Australia. Three genera (nine species) occur in South Africa, and another three genera (five species) are found in tropical America. Lachnanthes caroliniana occurs in eastern North America, from coastal Canada to Cuba.

Blancoa (1), Conostylis (45), Macropidia (1), Phlebocarya (3) and Tribonanthes (6); Haemodoroideae – Barberetta (1), Dilatris (4), Haemodorum (20), Lachnanthes (1), Pyrrorhiza (1), Schiekia (1), Wachendorfia (4) and Xiphidium (c. 3).

Phylogeny and evolution: The stem lineage emerged c. 81 million years ago. Two subfamilies are recognised, which diverged during the early Eocene, c. 47 million years ago.

Uses: Roots of Haemodorum produce a red pigment, which was used to make a drink by the Australian aborigines. It is said to have antitumor and antibacterial properties. Roots of other genera also produce colourful pigments. Many species are valuable ornamental plants, especially species of Anigozanthos, of which many named hybrids are available in the plant trade. They are also sold as cut flowers.

Genera and species: This family comprises 14 genera with 102 species in two subfamilies: Conostylidoideae – Anigozanthos (11),

Etymology: Haemodorum is derived from the Greek αίμα (haima), blood, and δώρων (doron), a gift.

ZINGIBERALES Families 118 to 125 comprise the order Zingiberales. This clade is characterised by giant herbs that have leaves that are rolled up in bud with a distinct petiole and a sheathing base that in many cases forms or clasps a pseudostem. Their blades have a strong midvein and parallel S-curved sideveins. Inflorescence bracts are usually large and persistent, and the flowers are often large, monosymmetric, with variously fused petals and inferior ovaries. The clade evolved roughly 80 million years ago. This age is corroborated by a fossil seed (Spirematospermum) from the Late Cretaceous. These families are all similar in floral and vegetative structure and could be merged. All were traditionally placed together in the ‘family’ Scitaminae, but the separate families were accepted in APG IV because of convention.

118. STRELITZIACEAE Traveller’s-palm family

This is a family of perennial evergreen, often arborescent herbs with axillary suckers or underground runners. The stem is 184

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subterranean and dichotomously branching or erect and woody, with leaves in a terminal, fan-like cluster at the top. Leaves are alternate, arranged in a plane (distichous) and have a sheathing base, a distinct petiole and a pinnately veined blade with parallel, sigmoid lateral veins with crossveins. The basal sheath lacks stipules. The blade is entire, but may be ripped into irregular leaflets by the wind. Inflorescences are terminal or lateral thyrses of one to many spathe-like bracts in which flowers are arranged in axillary cincinnae. Flowers are bisexual, zygomorphic and subtended by ridged bracteoles. The six petals are organised in two unequal

whorls, the outer three free and inner three more or less fused. In Strelitzia, two inner petals are fused to form an arrow-shaped structure that encloses stamens and style in a central groove. Stamens are basically six (in Ravenala), often reduced to five (in other taxa), but no staminode is present. Anthers are basifixed, and the two thecae open by slits. The inferior ovary is trilocular and bears a single filiform style and sunken nectaries. The fruit is a capsule that splits in three, and seeds have brightly coloured, furry arils, orange in Strelitzia, blue in Ravenala and red in Phenakospermum. Flowers are pollinated by birds and lemurs.

ZINGIBERALES

MONOCOTS

Strelitzia reginae, Royal Ravenala madagascariensis, Kusu Botanic Gardens, Kew, UK [118] Island, Singapore [118]

Distribution: Tropical South America, coastal South Africa and Madagascar. Phylogeny and evolution: Previously, the family was treated as a tribe of Musaceae, often placed together with Heliconia. Both are now segregated in separate families because to make Musaceae monophyletic in the traditional sense, Marantaceae and Zingiberaceae would also have to be included. Molecular phylogenetic analyses place Strelitziaceae as sister to the remainder of the order. Strelitziaceae diverged c. 59 million years ago.

Strelitzia nicolai, Santa Barbara, California, USA [118]

the United Kingdom. She was an amateur botanist who lived in Kew Palace for many years and contracted William Aiton and Joseph Banks to develop the gardens at Kew initiated by her mother-in-law, Princess Augusta; these would later become the Royal Botanic Gardens.

119. LOWIACEAE Orchid-lily family

Genera and species: This family includes three genera and seven (or more) species: Phenakospermum (1), Ravenala (1, possibly more) and Strelitzia (5). Uses: Traveller’s pal m, Ravenala madagascariensis, is commonly planted as a garden ornamental in the tropics. Strelitzia nicolai and S. reginae are also commonly available as garden plants. Inflorescences of bird-of-paradise-flower (S. reginae) are popular in the cut-flower trade, but relatively expensive. A yellow-flowered form found at Stellenbosch was named ‘Mandela’s Gold’ in honour of Nelson Mandela. Etymology: Strelitzia was named by Joseph Banks in honour of Charlotte of MecklenburgStrelitz (1744–1818), Queen to George III of

Orchidantha maxillarioides, Royal Botanic Gardens, Kew, UK [119]

long-lived and can bear new branches. The bisexual flowers are zygomorphic, fragrant, usually having a foul odour that attracts beetles. The three sepals are almost equal, narrow and fused into a tube at the base. The three petals are unequal in size and shape, the two lateral ones narrow and small and the middle one enlarged, forming a lip (labellum). The five free stamens have short filaments fused to the petal bases. Anthers have a short connective tip and open by slits. The inferior ovary is composed of three fused carpels that bear nectaries and a elongate style with a trilobed stigma. Fruits are dry capsules that split in three upon ripening, bearing numerous round hairy seeds. Distribution: Restricted to tropical Asia, this family can be found in China, Hainan, Indochina, Malaya, Borneo and Luzon.

A family of glabrous herbs, Lowiaceae are generally terrestrial along streams and near waterfalls in evergreen rainforest or forest clearings. They grow from a horizontal or vertical rhizome that bears scale-like leaves. Leaves are formed at the rhizome apices in distichous tufts. The base is sheathing without stipules, the petiole well-defined and blades lanceolate and entire with pinnate venation composed of a strong central vein and parallel lateral veins that are connected by crossveins. Flowers are solitary on terminal branches (paracladia), held upside down and lasting for only a single day, although inflorescences are

Phylogeny and evolution: This family is most closely allied to Strelitziacae. Genera and species: The single genus Orchidantha has c. 18 species. Etymology: Lowia is named after British colonial administator and naturalist Sir Hugh Lowe (1824–1905), the son of Scottish horticulturalist Hugh Lowe. He made the first documented ascent of Mount Kinabalu in Borneo. Lowia is a later synonym of Orchidantha, which is Greek for ‘orchid flower’. Plants of the World

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ZINGIBERALES

120. HELICONIACEAE Parrot-flower family

This family consists of large, evergreen herbs that grow from a rhizome or produce short erect trunks formed by leaf sheaths. Leaves are sheathing at the base, the sheath without stipules and petioles well defined, often elongate. Blades are often large with a strong midvein and parallel, sigmoid side veins that fuse at the margins and are cross-connected by parallel tertiary veins. Blades are entire, but large ones may be irregularly ripped along

Heliconia psittacorum, cultivated, Réunion [120]

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lateral veins. Inflorescences are terminal and composed of thyrsoid, distichous or helical racemes formed by spathe-like bracts in which cincinnae of flower clusters are formed. The obliquely symmetrical, bisexual flowers are subtended by crested bracts. The six petals and filaments are fused at the base, but five of these petals are fused completely to form a sheath; the remaining petal is free. One of six stamens is staminodial. Filaments are fused to petals, and the anther is basifixed with two thecae opening by slits. The inferior ovary is trilocular with a single filiform style and a capitate stigma. The fruit is a three-seeded berry that is often a peculiar blue or metallic colour. Flowers are pollinated mostly by hummingbirds in the New World and bats in the Pacific. Distribution: Most species are found in tropical America, but a few species are known from the Pacific, Sulawesi and New Guinea to Samoa, Fiji and New Caledonia.

Phylogeny and evolution: Heliconiaceae diverged from Zingiberaceae c. 88 million years ago and diversified c. 32 million years ago. The clade has no fossil record. Traditionally, they have been included in a broader Musaceae, but this would be non-monophyletic if Zingiberaceae and Marantaceae were excluded. Genera and species: This family consists of the single genus Heliconia with 197 species. Uses: In times of famine, flower buds have been eaten in Melanesia. Many species are grown as garden ornamentals or for the cut-flower industry, the colourful bracts lasting for a long time in flower arrangements. Etymology: Latin Heliconia is derived from Ελίκων (Helikon), a mountain in Greece of mythological importance, which in turn is derived from ήλιος (helios), the sun, and κώνος (konos), a cone.

Heliconia bihai, Guadeloupe [120]

Heliconia caribaea, Guadeloupe [120]

Heliconia rostrata, fruit, Tapirai, Brazil [120]

Heliconia rostrata, cultivated, Réunion [120]

ZINGIBERALES

MONOCOTS

Musella lasiocarpa, Kunming Botanical Garden, China [121]

Musa sikkimensis ‘Red Tiger’, male flowers, Royal Horticultural Society Garden, Wisley, UK [121]

Ensete ventricosum, Wundanyi, Taita Hills, Kenya [121]

121. MUSACEAE Banana family

This is a family of giant herbs, some of which are among the largest herbs known. Rhizomes are single or freely suckering, the stems monocarpic (dying after flowering). Alternate, spirally arranged leaves are sheathing at the base, forming a pseudostem, with elongate petioles. The large blades are entire but often split by the wind to the midrib along the parallel sigmoid side veins.

Musa basjoo, Royal Botanic Gardens, Kew, UK [121]

Lateral veins fuse near the margin and are interconnected with smaller cross-veins. The inflorescence is a terminal, indeterminate thyrse that bears a cincinnus of one or two rows in the axils of fleshy spathe-like bracts that recurve when f lowers are opening. The usually functionally unisexual (rarely bisexual) flowers lack bracteoles, and the females form at the base of the inflorescence, whereas the males occur near the apex. The zygomorphic flowers are composed of six petals, of which the outer three and the inner two are fused into a five-lobed or five-toothed sheath, the inner free petal is usually small and opposed. Stamens are usually five, often with a staminodium, which in some cases can be fertile. Filaments are free, and anthers are basifixed with two thecae opening by slits. The inferior ovary is trilocular with a single style that is often wider at the stigma. The fruit is a starchy berry with a leathery skin

Musa acuminata ‘Cavendish’, female flower, Tapirai, Brazil [121]

Musa laterita, Royal Botanic Gardens, Kew, UK [121]

and many large, hard seeds. In some cases, the fruit is ‘self-peeling’. Distribution: Musaceae occur in tropical Africa, Madagascar, South and Southeast Asia, southern China, Malesia, the Philippines, New Guinea, Queensland and Melanesia. Phylogeny and evolution: The crown group is estimated to be c. 40–60 million years old. Eocene seeds of Ensete are known from Oregon, and fossil Spirematospermum seeds from the Tertiary of Europe are also linked to Musaceae. Fossil banana-like leaves from the Early Tertiary of Greenland, Musopsis, could be in this family or another one in Zingiberales. Musella is sister to the rest of the family. Genera and species: This family has three genera with c. 80 species: Ensete (6), Musa (c. 70) and Musella (1).

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ZINGIBERALES Uses: A great variety of bananas are in cultivation. The most popular are the seedless cultivars that are usually triploid and derived from hybridisation between Musa acuminata and M. balbisiana or from just one of these species. The hybrid is often given the name M. ×paradisiaca, which refers to an ancient belief that this was the ‘apple of paradise’, the forbidden fruit of wisdom (which includes the synonym, M. sapientum). Bananas have a history of cultivation and hybridisation for thousands of years in Southeast Asia and China but are now a global commodity. Bananas are, after citrus, the most important fruit crop in the world, typically grown in large monocultures of the cultivar ‘Cavendish’. Fruits of dessert bananas are eaten raw or are juiced, deep-fried, cooked, frozen or dried and chipped. They are even used to make a ketchup-like product (with food colouring to make it red) or to brew beer. Plantains (starch bananas) are grown as a minor crop often in backyard vegetable gardens in the tropics. Plantains are generally boiled or fried before eating. Bananas and plantains are a staple food in most tropical countries. Male buds of Musa inflorescences are cooked as a vegetable in Asia. Leaves are used to wrap food when cooking and are sometimes used as an alternative for a plate or instead of an umbrella in rain. Swollen stems of Ensete ventricosum are used as a source for starch in East Africa. It has been introduced for this reason to the American tropics, where it now naturalises. Manila hemp, Musa textilis, originating in Borneo, is an important fibre crop in the Philippines and Pacific used to make rope and textiles. The Japanese fibre banana, Musa balbisiana var. liukiuense, is a cold hardy selection from the plantain that yields a fine, high-quality fibre highly valued in Japan. Several species such as Musa coccinea, M. ornata, M. splendida, M. velutina and other species of Musa, Ensete and Musella are popular garden ornamentals. Musa basjoo, M. sikkimensis, M. yunnanensis and Musella lasiocarpa have some frost tolerance, making them popular for a tropical touch in temperate gardens. Some of these are now being used to make hardy seedless banana cultivars.

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Etymology: Musa is the Latinised form of ‫زوم‬ (mauz), the Arabic name for banana. Giant banana: Musa ingens from New Guinea can grow over 15 m tall and can have trunks with a two metre circumference. It has leaves more than five metres long and a metre wide (almost as large as the 5.51 m2 of Victoria amazonica, Nymphaeaceae) and produces a giant bunch of inedible bananas with up to 20 hands weighing around 60 kg. It is the largest herb in the world.

122. CANNACEAE Canna-lily family

The fruit is usually a tuberculate, bristly capsule splitting into three. Seeds are hard and round and have been reported to remain viable for 600 years. Distribution: This family occurs in tropical and subtropical areas of the Americas from eastern North America (South Carolina) to southern South America (northern Argentina). They are now cultivated worldwide. Phylogeny and evolution: Morphological and molecular analyses show a close relationship of Cannaceae to Marantaceae. These two are maintained mainly because of their differences in stamen structure. Cannaceae are c. 30 million years old. Canna flaccida from North America is sister to all other species of Canna. Genera and species: The family consists of the single genus Canna with c. 10 species.

This is a family of perennial herbs with tuberous, starchy rhizomes and slime canals in all vegetative tissues. Leaves are alternate, often spiral, but sometimes in one plane (distichous). They are sheathing at the base with an open sheath, lacking stipules, that gradually merges into distinct petioles. The blade is rolled up in bud and is simple and entire with pinnate venation. The midvein is prominent with secondary veins parallel and merging into a vein along the margin. Inf lorescences are terminal thyrses with bracteate cincinnae of flowers held above the leaves. By reduction, this inflorescence may look like a raceme. The bisexual flowers are asymmetrical. The three sepals are free, green or purplish and persistent in fruit. The three petals are unequal in size, one smaller, fused into a tube at the base. The single stamen is petal-like, bearing a single theca along the edge. Other male parts are staminodial, the one to five staminodes petallike, one large one opposite the fertile stamen (labellum) and the others smaller (wings). The inferior ovary is trilocular with a petal-like style and a wet papillate stigmatic surface.

Uses: Edible rhizomes of achira, Canna discolor (syn.: C. edulis), are cultivated throughout the tropics as a staple for humans and livestock. Its starch granules are large and can be seen with the naked eye. Originating in the northern Andes, where it was domesticated around 2500 BC, plants are now grown commercially mainly in India and Australia, where it is known as Queensland arrowroot. A favourite bedding plant in tropical and temperate gardens, there are now thousands of Canna cultivars available, mostly of complex hybrid origin. Canna ×generalis cultivars are hybrids derived initially from C. glauca and C. indica, later also involving C. iridiflora and C. warszewiczii. Hybrids with large lipped flowers were then bred by hybridising C. ×generalis with C. flaccida, forming a cultivar group called C. ×orchidoides. Over time, these two cultivar groups were interbred, and horticultural varieties of Canna are now called only by their cultivar name without a hybrid or species epithet. Etymology: Latin canna (cane) originated from cána, the Irish Celtic word for a cane or reed.

ZINGIBERALES

MONOCOTS

Stromanthe sanguinea, Hong Kong Botanical Garden [123]

Canna glauca ‘Panache’, University of Wisconsin Botanical Garden, Madison, USA [122]

Canna indica var. limbata, Royal Botanic Gardens, Kew, UK [122]

Schumannianthus dichotomus (WA) [123]

Thalia geniculata, Royal Botanic Gardens, Kew, UK [123]

Goeppertia loeseneri, Singapore Botanical Garden [123]

Maranta arundinacea, Singapore Botanical Garden [123]

123. MARANTACEAE

aquatics. They are rosulate and stemless or with elongate branched stems and then lianescent or shrubby. Arial stems are solid and can be bamboo-like with swollen nodes and dry leaf sheaths. Leaves are arranged alternately, usually in one plane (distichous), and are divided into a basal clasping sheath, a distinct petiole that may be absent in some species, a thickened pulvinus and a simple leaf blade. The blade is entire and pinnately veined with a strong midvein and closely set S-shaped secondary veins. The base of the blade is often oblique or sides of the leaf blade are of different size. Blades can be uniformly green, or variously coloured, mottled, striped

or spotted. Flowers are formed in clusters of cymules subtended by a prophyll and arranged in bracteate heads or spikes (thyrses) that are simple or compound. The bisexual flowers are asymmetrical. The three sepals are free, but petals, stamens and staminodes are fused into a tube at least at the base. An outer whorl of staminodes is present and petal-like or rudimentary. An inner whorl is fused at the base into a tube that exceeds the outer tube. It is three-parted: one is fertile and often has a petal-like appendage, one is hood-shaped and the third is firm and fleshy. The inferior ovary is trilocular with two locules often reduced and infertile. The single terminal style is

Prayer-plant family

This is a family of variably sized herbs with rhizomes that are sometimes starchy. They occur in the forest understory and as emergent

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189

ZINGIBERALES fused with the floral tube and often covered by a hood-like staminode in trigger flowers. The stigma is a funnel-shaped depression at the style tip. The fruit is usually a dehiscent capsule, rarely a fleshy berry or nutlet. Distribution: This is a pantropical family of rainforests extending into the warm temperate zones of North and South America.

190

MONOCOTS

horticulturally important species) is to be treated under the genus Goeppertia, making it the largest genus in the family.

and cultivars of these, are popular houseplants or garden plants in the tropics. Thalia dealbata and T. geniculata are popular pond plants.

Genera and species: Marantaceae include 27 genera and c. 555 species: Afrocalathea (1), Calathea (37), Goeppertia (c. 250), Halopegia (3), Haumania (5), Hylaeanthe (5), Hypselodelphys (8), Indianthus (1), Ischnosiphon (36), Koernickanthe (1), Maranta (43), Marantochloa (18), Megaphrynium (5), Monophyllanthe (2), Monotagma (39), Myrosma (1), Phrynium (38), Pleiostachya (3), Sanblasia (1), Saranthe (10), Sarcophrynium (6), Schumannianthus (2), Stachyphrynium (9), Stromanthe (20), Thalia (6), Thaumatococcus (2) and Trachyphrynium (1).

Etymology: Maranta is named for Italian physician and botanist Bartolomeo Maranta (1500–1571), who is most famous for a book on poison antidotes.

124. COSTACEAE

Phylogeny and evolution: Marantaceae are estimated to have diverged from Cannaceae c. 79 million years ago, and the crown group of Marantaceae diversified some 57 million years ago, although sampling of Marantaceae in molecular dating studies has not been adequate for good estimation. They may have originated in Africa, where the diversity of the family is poor, but from there they dispersed to Asia and South America where they diversified. Generic delimitation in Marantaceae is still problematic and even though molecular studies have helped to realign some of these, there remain many issues. Phrynium is paraphyletic. Stachyphrynium and Donax have been readjusted to reflect monophyly, and Calathea was found to be polyphyletic with regard to Ischnosiphon, Monotagma, Pleiostachya and Sanblasia. Instead of merging these three genera with Calathea, it was decided that the majority of Calathea (including the

Uses: Economically, the most important species is arrowroot, Maranta arundinacea, a species from the Caribbean that yields a high-quality starch from the rhizomes. Sweet corn root, Goeppertia allouia, is sometimes grown in the West Indies for its edible tuber. Some species of Maranta and Goeppertia are grown by local peoples for their starchy rhizomes and inflorescences. Larger species of Ischnosiphon, Marantochloa and Schumannianthus are sometimes used for basketry. A great diversity of prayer plants, especially Goeppertia lancifolia, G. makoyana, Maranta bicolor, M. leuconeura

These terrestrial and rarely epiphytic, hairy herbs are perennial and rhizomatous, without oil cells. Stems are round and not or irregularly branched, and often spirally contorted. Leaves are spirally arranged and have a closed tubular sheath with a ligule that extends around the stem above the petiole insertion. Petioles are short and winged, the wing grading into the elliptic to linear, entire blade, which is rolled in bud. Venation is pinnate, the midvein strong, lateral veins

Costus arabicus in fruit, Royal Botanic Gardens, Kew, UK [124]

Hellenia speciosa, Singapore Botanical Garden [124]

Chamaecostus cuspidatus, private collection, Kingston upon Thames, Surrey, UK [124]

Christenhusz, Fay & Chase

Spiral-ginger family

ZINGIBERALES

MONOCOTS

arching toward the apex and partly running parallel to the midvein. The inflorescence is a cone-like spike terminating in a leafy shoot or on separate leafless shoots. Flowers are rarely solitary in the leaf axils (Monocostus). Bracts of inflorescences are imbricate and spirally arranged, often fleshy with nectar glands covering one or two flowers. The bisexual, zygomorphic flowers have sepals fused into a bi- or tri-lobed tube. The petals are fused into a trilobed corolla, the lobes unequal in size. The single stamen is often petal-like and bears two thecae that open by slits. The often more or less trilobed labellum, formed from fused petal-like staminodia, opposes the stamen and is often larger than the corolla. The labellum is fused with the stamen into a small tube. The inferior ovary is tri- (or bi-) locular with two nectaries and a filiform style that is usually enclosed by the thecae of the stamen. The fruit is a capsule that splits into three (or two) or breaks up irregularly. Distribution: This is a pantropical family. Phylogeny and evolution: The clade has been dated to c. 47 million years old. It is most closely related to Zingiberaceae, in which it was previously included as a subfamily. Cheilocostus is a synonym of Hellenia. Genera and species: This is a family of seven genera and 137 species: Chamaecostus (7), Costus (106), Dimerocostus (3), Hellenia (2), Monocostus (1), Paracostus (2) and Tapeinochilos (16). Uses: Unlike the closely related and similar Zingiberaceae, the plants are not aromatic and are therefore not used for spice, although some are used in traditional medicine. Fruit of Dimerocostus is eaten locally in South America. Crepe ginger, Hellenia speciosa, from tropical Asia is used for starch in India, and this species is also the source of diosgenin, used in the production of steroids. Some species of Costus, Hellenia and Tapeinochilos are commonly grown as garden ornamentals in the tropics. Etymology: Costus is Latinised Ancient

Greek meaning “from the East”, referring to the Indian lower Himalaya region from where spices were imported into Greece and Rome.

125. ZINGIBERACEAE Ginger family

This family of perennial herbs has numerous oil cells, making all parts aromatic. Creeping rhizomes are often branched and succulent, or fleshy roots (stilt-roots) can sometimes elevate the rhizome above the ground. Leaves are alternate in a single plane (distichous), either in the same plane as the rhizome or perpendicular to it. Leaf bases have an open sheath clasping the stem; ligules may be present at the tip of the sheath. Blades are usually sessile (or blades are absent at the base of shoots) and simple, elliptic to linear, rolled up in bud with an entire margin. A pulvinus is absent except in Zingiber. Venation is pinnate with a strong midvein and parallel, sinusoid lateral veins. Inflorescences either terminate a leafy shoot, or leaves and inflorescences occur on separate shoots. Inflorescences are lax or contracted racemes, spikes, heads or cone-like, or with the flowers in cincinnae in axils of a bract, forming a thyrse. Bracts are separate and papery or imbricate and fleshy. The bisexual, zygomorphic flowers have a tubular calyx composed of three fused sepals that are trilobed or three-toothed and sometimes split along one side. The three petals are fused into a trilobed corolla, which is tubular basally and has lobes that vary in shape and size, often one much larger than the other two. Only one of six stamens is fertile. In the outer whorl, one staminode is rudimentary, and the other two are petal-like. In the inner whorl, two staminodes are fused into a labellum-like structure. The remaining fertile stamen may not have a filament but

bears an anther with two thecae opening by slits or rarely by pores. The connective is often prolonged into a crest and sometimes spurs at the base. The inferior ovary is initially trilocular, but during development may become unilocular. The style is thin and almost always placed between the thecae, the stigmas are wet and funnel-shaped, often also crested and situated on top of the stamen. Two nectaries occur at the top of the ovary at the base of the style. The fruit is usually a dry or fleshy capsule that splits regularly in three or irregularly in an undefined number, sometimes not dehiscent. Sometimes fruits are united into a fleshy syncarp, which is dispersed as a single unit. Distribution: This is a pantropical family with species extending into the subtropics and temperate zones in Japan, the Himalayas and South Africa. Renealmia is the only American genus and arrived there less than 16 million years ago. Phylogeny and evolution: An age of 65 million years has been suggested for this family, verified by the fossil Zingiberopsis of the Upper Cretaceous or Lower Eocene of North America, but the major diversification took place 26 million years ago. The family is a member of a clade with Cannaceae, Costaceae and Marantaceae, which share petal-like staminodes. Several genera were found to be paraphyletic, and studies on some genera have resulted in a broader concept of, for instance, Globba (including Mantisia) and Curcuma (with inclusion of Laosanthus, Paracautleya, Stahlianthus, Smithatris and some Kaempferia and Hitchenia species). Pyrgophyllum is included in Camptandra. Large genera like Alpinia and Amomum are polyphyletic and are in need of revision. Genera and species: This family has 51 genera and c. 1,600 species in four subfamilies: Siphonochiloideae – Aulotandra (6) and Siphonochilus (11); Tamijioideae – Tamijia (1); Alpinioideae – Aframomum (55), Alpinia (241, polyphyletic), Amomum (188, polyphyletic), Burbridgea (5), Cyphostigma (1), Elettaria (11), Elettariopsis (23), Etlingera (100),

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ZINGIBERALES

MONOCOTS

Alpinia zerumbet, Hong Kong Botanical Garden [125]

Hedychium coronarium, Royal Botanic Gardens, Kew, UK [125]

Roscoea bhutanica, Royal Botanic Gardens, Kew, UK [125]

Globba patens, Mt Datuk, Malaysia [125]

Zingiber zerumbet, New Caledonia [125]

Etlingera elatior, Singapore Botanical Garden [125]

Geocharis (6), Geostachys (25), Hornstedtia (33), Leptosolena (1), Paramomum (1), Plagiostachys (27), Pleuranthodium (23), Renealmia (85), Riedelia (76), Siamanthus (1), Siliquamomum (2) and Vanoverberghia (2); Zingiberoideae – Boesenbergia (68), Borneocola (8), Camptandra (4), Caulokaempferia (30), Cautleya (2), Cornukaempferia (3), Curcuma (109), Distichochlamys (4), Gagnepainia (3), Globba (c. 100), Haniffia (3), Haplochorema (6), Hedychium (93), Hemiorchis (3), Hitchenia (1), Kaempferia (33), Kedhalia (1), Larsenianthus (4), Myxochlamys (2), Nanochilus (1), Newmania (2), Parakaempferia (1), Pommereschea (2), Rhynchanthus (4), Roscoea (21), Scaphochlamys (25), Stadiochilus (1) and Zingiber (144).

a desired commodity in classical times, probably being one of the first exotic spices to be used in Europe, already widely in use by wealthy Romans. The young, mild and fragrant rhizomes are often pickled, candied or cooked fresh in a number dishes. With honey it makes a delicious, anti-inflammatory tea. Sugary stem ginger or candied ginger is frequently used in baked goods or made into a spicy candy. When dried, it is a fragrant, hot spice, used in curries and biscuits, including ginger nuts, gingerbread, ginger snaps and speculaas, and for the production of ginger beer, ginger wine, canton and ginger ale. Ginger is also sliced finely and pickled in Japanese cuisine often served with sushi. Outside Asia, ginger is cultivated on a large scale in Jamaica. Turmeric (Curcuma longa and, to a lesser extent, C. aromatica) has been cultivated in

India for millennia and is used fresh as a spice or dried under heat as a spice or dye. It is native to tropical Asia, where it is commonly grown and used in local cuisine (curries etc.). Fresh leaves can also be used to wrap food before cooking. As a food dye it is known in the EU under E100 and gives a custard yellow colour found in many processed foods and canned beverages. It is often confused with saffron (Crocus sativus, Iridaceae), which has a different hue and flavour. Even though turmeric makes a poor fabric dye because it is light-unstable, it is frequently used for colouring robes of Buddhist monks and Indian saris. Zedoary or white turmeric (C. zedoaria) is also used as a spice, but in Europe where it was already known in the 6th century, it was replaced by ginger. It smells like mango, tastes like ginger, is bitter and is still used in Indian and Thai cuisine.

Uses: Ginger (Zingiber officinale) has been used since ancient times in India and was

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Native to tropical Asia, immature fruits of green cardamom (Elettaria cardamomum) have been used as a spice in local cuisines since ancient times. It was first introduced to Europe in the late 13th century and has ever since been a popular spice, especially in Scandinavia to flavour traditional cakes and pastries and in the Middle East to flavour coffee and tea. Grains of paradise (Aframomum melegueta) is a spice with a pungent, citrusy flavour. Common in West and North African cuisines, it was known in the Mediterranean during ancient times. Today, it is known for its digestive properties and is commercially used to flavour beer, gin and Norwegian aquavit. It is also used in the Caribbean in voodoo rituals. It is commonly used as a spice in East African cuisine and locally to flavour coffee. A number of other species are also used as spice. Nepal cardamom (Amomum subulatum) is used mostly in India and Pakistan and black cardamom (Hornstedtia

costata) in foods in Vietnam and Sichuan. Amomum villosum is cultivated in China for its cardamom-like fruits. Greater galangal or laos (Alpinia galanga) is a spice popular in Thai and Indonesian cuisines, whereas lesser galangal (Alpinia officinarum) flavours Southeast Asian curries, drinks and perfumes, and Russian vinegar and Nastoika liqueur. The Australian ginger (A. caerulea) has edible fruits and can be used as a spice, but is not grown commercially. Korarima or false cardamom (Aframomum corrorima), araththa (Alpinia calcarata), Chinese cardamom (A. globosa), rathkihiriya (A. malaccensis), black ginger (A. nigra), shell ginger (A. zerumbet), fingerroot (Boesenbergia rotunda), kencur (Kaempferia galanga), myoga (Zingiber mioga), cassumunar ginger (Z. montanum), awapuhi (Z. zerumbet) and Renealmia alpina are all minor culinary herbs or spices. The young shoots of some species, such as the porcelain rose (Etlingera elatior) and Tamijia flagellaris, are eaten as a vegetable.

Alpinia nutans is added to food as a preservative in Japan. Many species are cultivated as garden ornamentals in the tropics and warmer temperate regions, especially Alpinia purpurata, A. speciosa, Curcuma alismatifolia, Etlingera elatior and other species of Alpinia, Globba and Hedychium. Several species of Roscoea and Hedychium, Cautleya robusta and Zingiber mioga are frost-tolerant and can be grown in temperate gardens. Hedychium species can become problematic invasives in tropical mountain forest habitats, crowding out the native vegetation and preventing regeneration of natural woodland. Etymology: Zingiber is derived from Sanskrit srngam, a horn, and vera, a body or root, which came into Greek as ζιγγίβερης (ziggiberis), into Latin as zingiberi and gingiber, which became gingimbre in Old French and ginger in modern English, all names for the spice that originated in India.

POALES Families 126 to 139 represent the order Poales. This order evolved c. 100 million years ago (based on molecular calculations), and the crown group is dated to 83 million years, although some fossils are older, such as Protoananas lucenae (Bromeliaceae) from Brazil which is c. 113 million years old and Protograminis laminatus (Poaceae, Pooideae) from Burma, which is estimated to be c. 105 million years old. Therefore, the lineage may be much older than suggested, unless these fossils have not been properly identified. It is unusual to have fossil age estimates for clades that are older than those estimated from molecular clocks.

126. TYPHACEAE Bulrush family

This is a family of glabrous, amphibious herbs with scaly, starchy, creeping rhizomes. They

are usually emergent, growing in muddy soil, or submerged with floating leaves. Leaves are alternate, distichous (in one plane) and sheathing basally. The sheath sometimes forms a short erect stem. Leaf blades are sessile and linear with parallel venation. Inflorescences are racemose, but the unisexual flowers are clustered in dense heads or in cylindrical, cigar-like spikes. Petals are one, three or many and can be small and scale-like or slender and bristle-like. Male flowers usually have three stamens (sometimes fewer or more) with free or fused filaments and basifixed anthers. Female flowers have a superior ovary composed of one

to three fused carpels with a single style and one to three stigmas. The fruit is an indehiscent nutlet, which is plumed in Typha. Distribution: The family is nearly cosmopolitan, apart from the desert regions of Arabia, the Gobi and the Sahara. It is also absent from Borneo and Sulawesi. Phylogeny and evolution: The family is dated to at least 80 million years, and the two genera diverged c. 72 million years ago. Diversification within the genera is more recent. The family has a rich fossil history,

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MONOCOTS

Typha dominguensis in fruit, Palm Springs, California, USA [126]

Typha minima, Royal Botanic Gardens, Kew, UK

Sparganium erectum, near Cambridge, England, UK [126]

Sparganium angustifolium, Isokari, Finland [126]

the earliest possibly dating from the end of the Cretaceous. Sparganium is sometimes placed in its own family, Sparganiaceae, but the two genera share many characters and are merged in APG IV.

127. BROMELIACEAE

and large. Flowers are bisexual or functionally unisexual, usually regular, sometimes slightly oblique, with three fused or free sepals and three fused or free petals. The six stamens form two whorls of three and have free or fused filaments. Anthers have two thecae and open by latrorse slits. The inferior to superior ovary is trilocular and bears a single style with three variously ornate stigmas. Fruits are usually septicidal capsules that split in three or a berry. Seeds are sometimes winged or plumed. Pollination is predominantly by hummingbirds, but other birds, bats and insects (bees, bumble bees, moths and butterflies) are also reported to visit flowers.

[126]

Pineapple family

Genera and species: This family has two genera with c. 50 species: Sparganium (c. 20) and Typha (30). Uses: All parts of Typha are edible, and in the past rhizomes and young male spikes, even pollen, have been consumed. Typha is, like reeds (Phragmites, Poaceae), used for biofiltration of surface water. Both Typha and Sparganium are sometimes grown as pond ornamentals. Etymology: Τυφά (Typha) is the Greek name for cattail, possibly derived from τύφος (tyfos), smoke, in reference to the seeds that may form a cloud when dispersing.

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This is a family of epiphytic and terrestrial, rosette-forming herbs with short or elongate stems that sometimes become woody and tree-like. Roots are usually present, but absent in some epiphytes. Leaves are spirally arranged, simple, entire or prickly toothed, broadened and clasping or sheathing at the base without stipules or ligules. Flowers are formed in terminal, bracteate spikes or condensed heads, the bracts often colourful

Distribution: The family is widespread in the tropical and warm temperate Americas, north to Virginia and south to Patagonia. A single species (Pitcairnia feliciana) occurs in West Africa, a result of recent long-distance dispersal.

POALES

MONOCOTS

Phylogeny and evolution: The stem lineage of Bromeliaceae is dated to 100 million years, but the crown group is much more recent, c. 20 million years old, the majority of the diversification in the family having occurred much more recently. A fossil from the mid-Tertiary of Costa Rica, Karatophyllum bromelioides, is until now the only fossil that can reliably be assigned to this family. Other fossils like the 113 million-year-old fossil of Protoananas has a more dubious relationship to Bromeliaceae. Generic delimitation needs attention, with several found to be non-monophyletic in recent molecular analyses. Eight subfamilies are sometimes accepted. Genera and species: Bromeliaceae include c. 62 genera with c. 3,475 species: Acanthostachys (2), Aechmea (283), Alcantarea (36), Ananas (3), Androlepis (2), Araeococcus (9), Billbergia (65), Brewcaria (6), Brocchinia Guzmania megastachya, Guadeloupe [127]

Bromelia antiacantha, Royal Botanic Gardens, Kew, UK [127]

(20), Bromelia (63), Canistropsis (11), Canistrum (13), Catopsis (19), Cipuropsis (10), Connellia (6), Cottendorfia (1), Cryptanthus (78), Deinacanthon (1), Deuterocohnia (20), Disteganthus (2), Dyckia (163), Edmundoa (3), Eduandrea (1), Encholirium (28), Fascicularia (1), Fernseea (2), Fosterella (31), Glomeropitcairnia (2), Goudaea (2), Gregbrownia (4), Greigia (36), Guzmania (218), Hechtia (64), Hohenbergia (68), Hohenbergiopsis (1), Lapanthus (3), Lindmania (39), Lymania (9), Navia (93), Neoglaziovia (3), Neoregelia (125), Nidularium (47), Ochagavia (4), Orthophytum (67), Pitcairnia (c. 410), Portea (8), Pseudaechmea (1), Puya (230), Quesnelia (23), Racinaea (76), Ronnbergia (13), Sequencia (1), Sincoraea (1), Steyerbromelia (6), Stigmatodon (18), Tillandsia (c. 700), Ursulaea (2), Vriesea (c. 225), Werauhia (96), Wittrockia (7), Waltillia (1), and Zizkaea (1).

Uses: Pineapple, Ananas comosus, is the most important economic plant of this family. The fruit is composed of berries that coalesce with the fleshy bracts and rachis of the inflorescence, forming a fleshy, juicy compound fruit. Pineapples can be eaten fresh or cooked, juiced, dried or jammed, and are used in many cuisines. Pineapples are rich in bromelain, an enzyme that breaks down proteins and can be used as a meat tenderiser and in the pharmaceutical industry. Some species of Bromelia also have edible fruits but are only consumed locally. Leaves of pineapple plants are sometimes used to make fibre used in wallpaper etc. Several species produce useful, strong fibres, especially caroá, Neoglaziovia variegata, and caraguá, Ananas lucidus, which are commonly used in South America. Puya chilensis fibres are used to make durable fishing nets.

Ananas comosus, plantation in Paraná, Brazil [127]

Tillandsia usneoides clothing trees in North Carolina, USA [127]

Pitcairnia macrobotrys, Royal Botanic Gardens, Kew, UK [127]

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POALES Spanish moss, Tillandsia usneoides, is a rootless epiphyte that was formerly important in the southern USA as packing material and for stuffing of pillows and cushions. Spanish moss and other epiphytic Tillandsia species are good indicators of air quality. Many bromeliads are cultivated as ornamentals. Giants, dwarfs and carnivores: Queen of the Andes, Puya raimondii, is the largest of all bromeliads, making a vegetative growth to c. 4 metres and up to 10 metres tall when flowering, with an inflorescence bearing over 3,000 flowers. The related Puya chilensis is infamous for being hazardous to sheep and birds, which can get impaled or otherwise caught by its leaves. It has therefore been considered to be inadvertently carnivorous because the decaying corpses can provide nutrients to the plants. Many species of bromeliads collect water in the middle of their rosette of tightly overlapping leaves. These cups,

Guacamaya superba, habit, Colombia (MF) [128]

Schoenocephalium teretifolium, Colombia (MF) [128]

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called phytotelmata, often house entire ecosystems, including treefrogs, land crabs, insects, earthworms, protists and even aquatic angiosperms (e.g. Utricularia, Lentibulariaceae) and algae. A highly endemic fauna is found in these tanks. A few species are believed to have perfected this cup into a trap for insects to gain additional nutrients. Most famous is Brocchinia reducta, which has UV-reflecting scales on the insides of the cup that lure insects that fall into the liquid, where they drown and decompose. The plant does not produce enzymes as do some carnivorous plants in other families, but it is considered a carnivorous plant in the true sense because it absorbs the nutrients through adventitious roots. Similarly, Catopsis berteroniana has also been suspected of carnivory. Tillandsia includes some of the smallest bromeliad species, and being air plants lacking roots, species like T. recurvata can cover electricity and telephone wires in tropical America.

Etymology: Bromelia commemorates Swedish physician and botanist Olaus Bromelius (Olof af Bromell, 1639–1705), who was famed for owning a large botanical library.

128. RAPATEACEAE Tow-tow family

Terrestrial and epiphytic, sometimes emergent aquatic, perennial herbs comprise this family. They have creeping rhizomes with leaves clustered at the tip of a short stem emerging from the rhizome. Leaves are mostly in one

Rapatea paludosa, Colombia (MF) [128]

Duckea cyperaceoidea, Colombia (MF) [128]

Monotrema bracteatum, Colombia (MF) [128]

POALES

MONOCOTS

Xyris sp., habit, Bertioga, Brazil [129]

Xyris sp., Bertioga, Brazil [129] Xyris involucrata, Colombia (MF) [129]

plane (distichous), and fan-shaped, but are sometimes somewhat spiralling or twisting, the bases asymmetrical and flattened, with ligules or petioles rarely present at the tip, sometimes with the blades perpendicular to bases. Leaf blades are grass-like, linear or lanceolate, sometimes elliptic, with parallel venation, the veins meeting at the tip and base of the leaf. Inflorescences are terminal or axillary from a leaf sheath and solitary or many in a head. Flowers are formed in bracteate spikelets, which in turn are aggregated in heads that are surrounded by two, rounded or flattened bracts. The bisexual flowers are regular or slightly zygomorphic. They have three sepals and three petals that are fused at the base into a small tube. The six stamens are in two whorls, and the filaments are basally fused to each other or to the corolla tube. The anthers are basifixed, often with apical appendages and two thecae that open by one, two or four apical pores. The superior ovary is trilocular (or unilocular by abortion of the other two) and bears a single erect style with a capitate stigma. The fruit is a septicidal capsule.

Monot remoideae, Rapateoideae and Saxofriedericioideae. Rapateaceae occupy an isolated position in Poales, as sister to what some authors have termed core Poales (the combined cyperid and graminid clades). Genera and species: Rapateaceae have 17 genera with 94 species: Amphiphyllum (1), Cephalostemon (5), Duckea (4), Epidryos (3), Guacamaya (1), Kunhardtia (2), Marahuacaea (1), Maschalocephalus (1), Monotrema (4), Phelpsiella (1), Potarophytum (1), Rapatea (22), Saxofridericia (8), Schoenocephalium (4), Spathanthus (2), Stegolepis (33) and Windsorina (1). Etymology: Rapatea is derived from an Amerindian name for the plant, probably from the Wayampi language in French Guiana.

129. XYRIDACEAE Yellow-eyed-grass family

Distribution: This family occurs in tropical South America and coastal West Africa. The single African representative, Maschalocephalus dinklagei, arrived there by recent long-distance dispersal. Phylogeny and evolution: The origin of Rapateaceae is dated to c. 79 million years ago. It has no known fossil record. The family is sometimes split into three subfamilies:

Abolboda pulchella, Colombia (MF) [129]

unifacial, laterally flattened, rarely rounded or angular. Lateral or terminal inflorescences have a long peduncle, emerging from a specialised sheath or from the inner leaves, with or without bracts along the stem. Bracts are present below the flowers. Each peduncle bears one-, rarely two bracteate spikes or heads, or sometimes a panicle composed of spikes. The actinomorphic, bisexual flowers are one to many per inflorescence, each singly in the axils of a bract. Usually there is only one or a few open in the inflorescence at any time. The three sepals are all equal in size and shape, or one is reduced to a scale or absent. In Xyris the sepals are thin and wrapped around the corolla. Three petals are usually equal, sometimes unequal, free or fused into a two-lipped corolla, the bases clawed or fused into a narrow tube. Three (rarely six) stamens oppose the petals. Anthers are bilocular, opening by slits. Staminodia are usually three, sometimes one, filamentous or branched, often densely hairy. The superior ovary is unilocular or partly trilocular with a terminal three-branched or simple style with a variably shaped stigma. The fruit is a capsule that usually splits into three. Distribution: This is a pantropical family, extending to temperate eastern North America (Great Lakes and Nova Scotia) and temperate Australia and Tasmania.

This is a family of perennial herbs with leaves emerging from a creeping or erect rhizome. Leaves are spirally arranged or in a fan, the bases broad and open, sheathing, the sheaths often flat and keeled, and the blades usually

Phylogeny and evolution: Xyridaceae are most closely related to Eriocaulaceae and Mayacaceae, with which they share several characters and habitat preferences. In this part of the graminid/cyperid clade, relationships Plants of the World

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have not always been well resolved or well supported. The stem group of Xyridaceae is c. 87 million years old. The family is sometimes divided into two subfamilies, Abolbodoideae and Xyridoideae. Genera and species: Xyridaceae include five genera and c. 395 species: Abolboda (21), Achlyphila (1), Aratitiyopea (1), Orectanthe (2) and Xyris (c. 370). Etymology: Xyris is derived from the Greek ξυράφι (xyrafi), a razor, in reference to the sharply edged leaves of some species.

130. ERIOCAULACEAE Pipewort family

This is a family of perennial and sometimes annual, terrestrial and aquatic herbs. Plants are usually rosulate and tufted but can have all leaves along an erect stem in some genera. Stems are usually hairy. Leaves are usually spirally arranged, sometimes two-ranked, with or without a clasping base, linear to

Syngonanthus caulescens, Colombia (MF) [130]

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lanceolate with parallel venation. Flowers are borne in dense head-like spikes on thin peduncles that may be solitary or compound in various several-times-umbellate structures composed of several to hundreds of these heads. Each peduncle has a closed sheath at the base. Each head is subtended by fused bracts (involucre) and contains ten or more usually unisexual flowers, each subtended by smaller free bracts that can be petal-like. Flowers are usually regular but can be zygomorphic. Sepals are two or three and various in colour, the petals are two or three, usually colourless or white. Stamens can be in one whorl, then only two or three, or in two whorls, then four or six or reduced to a single stamen. Anthers are dorsifixed or basifixed and have two cells. The superior ovary is bi- or trilocular with the same number of styles. The fruit is a loculicidal capsule spliting on the side. Distribution: This is a pantropical family, extending into the temperate zones in southern California, north to the Great Lakes and Newfoundland in eastern North America, south to Argentina; also in Scotland, Ireland, Sub-Saharan Africa, Madagascar, Asia north to the Himalayas, Manchuria, eastern Siberia and Japan and south to eastern Australia (Victoria). The family is especially diverse in the Guiana Shield and Atlantic Brazil. Phylogeny and evolution: Because of their unique inflorescence and pollen type, Eriocaulaceae have always been considered a

Tonina fluviatilis, Bertioga, Brazil [130]

natural entity. The clade (stem node) is at least 58 million years old. Generic delimitation in Eriocaulaceae is debated and still unclear, and genera such as Syngonanthus may have to be divided or others may have to be merged with it. They are closely related to Xyridaceae. Genera and species: This family has ten genera and c. 1,207 species: Actinocephalus (33), Comanthera (41), Eriocaulon (c. 480), Lachnocaulon (7), Leiothrix (64), Mesanthemum (15), Paepalanthus (420), Rondonanthus (6), Syngonanthus (140) and Tonina (1). Uses: Syngonanthus chrysanthus is occasionally grown as a houseplant. Inflorescences of several species are collected in Brazil for dried flower arrangements, although this harvesting is not believed to be sustainable. Golden grass (Syngonanthus nitens) is used for handicrafts. Tonina fluviatilis is occasionally cultivated as an oxygenating plant for tropical ponds or aquaria. Carnivory: Paepalanthus bromelioides shares the habitat of Brocchinia reducta (Bromeliaceae), its leaves also forming a tank that may similarly function to catch insects, although this is speculative. No study has been carried out on possible trapping, digesting or absorbing mechanisms in this species. Etymology: Eriocaulon is composed from the Greek έριο (erion), wool, and καυλος (caulos), stem or penis.

Eriocaulon decangulare by W. J. Hooker from Curtis’s Botanical Magazine vol. 59: 3126, 1832 [130]

Paepalanthus bromelioides, Caraguatatuba, Brazil [130]

POALES

MONOCOTS

Mayaca fluviatilis, Colombia (MF) [131]

Mayaca fluviatilis, Singapore (KH) [131]

Thurnia sphaerocephala, Guyana (PM) [132]

Prionium serratum, Karatara River, Western Cape, South Africa (CD) [132]

131. MAYACACEAE

is unilocular with a single style and short stigma. The fruit is a loculicidal capsule that splits in three.

132. THURNIACEAE

Bog-moss family

Palmiet family

Distribution: Mayacaceae occur in tropical America from southern North America to Uruguay and southern tropical Africa.

This is a small family of rooted, submerged, aquatic herbs that lack vessels in stems and leaves. Narrow, linear or filiform leaves are spirally arranged and sessile, and often have two teeth at the tip. Flowers are solitary in leaf axils, although seemingly terminating a stem and growing from a broad prophyll or bract. Flowers occur on pedicels that lift the flower above the water surface. Flowers are bisexual and have a differentiated calyx and corolla. The three sepals are green, the three petals white and narrowed at the base. The three stamens alternate with the petals and have slender filaments and basifixed anthers that open by apical pores. The superior ovary

Phylogeny and evolution: Previously, the family was thought to be related to Commelinaceae, but it is now known to be sister to a clade of Thurniaceae, Juncaceae and Cyperaceae or to Eriocaulaceae plus Xyridaceae. The stem lineage of Mayacaceae diverged from their relatives c. 100 million years ago. Genera and species: The family consists of the single genus Mayaca with six species. Uses: Mayaca fluviatilis is sometimes used as an aquarium plant. Etymology: Mayaca is an Amerindian name for the plant, probably from the Wayampi language as its name was first recorded in French Guiana.

This family of perennial herbs has erect woody trunks and subterranean rhizomes. Leaves are spirally arranged or three-ranked and clustered near the apex of the stems. They are linear and sheathing or clasping the stem at the base. Venation is parallel, the margins serrate. The inflorescence is a terminal bracteate panicle or a dense, globose head subtended by leafy bracts. Flowers are bisexual with six equal and dry petals in two whorls. The six stamens are free, the filaments slightly fused with the petals, and the basifixed anthers open by slits. The superior ovary is tricarpellate, the tip of the ovary elongated into a style with three free branches and dry Plants of the World

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POALES stigmas. The fruit is a loculicidal capsule that splits in three. Distribution: Thurniaceae occur in Guiana Shield in South America (Thurnia) and South Africa (Prionium). Phylogeny and evolution: The family is closely related to Juncaceae and Cyperaceae and was previously included in the former by many authors. They share tetrad pollen with Juncaceae, but Juncaceae lack the silica bodies common in Thurniaceae and Cyperaceae. The family is c. 98 million years old, but the crown node diverged c. 33 million years ago. Genera and species: This family has two genera with four species: Prionium (1) and Thurnia (3). Etymology: Thurnia is named in honour of the discoverer of the genus, British botanist, explorer and photographer Sir Everard Ferdinand im Thurn (1852–1932), who was curator of the British Guyana Museum, one of the first naturalists to climb Mount Roraima and later governor of Fiji.

133. JUNCACEAE Rush family

MONOCOTS

is formed of three fused carpels, topped by a three-branched filiform style. The fruit is a loculicidal capsule that splits in three. Distribution: The family occurs worldwide but is remarkably absent from the Amazon region and large parts of subtropical Africa, Madagascar and northern Australia. Phylogeny and evolution: Juncaceae are not well represented in the fossil record. Luzula is known from the Upper Oligocene in Germany, and there are Miocene fossils and fossil pollen known from Russia and North America. Molecular dating estimates this family to be c. 74 million years old. Genera and species: The family has seven genera and 460 species: Distichia (3), Juncus (325), Luzula (120), Marsippospermum (4), Oxychloë (5), Patosia (1) and Rostkovia (2). Uses: Juncaceae are often important pasture plants, but they have little other use. Juncus effusus and J. rigidus are sometimes woven into mats and into children’s toys (in a similar manner to corn dollies made from cereal straw). The pith of J. effusus can be used as a lamp-oil wick. Distichia is used for fuel in the Andes. Some species of Luzula are occasionally offered as garden ornamentals. Etymology: Juncus is the classical Latin name for rush.

134. CYPERACEAE Sedge family

Flowering stems of these plants are erect, the inflorescences terminal or appearing lateral, often composed of compound open cymose panicles or contracted into heads. Flowers are sometimes solitary. The bisexual or unisexual flowers usually have six (rarely four) free petals that are arranged in two whorls. The usually six stamens are also in two whorls, although the inner whorl may be absent. Filaments are filiform or flattened, and the anthers have two locules opening by slits. The superior ovary

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rhizomes that sometimes form stolons, bulbs or an erect trunk. Stems with nodes emerge from a leaf sheath, which is sometimes thickened at the base, often triangular in cross-section, or round to slightly flattened and sometimes winged. Leaves form a basal rosette, or there are few to many along the stem, rarely all cauline, usually spirally arranged in three (sometimes two) ranks. Lowermost, often underground, leaves are frequently reduced to sheaths. Leaf sheaths are usually closed, with or without a ligule on top. Blades are usually linear, sometimes a little broader and oblong, or inrolled and round in cross-section with parallel venation. The bisexual or functionally unisexual flowers occur in spikelets, which in turn form capitula, corymbs, panicles, spikes or umbels or are reduced to a single solitary spikelet. Spikelets are composed of bracts arranged along a slender axis (called a rachilla), with one to many bracts (glumes) subtending florets. Spikelets usually have a prophyll, which in some taxa (e.g. Carex) becomes bottle-shaped and encloses the rachilla and single flower. Flowers lack petals, but often have three to six (or many) scales that are bristle-like or rarely flat and laminar. There are usually three stamens, but sometimes one, two, four or six (or more), the filaments often elongating, usually free, with a basifixed anther that has an arrow-shaped base and connective often forming a crest. The superior ovary is usually unilocular, formed by three fused carpels and is therefore more or less triangular, or when of two carpels, somewhat flattened. The style is usually long and has three papillate stigmatic branches. The fruit is usually a nutlet, rarely a small drupe. Distribution: This is one of the most widespread families. They are cosmopolitan but absent from Antarctica and the permanent ice sheet of Greenland.

These annual and perennial herbs, shrubs and lianas grow terrestrially or sometimes epiphytically, often on wet ground or in aquatic habitats. Perennial species usually have

Phylogeny and evolution: Cyperaceae are sister to Juncaceae. The fossil record of Cyperaceae pollen and fruit starts in the Palaeocene. It is well represented in Cenozoic deposits of North America and Europe. The crown node of Cyperaceae is dated to be c. 76 million years old, and Cyperoideae diversified

POALES

MONOCOTS

Luzula campestris, near Turku, Finland [133]

Luzula alopecurus, Royal Botanic Gardens, Kew, UK [133]

Mesomelaena tetragona, Western Australia [134]

Caustis dioica, Mt Benia, Western Australia [134]

Luzula luzuloides, Royal Botanic Gardens, Kew, UK [133]

Cyperus papyrus, Jardin des Plantes, Paris [134]

Rhynchospora alba, Lake District, England, UK [134]

Juncus effusus, Lake District, England, UK [133]

Eriophorum angustifolium, Lake District, England, UK [134]

Schoenoplectus californicus var. paschalis, Easter Island [134]

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POALES c. 77 million years ago. Mapanioideae diversified later, c. 33 million years ago. Mapanioid fossils are known from the Eocene of Europe. Genera and species: Cyperaceae include 100 genera with c. 5,500 species and are divided in two subfamilies: Mapanioideae – Capitularina (1), Chorizandra (5), Chrysitrix (4), Diplasia (1), Exocarya (1), Hypolytrum (60), Lepironia (1), Mapania (85), Paramapania (7), Principina (1) and Scirpodendron (2); Cyperoideae – Actinoschoenus (3), Actinoscirpus (1), Afroscirpoides (1), Afrotrilepis (2), Amphiscirpus (1), Androtrichum (2), Arthrostylis (1), Ascolepis (22), Becquerelia (7), Bisboeckelera (3), Blysmus (4), Bolboschoenus (15), Bulbostylis (215), Calyptrocarya (8), Capeobolus (1), Carex (1,829), Carpha (15), Caustis (5), Cephalocarpus (4), Cladium (3), Coleochloa (8), Costularia (24), Crosslandia (1), Cyathochaeta (5), Cyathocoma (3), Cymophyllus (1), Cyperus (704), Cypringlea (3), Diplacrum (9), Dracoscirpoides (3), Dulichium (1), Eleocharis (284), Epischoenus (8), Eriophorum (16), Erioscirpus (2), Evandra (2), Everardia (11), Ficinia (77), Fimbristylis (305), Fuirena (59), Gahnia (41), Gymnoschoenus (2), Hellmuthia (1), Isolepis (76), Karinia (1), Khaosokia (1), Kobresia (58), Koyamaea (1), Kyllinga (80), Lagenocarpus (30), Lepidosperma (74), Lipocarpha (35), Machaerina (50), Mesomelaena (5), Microdracoides (1), Morelotia (2), Neesenbeckia (1), Nelmesia (1), Nemum (8), Neoscirpus (1), Oreobolopsis (3), Oreobolus (17), Phylloscirpus (3), Pleurostachys (33), Pseudoschoenus (1), Ptilothrix (1), Pycreus (118), Reedia (1), Rhodoscirpus (1) Rhynchocladium (1), Rhynchospora (354), Schoenoplectiella (51), Schoenoplectus (29), Schoenoxiphium (21), Schoenus (108), Scirpoides (3), Scirpus (50), Scleria (252), Sumatroscirpus (1), Tetraria (54), Trachystylis (1), Trianoptiles (3), Trichophorum (12), Trichoschoenus (1), Tricostularia (5), Trilepis (5), Uncinia (69), Volkiella (1) and Zameioscirpus (3). Uses: Pith of papyrus (Cyperus papyrus) is still used to make a paper-like material.

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Papyrus is a native of the Upper Nile in South Sudan but was already in ancient times brought to the Lower Nile in Egypt, where it soon naturalised and covered the shores. It is probably the earliest account of an invasive species caused by human introduction. The oldest known papyrus documents date from c. 2555 BC, but the method of using papyrus probably developed some 500 years earlier. Stems are first peeled and the inner pith cut lengthwise into thin slices, which are then placed on a flat surface with the edges overlapping. It is then covered with a second layer at right angles to the first. While still moist, the two sticky layers are pounded together, and then pressure is applied under which the sheet dries. When dry the sheet is polished with a round, smooth stone or wooden tool. Papyrus is suitable for use as paper in dry climates and was commonly used around the Mediterranean and the Middle East, but in damp climates it is not suitable because the cellulose soon becomes mouldy. In the 1st century, papyrus became more expensive and was soon replaced by the cheaper and more durable parchment and later by wood-based paper. By the 18th century, C. papyrus had become extinct in Egypt. Machine-manufactured papyrus was developed in the 1960s, and currently there are minor papyrus industries in Egypt and Sicily to satisfy the tourist market. In Central Africa papyrus is also used to make handicrafts, mats, furniture etc. It has also been under consideration for production of biofuel in some African countries. When harvested sustainably, this has great potential. It also has potential to be used for biofiltration of eutrophic or polluted water. The earth almond, or yellow nutsedge (Cyperus esculentus), has been cultivated for its edible tubers since 3000 BC in Egypt and spread from there across Europe, Africa and Asia. The tubers, called tigernuts, used to be sold as a treat in many European countries but have gone out of fashion. It is well known for the Spanish drink, horchata de chufa. The species is underused, especially because it is considered one of the worst invasive weeds. Tubers of the purple nutsedge (C. rotundus) can also be eaten but have a bitter taste.

Totora reeds (Schoenoplectus californicus) are locally woven into boats and rafts, especially in Lake Titicaca and on Easter Island. Some species of Schoenoplectus are used for water and sewage treatment. Several other species of Cyperaceae are locally used for weaving mats and baskets. A number of species are sometimes grown as ornamentals, mainly members of the genera Carex, Caustis, Cyperus, Eriophorum, Schoenoplectus and Uncinia. The umbrellaplant (Cyperus involucratus) is a common houseplant. Etymology: Cyperus is the Latinised form of the classical Greek name for nutsedge, κύπερος (kyperos).

135. RESTIONACEAE Fynbos family

This family consists of evergreen, perennial, r ush-like, usually unisexual plants, sometimes diminutive and ephemeral. Their photosynthetic stems are arranged in clumps arising from creeping rhizomes. The round, flattened or quadrangular stems are simple, forking or branching in whorls and can be confused with horsetails (Equisetaceae) or bamboos (Poaceae). Leaves are usually reduced to yellow, tan or brown sheaths surrounding nodes, split to the base on one side. Sheath tips often have thin lobes or a membranaceous margin, sometimes extending into an awn or reduced leaf blade. On juvenile stems leaf blades are often more developed, sometimes also persistent in adult plants, or in some taxa leaves are normally developed, in basal rosettes with laterally flattened (Anarthria) or terete blades with open sheaths (Centrolepidoideae). The flowers are arranged in bracteate spikes or branched

POALES

MONOCOTS

Anarthria scabra, male, near Albany, Western Australia [135]

Anarthria scabra, female, near Albany, Western Australia [135]

Elegia capensis, female, National Botanic Gardens of Ireland, Glasnevin [135]

Rhodocoma capensis, National Botanic Gardens of Ireland, Glasnevin [135]

Lyginia imberbis, female, near Perth, Western Australia [135]

Alexgeorgea subterranea, Mt Benia, Western Australia [135]

racemes, often sexually differentiated. Flower spathes can be large, covering the f lowers or small and unremarkable, sometimes condensed into two-ranked or head-like spikelets. Flowers can be solitary or arranged in spikelets and surrounded by one or two bractlets, often with the lower bractlet flowerless (as in grasses). Flowers are usually unisexual with a six-parted perianth, usually in two whorls of three, sometimes fewer or perianth absent. Male flowers have six or three stamens with free (sometimes fused) filaments and dorsifixed dangling anthers and usually a minute pistillode, in Centrolepidoideae reduced to a single stamen. Female f lowers have staminodes or lack stamens altogether. The ovary is superior, uni- to trilocular and free, with one to three, bushy styles that are sometimes fused at the base and have finely branched stigmas, or are filiform and covered with stigmatic papillae. In Centrolepidoideae the female f lowers

are single carpels with a unilocular ovary, sometimes united with adjacent carpels into pseudanthia. Fruits are capsules, nutlets or follicles opening by a slit. Distribution: Well known from southern Africa, the family also occurs in Madagascar, Southeast Asia, Malesia, Australia, New Zealand, Melanesia, southern South America and the Falkland Islands. It is particularly diverse in South Africa and Australia. Phylogeny and evolution: Distinctive fossil pollen shows that the family dates back at least to the Late Cretaceous and may have a Gondwanan origin, which corresponds with its modern-day distribution, although because the crown group of Restionaceae is dated to 74 million years old, its current distribution is more likely to be a result of long-distance dispersal. Fossil pollen shows that it may have been widespread in the Northern

Desmocladus fasciculatus, male, near Perth, Western Australia [135]

Centrolepis aristata, near Perth, Western Australia [135]

Hemisphere during the Tertiary, although the identity of the pollen is uncertain, and it may belong to any of the related families. Former Anarthriaceae (including Hopkinsiaceae and Lyginiaceae) form a sister lineage to other Restionaceae and can be maintained as a family, but these are so similar that it seems obvious to unite the two. The clade evolved c. 55 million years ago. Molecular studies place Centrolepidaceae on a long branch as sister to Australian Restionaceae (Leptocarpoideae and Sporadanthoideae), together forming a clade sister to the African Restionoideae. Therefore they are included in Restionaceae in their own subfamily. Centrolepidoideae split from other Australian Restionaceae c. 45–97 million years ago. Hydatellaceae, which are superficially similar, were previously treated as part of this family, but are now known to belong to Nymphaeales, a case of convergent evolution.

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POALES Genera and species: This is a family with five subfamilies including c. 55 genera and c. 572 species: Anarthrioideae – Anarthria (6), Hopkinsia (2) and Lyginia (3); Restionoideae – Anthochortus (15), Askidiosperma (12), Calopis (23), Cannomois (12), Ceratocaryum (6), Coleocarya (1), Dielsia (1), Elegia (51), Hydrophilus (1), Hypodiscus (15), Ischyrolepis (48), Mastersiella (13), Nevillea (3), Platycaulos (c. 8), Restio (c. 100), Rhodocoma (7), Soroveta (1), Staberoha (c. 9), Thamnochortus (34) and Willdenowia (12); Centrolepidoideae – Aphelia (6), Centrolepis (26) and Gaimardia (4); Leptocarpoideae – Alexgeorgea (3), Apodasmia (4), Baloskion (8), Catacolea (1), Chaetanthus (3), Chordifex (20), Cytogonidium (1), Dapsilanthus (4), Desmocladus (15), Empodisma (3), Eurychorda (1), Harperia (4), Hypolaena (8), Kulinia (1), Lepidobolus (c. 10), Leptocarpus (3), Loxocarya (5), Meeboldina (11), Melanostachya (1), Onychosepalum (3), Platychorda (2), Stenotalis (1), Taraxis (1), Tremulina (2), Tyrbastes (1) and Winifredia (1); Sporadanthoideae – Calorophus (2), Lepyrodia (22) and Sporadanthus (8), Uses: Few species are used commercially. Thamnochortus insignis is locally used in South Africa for thatching. Fynbos vegetation has contributed to the formation of peat in South Africa, which is being exploited unsustainably. Some species are used as garden ornamentals, with Elegia capensis being the most commonly used species due to its frost tolerance, although other Cape Restionoideae are gaining favour.

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136. FLAGELLARIACEAE Whip-vine family

Etymology: Flagellaria is named for the leaves that have a flagelliform or whip-shaped tip (a tendril).

137. JOINVILLEACEAE Ohe family

This family includes herbs with perennial shoots and clustered rhizomes. Stems are solid and glabrous. Leaves are alternate, clasping the nodes, in a plane (distichous). Leaf bases are sheathing with the sheath closed around the stem. Blades are lanceolate, with a constricted base and a long-acuminate tip that is curled up into a tendril (flagellum), with which it climbs. Inflorescences are spikes composed of a terminal panicle-like racemes. Flowers are bisexual with six persistent tepals in two whorls, the bases of which are slightly fused and thickened. Each tepal has a single vein. The six stamens are in two whorls with short filaments and the linear basifixed anthers open by slits. The superior ovary is composed of three fused carpels, trilocular and topped by three short spreading styles. Fruits are red or black, one-seeded berries. Distribution: This family is found in tropical and southern Africa, throughout the Indian Ocean islands to India, Malesia and the western Pacific.

Underground: Alexgeorgea subterranea was discovered by Sherwin Carlquist in 1974 near Jurien Bay in Western Australia. At first he only found male plants of this new species, but he needed female plants to describe it. He then noted purple thread-like structures emerging from the sand, which, after some digging, turned out to be the styles of the otherwise underground female flowers.

Phylogeny and evolution: Previously, Hanguana and Joinvillea were included in this family, but the relationship with Hanguana (Commelinales) is remote, and Joinvillea, even though in Poales, is also excluded on morphological and phylogenetic grounds. Flagellaria is sister to Joinvilleaceae plus Poaceae/Ecdeiocoleaceae. Flagellaria-type pollen is known from the Upper Miocene with certainty, and Eurasian material from the Tertiary probably also belongs here. The stem node is c. 90 million years old.

Etymology: Restio means ‘maker of rope’ and is derived from Latin restis, a rope or cord.

Genera and species: The sole genus, Flagellaria, has four species.

Christenhusz, Fay & Chase

Uses: Some species are used for basketry. Flagellaria indica has some medicinal properties.

This family of large herbs grows from a short rhizome and has unbranched, hollow, round or slightly flattenened shoots with solid nodes. Leaves surround the stem at nodes and are sheathing basally, the sheath opens with a ligule-like structure. Blades are linearlanceolate with parallel venation in which veins merge at the margin and tip, and the lamina is plicately folded. Blades are usually hairy. Inflorescences terminate stems and are many-flowered, pendent panicles composed of spikes or racemes. Flowers are bisexual, regular, and trimerous. The six free or nearly free tepals are bract-like, dry and persistent in fruit. Each tepal has three veins. Stamens are free or somewhat fused with the tepal bases, the anthers basifixed, arrow-shaped and opening by slits. The superior ovary is trilocular and has three free styles with feathery stigmas. Fruits are drupes with one to three hard seeds. Distribution: This family occurs on islands in Malesia into the Pacific, east to Hawaii and south to New Caledonia. Phylogeny and evolution: Joinvillea was previously included in Flagellariaceae and was only segragated in 1970 on the basis of growth form, indument, leaf morphology and some other anatomical characters. It is a relictual group, c. 89 million years old (stem

POALES

MONOCOTS

Flagellaria indica, along the Daly River, Northern Territory, Australia [136]

Flagellaria neocaledonia, New Caledonia [136]

Joinvillea plicata, New Caledonia [137]

node), and sister to Ecdeiocoleaceae plus Poaceae. Fossils are known from the Upper Miocene of New Zealand. Genera and species: This family includes only the genus Joinvillea, which has four acceped species, although there are likely to be additional cryptic species in New Caledonia and other Pacific islands. Etymology: Joinvillea is named for François Ferdinand Philippe Louis Marie d’Orléans, Prince de Joinville (1818–1900), son of King Louis Philippe of France. As admiral of the French navy he was entrusted with the task of

Joinvillea gaudichaudiana in fruit, New Caledonia [137]

Georgeantha hexandra, female flower, Mt Benia, Western Australia [138]

returning the remains of Napoleon from Saint Helena to Paris in 1840.

138. ECDEIOCOLEACEAE Kwongan-rush family

Ecdeiocolea monostachya, male flowers, Mt Benia, Western Australia [138]

This is a family of rush-like, perennial, bisexual herbs with creeping, hairy rhizomes. They form clumps and can be locally dominant in the vegetation. Leaves are reduced to sheaths that are split on one side to the base. Sheaths surround green culms or inflorescences, which are round in cross-section. Inflorescences are composed of one to three branches terminated by one to four spikelets. Gl umes (bracts) are mostly fertile, and the gender of flowers alternates one to several times in the spikelet. Flowers are unisexual, flattened, and subtended by a glossy, rigid bractlet. The six undifferentiated petals are in two whorls, the outer two

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POALES

Grass family

Stems are jointed with hollow or (rarely) solid internodes. Stems and branches arise from nodes and are subtended by a leaf sheath. Leaves are arranged alternately along the stem, often in two ranks. Leaf bases are sheathing, surrounding the stem internode, usually open on one side, or partially fused. Blades are usually long and narrow, but can be inrolled and linear to broadly ovate. Veins are parallel, in bamboos sometimes with cross-connecting veinlets. The ligule (an appendage at the sheath-blade junction) is thin and membranaceous, sometimes reduced to a line of hairs or absent. Inflorescences are spikelets, organised in lax or contracted spikes, panicles or racemes, occasionally solitary. Spikelets are composed of bracts arranged along a slender axis (a rachilla), the two lower bracts (glumes) without flowers, subtending one to many florets Florets are composed of two bracts enclosing the flower. The outer bract (lemma) is usually more sturdy and enclosing the more delicate inner bract (palea), which is usually two-keeled. Lemma and glumes often have an elongated bristle (awn; see Fig.38 in Glossary). Flowers are unisexual or bisexual and are composed of (zero to) two (or three to many) highly reduced petals (lodicules), stamens and/or a superior ovary. Stamens are usually three, sometimes one, two, six or many, with a hollow elongate filament and a dangling anther. The ovary is unilocular with (one or) two (rarely three) free or basally united styles with feathery elongate stigmas. Styles and anthers are exserted from the florets. Fruits are normally a dry indehiscent nutlet called a caryopsis, sometimes fleshy in bamboos.

Grasses are a large family of annual and perennial, herbaceous and woody plants (bamboos) with often elongate rhizomes.

Distribution: Poaceae occur globally in every habitat and on every continent, including Antarctica and Greenland. The family is represented in the Antarctic Peninsula and throughout the Arctic and also occurs in deserts, forests, dunes, mangroves, salt swamps, brackish and fresh water, and of course in grasslands. A species of Caryophyllaceae (Colobanthus quitensis) and Antarctic hair grass (Deschampsia antarctica) are the only vascular plants found on the Antarctic mainland.

keeled, the inner four flat. Male flowers have a minute pistillode and four or six stamens with slender free filaments and bilocular, dorsifixed anthers that open by slits. Female flowers have staminodes and a superior ovary topped with two or three bushy styles. The fruit is a trilocular capsule or nut. Plants are wind pollinated and survive fires by resprouting from their rhizomes. Distribution: This family is restricted to nutrient-poor, sandy soils in Western Australia. Phylogeny and evolution: Ecdeiocolea was included in Restionaceae before it was recognised as a separate family; a second genus, Georgeantha, was later added. They were separated on the basis of their culm morphology, embryology and pollen morphology. Molecular studies place these as sister to Poaceae. They are estimated to have evolved c. 73 million years ago (stem node). Genera and species: This is a family of two genera and three species, Ecdeiocolea (2) and Georgeantha (1). Uses: There are no known uses, but the family may be of importance for habitat restoration in kwongan soils of Western Australia. Etymology: Ecdeiocolea is composed of the Greek, εκτός (ektos), outside, δένω (déno), to bind and κολεός (koleos), a sheath or vagina.

139. POACEAE

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Christenhusz, Fay & Chase

Phylogeny and evolution: Grasses evolved during the Late Cretaceous in the forest understory. Bamboos evolved woodiness secondarily and were probably the first group of grasses to diversify and form dominant vegetation. The greatest diversification of true grasses occurred during the Tertiary. The Miocene spread and diversification of tundra, prairie and savanna grasses have been linked to the radiation of large grazing mammals around the same time, which evolved hard enamel on their teeth to cope with the increasing number of silica particles (phytoliths) common in grasses. Grasslands became successful because the growing buds were placed below or close to the ground, allowing them to be grazed or burnt. Grassland grasses started spreading during the Eocene, although some South American grasslands may be older (c. 25 million years old). Evolution of grasslands influenced the biosphere and was partly responsible for the rapid decline in atmospheric CO2 during the Oligocene (c. 33 million years ago), just when the first C4 grasses started appearing. Natural grasslands can be considered a carbon and water sink, with a consequence of global cooling in the longer term. However, the high flammability of dry grasses increased the chances of fire allowing the spread of grasslands even further, and increased temperatures favoured C4 grasses, allowing them to invade C3 -dominated grasslands leading to the development of savannas. Genera and species: This large family includes c. 792 genera and c. 11,000 species in 13 subfamilies, although Aristidoideae, Arundinoideae, Centrothecoideae, Chloridoideae, Danthonioideae, Micrairoideae and Panicoideae form a clade and are therefore treated here in a single subfamily (Panicoideae), instead of accepting seven subfamilies: Anomochloöideae – Anomochloa (1) and Streptochaeta (3); Pharoideae – Leptaspis (3), Pharus (7), Scrotochloa (2) and Suddia (1); Puelioideae – Guaduella (6) and Puelia (5); Ehrhartoideae – Chikusichloa (3), Ehrharta (37), Humbertochloa (2), Hygrochloa (2), Hygroryza (1), Leersia (18), Luziola (11), Maltebrunia (3), Oryza (18), Phyllorachis (1), Por tere sia (1), Potamophila (1),

POALES

MONOCOTS

Panicum miliaceum, leaf base and ligule, Provence, France [139]

Anomochloa marantoidea by W. H. Fitch Gigantochloa atter, new shoot, near from Curtis’s Botanical Magazine vol 88 Caraguatatuba, Brazil [139] (ser. 3, vol. 18): t. 5331 (1862) [139]

Coix lacryma-jobi var. ma-yuen, Royal Botanic Gardens, Kew, UK [139]

Scrotochloa urceolata, Mt Datuk, Malaysia [139]

Zea mais, private garden, Kingston upon Thames, Surrey, UK [139]

Oryza sativa ‘Axios’, Helsinki Botanical Garden, Finland [139]

Leymus arenarius, Isokari, Finland [139]

Aegilops speltoides, Ruissalo Briza major, Sicily, Italy [139] Botanical Garden, Turku, Finland [139]

Triticum aestivum, Ruissalo Botanical Garden, Turku, Finland [139]

Plants of the World

207

POALES Prosphytochloa (1), Rhynchoryza (1), Streptogyna (2), Zizania (4) and Zizaniopsis (5); Bambusoideae – Acidosasa (11), Actinocladum (1), Agnesia (1), Alvimia (3), Ampelocalamus (13), Apoclada (1), Arberella (17), Arthrostylidium (32), Arundinaria (3), Athroostachys (1), Atractantha (6), Aulonemia (45), Bambusa (148), Bashania (4), Bonia (5), Buergersiochloa (1), Cathariostachys (2), Cephalostachyum (12), Chimonobambusa (40), Chimonocalamus (16), Chusquea (156), Colanthelia (7), Cryptochloa (8), Cyrtochloa (7), Davidsea (1), Decaryochloa (1), Dendrocalamus (57), Diandrolyra (3), Didymogonyx (2), Dinochloa (37), Drepanostachyum (10), Ekmanochloa (2), Elytrostachys (2), Eremitis (1), Eremocaulon (4), Fargesia (86), Ferrocalamus (1), Filgueirasia (2), Froesiochloa (1), Gaoligongshania (1), Gelidocalamus (12), Gigantochloa (56), Glaziophyton (1), Greslania (4), Guadua (27), Hickelia (4), Himalayacalamus (9), Hitchcockella (1), Holttumochloa (3), Indocalamus (33), Indosasa (22), Kinabaluchloa (2), Lithachne (4), Maclurochloa (1), Maclurolyra (1), Melocalamus (12), Melocanna (2), Merostachys (46), Mniochloa (1), Myriocladus (13), Nastus (25), Neohouzeaua (8), Neololeba (3), Neomicrocalamus (4), Ochlandra (12), Oligostachyum (17), Olmeca (5), Olyra (24), Oreobambos (1), Otatea (8), Oxytenanthera (1), Parabambusa (1), Pariana (29), Parodiolyra (6), Phuphanochloa (1), Phyllostachys (51), Perrierbambus (2), Pinga (1), Piresia (5), Piresiella (1), Pleioblastus (22), Pseudosasa (21), Pseudostachyum (1), Pseudoxytenanthera (4), Racemobambos (19), Raddia (9), Raddiella (8), Rehia (1), Reitzia (1), Rhipidocladum (16), Sarocalamus (3), Sasa (57), Sasaella (1), Schizostachyum (62), Semiarundinaria (7), Shibataea (7), Sinarundinaria (2), Sinobambusa (14), Sinocalamus (6), Sirochloa (1), Soejatmia (1), Sokinochloa (7), Sphaerobambos (3), Sucrea (3), Temburongia (1), Temochloa (1), Thamnocalamus (4), Thyrsostachys (2), Valiha (2), Vietnamocalamus (1), Vietnamosasa (3), Yersinochloa (1), and Yushania (86); Poöideae – Aegilops (23), Agropyron (13), Agropyropsis (1), Agrostis (218), Agrostopoa (3), Aira (8), Airopsis (1), Alopecurus (39), Ammochloa (3), Ammophila (3), Ampelodesmos (1), Ancistragrostis (1),

208

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MONOCOTS

Aniselytron (2), Anisopogon (1), Anthosachne (9), Anthoxanthum (18), Antinoria (2), Apera (5), Aphanelytrum (2), Arctagrostis (2), Arctophila (1), Arrhenatherum (7), Australopyrum (6), Austrofestuca (3), Avena (21), Beckmannia (2), Brachyelytrum (3), Brachypodium (16), Briza (22), Bromus (154), Brylkinia (1), Calamagrostis (251), Castellia (1), Catabrosa (3), Catapodium (4), Chaetopogon (1), Cinna (4), Coleanthus (1), Colpodium (21), Connorochloa (1), Cornucopiae (2), Corynephorus (5), Crithopsis (1), Cutandia (6), Cyathopus (1), Cynosurus (9), Dactylis (2), Danthoniastrum (4), Dasypyrum (2), Deschampsia (40), Desmazeria (3), Diarrhena (5), Dichelachne (9), Dielsiochloa (1), Dissanthelium (14), Dryopoa (1), Dupontia (1), Dupontiopsis (1), Duthiea (3), Echinaria (1), Echinopogon (7), Elymus (191), Eremopoa (5), Eremopyrum (4), Erianthecium (1), Festuca (620), Gastridium (2), Gaudinia (4), Glyceria (40), Graphephorum (2), Hainardia (1), Helictotrichon (92), Henradia (2), Heteranthelium (1), Hierochloe (33), Holcus (10), Hordeleymus (1), Hordeum (36), Hypseochloa (2), Koeleria (46), Koordersiochloa (2), Lachnagrostis (4), Lagurus (1), Lamarckia (1), Leymus (53), Limnas (3), Limnodea (1), Lindbergella (1), Littledalea (4), Loliolum (1), Lolium (8), Lycochloa (1), Lygeum (1), Megalachne (2), Melica (89), Metcalfia (1), Mibora (2), Micropyropsis (1), Micropyrum (2), Milium (6), Narduroides (1), Nardus (1), Nephelochloa (1), Neuropoa (1), Oreochloa (4), Oreopoa (1), Oryzopsis (8), Parafestuca (1), Parapholis (6), Pentapogon (1), Periballia (3), Peyritschia (7), Phaenosperma (1), Phalaris (17), Phippsia (3), Phleum (16), Pholiurus (1), Piptatherum (33), Pleuropogon (6), Poa (533), Podophorus (1), Polypogon (21), Psammochloa (1), Psathyrostachys (11), Pseudodanthonia (1), Pseudosclerochloa (2), Psilurus (1), Puccinellia (109), Relchela (1), Rhizocephalus (1), Rhombolytrum (2), Rostraria (13), Schizachne (1), Sclerochloa (2), Scolochloa (1), Scribneria (1), Secale (9), Sesleria (31), Simplicia (2), Sinochasea (1), Sphenopholis (6), Sphenopus (2), Stephanachne (2), Stipa (486), Taeniatherum (1), Thinopyrum (11), Torreyochloa (5), Tovarochloa (1), Triniochloa (6), Triplachne (1), Trisetaria (15), Trisetum (84),

Triticum (12), Ventenata (8), Vulpia (26), Vulpiella (1), Wangenheimia (1) and Zingeria (5); Panicoideae – Acostia (1), Acrachne (3), Acritochaete (1), Acroceras (24), Adenochloa (14), Aegopogon (4), Aeluropus (6), Afrotrichloris (2), Agenium (3), Alexfloydia (1), Alloeochaete (6), Allolepis (1), Alloteropsis (5), Altoparadisium (2), Amphicarpum (2), Amphipogon (8), Anadelphia (14), Ancistrachne (4), Andropogon (104), Andropterum (1), Anthaenantia (2), Anthaenantiopsis (4), Anthephora (11), Apluda (1), Apochiton (1), Apocopis (16), Aristida (297), Arthragrostis (4), Arthraxon (22), Arthropogon (5), Arundinella (47), Arundo (4), Arundoclaytonia (1), Asthenochloa (1), Astrebla (4), Austrochloris (1), Austroderia (5), Axonopus (71), Baptorhachis (1), Bewsia (1), Bhidea (3), Blepharidachne (4), Blepharoneuron (2), Bothriochloa (36), Bouteloua (27), Brachiaria (119), Brachychloa (2), Bromuniola (1), Buchlomimus (1), Calamovilfa (5), Calderonella (1), Calyptrochloa (1), Canastra (2), Capeochloa (3), Capillipedium (17), Catalepis (1), Cathestecum (4), Cenchrus (26), Centotheca (3), Centrochloa (1), Centropodia (4), Chaetium (3), Chaetobromus (1), Chaetopoa (2) , Chamaeraphis (1) , Chandrasekharania (1), Chasmantium (5), Chasmopodium (2), Chevalierella (1), Chimaerochloa (1), Chionachne (9), Chionochloa (26), Chloris (59), Chlorocalymma (1), Chondrosum (15), Chrysochloa (4), Chrysopogon (48), Cladoraphis (2), Clausospicula (1), Cleistachne (1), Cleistochloa (3), Cleistogenes (13), Coelachne (11), Coelachyrum (6), Coelorachis (21), Coix (3), Cortaderia (20), Cottea (1), Craspedorhachis (3), Crinipes (2), Crypsis (9), Ctenium (20), Cyclostachya (1), Cymbopogon (53), Cynodon (9), Cyperochloa (1), Cyphochlaena (2), Cyphonanthus (1), Cyrtococcum (14), Dactyloctenium (13), Dallwatsonia (1), Danthonia (26), Danthonidium (1), Danthoniopsis (16), Dasychloa (1), Decaryella (1), Desmostachya (1), Dichaetaria (1), Dichanthium (20), Digitaria (261), Dignathia (5), Diheteropogon (4), Dilophotriche (3), Dimeria (60), Dinebra (5), Diplachne (4), Diplopogon (1), Disakisperma (3), Dissochondrus (1), Distichlis (10), Dregeochloa (2), Eccoptocarpha (1), Echinochloa (34), Echinolaena (7), Ectrosia (14), Eleusine (10), Elionurus (15), Ellisochloa (2),

POALES

MONOCOTS

Elymandra (6), Elytrophorus (2), Enneapogon (24), Enteropogon (17), Entolasia (6), Entoplocamia (1), Eragrostiella (6), Eragrostis (413), Eremochloa (13), Eriachne (48), Eriochloa (34), Eriochrysis (10), Erioneuron (2), Euclasta (2), Eulalia (36), Eulaliopsis (2), Eustachys (15), Exotheca (1), Farrago (1), Fingerhuthia (2), Garnotia (30), Geochloa (3), Germainia (9), Gerritea (1), Gilgiochloa (1), Glyphochloa (11), Gouinia (13), Griffithsochloa (1), Gymnopogon (14), Gynerium (1), Habrochloa (1), Hackelochloa (2), Hakonechloa (1), Halopyrum (1), Harpachne (3), Harpochloa (2), Hemarthria (12), Hemisorghum (2), Heterachne (3), Heteranthoecia (1), Heteropholis (4), Heteropogon (6), Hilaria (10), Hildaea (5), Holcolemma (3), Homolepis (5), Homopholis (1), Homozeugos (6), Hubbardia (2), Hubbardochloa (1), Hydrothauma (1), Hylebates (2), Hymenachne (7), Hyparrhenia (56), Hyperthelia (7), Ichnanthus (31), Imperata (11), Indopoa (1), Isachne (92), Ischaemum (84), Iseilema (25), Ixophorus (1), Jansenella (2), Jouvea (2), Kampochloa (1), Kaokochloa (1), Kellochloa (2), Kerriochloa (1), Lasiacis (15), Lasiurus (1), Lecomtella (1), Leptagrostis (1), Leptocarydion (1), Leptochloa (31), Leptocoryphium (2), Leptothrium (2), Lepturidium (1), Lepturopetium (2), Lepturus (12), Limnopoa (1), Lintonia (2), Lophachme (2), Lophatherum (2), Lopholepis (1), Lophopogon (2), Loudetia (25), Loudetiopsis (11), Louisiella (1), Loxodera (5), Lycurus (3), Manisuris (1), Megaloprotachne (1), Megastachya (2), Melanocenchris (3), Melinis (22), Merxmuellera (7), Mesosetum (26), Michrachne (5), Micraira (15). Microcalamus (1), Microchloa (6), Microstegium (26), Miscanthus (16), Mnesithea (4), Molinia (2), Monachather (1), Monelytrum (1), Monocymbium (3), Monodia (1), Mosdenia (1), Muhlenbergia (164), Munroa (5), Myriostachya (1), Neesiochloa (1), Nematopoa (1), Neobouteloua (2), Neostapfia (1), Neostapfiella (3), Neurachne (7), Neyraudia (4), Notochloe (1), Odyssea (1), Oedochloa (9), Ophiochloa (2), Ophiuros (4), Opizia (2), Oplismenopsis (1), Oplismenus (7), Orcuttia (5), Orinus (6), Oropetium (6), Orthacanthus (1), Orthoclada (2), Oryzidium (1), Otachyrium (8), Ottochloa (3), Oxychloris (1), Oxyrhachis (1), Panicum (446), Pappophorum (7), Paractaenium (1), Parahy-

parrhenia (6), Paraneurachne (1), Paratheria (2), Paspalidium (37), Paspalum (333), Pennisetum (83), Pentameris (80), Pentarrhaphis (3), Pereilema (4), Perotis (13), Phacelurus (9), Phaenanthoecium (1), Pheidochloa (2), Phragmites (4), Piptophyllum (1), Plagiantha (1), Plagiosetum (1), Plinthanthesis (3), Poecilostachys (11), Pogonachne (1), Pogonarthria (4), Pogonatherum (4), Pogonochloa (1), Pogoneura (1), Pohlidium (1), Polevansia (1), Polytoca (2), Polytrias (1), Pommereulla (1), Pringlechloa (1), Prionanthium (1), Psammag rostis (1), Pseudanthistiria (4), Pseudechinolaena (6), Pseudodichanthium (1), Pseudopentameris (4), Pseudoraphis (7), Pseudosorghum (2), Pseudozoysia (1), Psilolemma (1), Pyrrhanthera (1), Ratzeburgia (1), Redfieldia (1), Reimarochloa (3), Reynaudia (1), Rheochloa (1), Rhytachne (13), Richardsiella (1), Rottboellia (6), Rugoloa (3), Rytidosperma (73), Saccharum (35), Sacciolepis (25), Sartidia (5), Saugetia (2), Schaffnerella (1), Schedonnardus (1), Schismus (5), Schizachyrium (61), Schmidtia (2), Schoenefeldia (2), Sclerodactylon (1), Scleropogon (1), Scutachne (2), Sehima (5), Setaria (102), Setariopsis (2), Silentvalleya (2), Snowdenia (4), Soderstromia (1), Sohnsia (1), Sorghastrum (20), Sorghum (28), Spartina (15), Spartochloa (1), Spathia (1), Sphaerocaryum (1), Spheneria (1), Spinifex (4), Spodiopogon (16), Sporobolus (184), Stapfochloa (6), Steinchisma (7), Steirachne (2), Stenotaphrum (7), Stereochlaena (4), Steyermarkochloa (1), Stipagrostis (56), Streptolophus (1), Streptostachys (7), Styppeiochloa (3), Swallenia (1), Symplectrodia (2), Taeniorhachis (1), Tarigidia (2), Tatianyx (1), Tenaxia (8), Tetrachaete (1), Tetrachne (1), Tetrapogon (5), Thaumastochloa (8), Thedachloa (1), Thelepogon (2), Themeda (27), Thrasya (22), Thrasyopsis (2), Thuarea (2), Thyridachne (1), Thyridolepis (3), Thysanolaena (1), Toliara (1), Trachypogon (4), Trachys (2), Tragus (8), Tribolium (16), Trichloris (2), Tricholaena (4), Trichoneura (8), Trichopteryx (5), Tridens (16), Trilobachne (1), Triodia (67), Triplasiella (1), Triplasis (2), Triplopogon (1), Tripogon (42), Tripogonella (3), Tripsacum (14), Triraphis (8), Triscenia (1), Tristachya (20), Tuctoria (3), Uniola (5), Uranthoecium (1), Urelytrum (7), Urochloa (15), Urochondra (1), Vaseyochloa

(1), Veldkampia (1), Vietnamochloa (1), Viguierella (1), Vossia (1), Whiteochloa (6), Willkommia (4), Xerochloa (3), Yakirra (7), Yvesia (1), Zaqiqah (1), Zea (6), Zenkeria (5), Zeugites (10), Zonotriche (3), Zoysia (8) and Zygochloa (1). Uses: Poaceae are probably the economically most important family as they are used for food, spice, perfume, building material, biofuel, animal fodder, decoration and sports and recreation. Cereal grains were important in the development of modern human culture. Cultivation of cereal crops (derived from Ceres, the Greek goddess of harvest), allowed humans to become less nomadic and more settled, building houses and protecting their valuable crops against wild animals and human invaders. Grasses are responsible for the formation of the first cities because they permitted the storage of food throughout adverse periods and an excess of food to develop, which permitted professions to arise that were not concerned with food. Wheat (Triticum) provides a fifth of the calories eaten by humans today. It is the primary cereal of temperate regions. Einkorn wheat (T. monococcum) began to be domesticated c. 9000 BC in southwestern Turkey. Even though it was one of the first grains to be cultivated, it is currently rarely grown, usually only as animal food or to make bulgur (cracked wheat). It has a different type of gluten than conventional wheat, which may be of use for people with wheat gluten intolerance. Emmer wheat (T. dicoccum) was first domesticated 10,000 years ago in the Levant and Iran during the Neolithic period, possibly through natural hybridisation of T. dicoccoides and Aegilops speltoides. Emmer is still used to make bread in Italy and Switzerland, and it is used whole in soups in Tuscany. Because of the high fibre content of emmer wheat, it has become popular in the health food industry. Durum wheat (Triticum durum) is also tetraploid and a selection of emmer wheat from Central Europe and the Middle East, which first appeared in cultivation around 7000 BC. It has a high protein content, making it suitable for couscous, dry pasta, gruels, noodles, papadums, pizza, puddings, semolina,

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POALES soups etc. Wild emmer or durum wheat was hybridised with another wild grass (Aegilops tauschii) to produce hexaploid spelt (T. spelta) and modern bread wheat (T. aestivum). Rye (Secale cereale) is known from Neolithic sites in Turkey, but was not domesticated before the Bronze Age in Central Europe (c. 1800–1500 BC). It has been suggested that early wheat fields included a small amount of rye, and that when a wheat crop failed, rye survived and was thus selected. It has been considered inferior to wheat and barley, but because of its greater climatic tolerance, it became the staple crop in the colder climates of Europe. Rye is used for flour, bread, beer, crackers, pumpernickel, vodka, whisky and animal fodder. It can also be cooked whole, and it is low in glutenin, making it suitable for some people with gluten intolerance. Triticale (×Triticosecale) is a hybrid between wheat and rye, which was developed in the 19th century in Scotland and Sweden. It combines the yield and quality of wheat with the disease resistance and environmental tolerance of rye. It has a higher protein and a lower gluten content. It is used for bread and other baked goods and animal feed. Barley (Hordeum vulgare) is most important for the production of beer, whisky and other fermented distilled beverages and animal food. Wild barley (H. vulgare subsp. spontaneum) was first harvested c. 2300 BC in the Middle East, North Africa and Tibet. It was one of the first domesticated grains and is known from a Palaeolithic site in Israel, dated 8500 BC. It has been argued that the availability of barley, along with other domesticatable animals and crops such as wheat and rye, allowed Eurasian civilisations to thrive and conquer other areas. Barley was even used as currency in early times. Barley bread was considered a peasant food and has been largely replaced by potatoes. Asian rice (Oryza sativa) is the most widely consumed staple food, especially in Asia and Latin America. Rice flour has a great number of uses. Rice was independently domesticated in two places, Asian rice (O. sativa) in China c. 10,000 years ago and African rice (O. glaberrima) c. 2,500 years

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ago in the Niger valley of Mali. African rice has become more popular in recent times because it is tolerant of fluctuations in water levels, changes in temperature, pollution and herbivory. However, the grains are more brittle than those of Asian rice, making them difficult to process industrially, and in addition African rice has a lower yield. A hybrid between the two is now marketed as ‘nerica’ (= New Rice for Africa), with great potential. Domestication of maize or corn (Zea mays subsp. mays) occurred 8,700 years ago in seasonal tropical forests in southwestern Mexico, perhaps in the Balsas Valley. It is a selected form of teosinte (Z. mays subsp. mexicana). It is currently the most important cereal crop next to wheat and rice and a staple food in the Americas and Africa. It is the basis for industrial alcohol, babycorn, beer, bourbon, cooking oil, cornflakes, tortillas, tamales, cornflour and polenta. Sugar-rich varieties (sweet corn) are usually grown for human consumption, whereas starchy field corn is used for silage to feed livestock, making it also an important crop for the dairy industry. Even though native to the Fertile Crescent, oats (Avena sativa) were first domesticated in the Bronze Age of Europe. Oats are usually considered a secondary crop, although, like rye, they are highly valued in the colder climates of Europe. Oats are usually rolled or crushed into oatmeal or ground into flour. Oatmeal is eaten as a porridge or made into cookies, digestive biscuits and other baked goods. They also are a major component of breakfast cereals such as muesli and granola. Oats are valued for their capacity to lower cholesterol. Oat cakes are especially popular in Scotland. Oat is also used to feed horses when they need an extra energy boost. Winter oats are grown as a green manure. Samuel Johnson, in his dictionary published in the 18th century, notoriously defined oats as ‘a grain, which in England is generally given to horses, but in Scotland supports the people.’ Of southern African origin, sorghum (mainly Sorghum bicolor, but other species are also used) has been a staple food for millions of people in Africa, Asia and Latin America. It can be grown without additional fertilisers and is the fifth most important cereal crop.

Even though it is used for human consumption in impoverished countries, its main use is for poultry and cattle feed. Millet is a group of small-seeded grasses used as cereal crops and animal fodder. They are important local crops, especially in developing countries because of their short growing season and ability to cope with drought and high temperatures. They encompass grasses from a variety of genera, for example: Guinea millet (Brachiaria deflexa), Job’s tears (Coix lacryma-jobi), Japanese millet (Echinochloa esculenta), Indian barnyard millet (E. frumentacea), burgu millet (E. stagnina), finger millet (Eleusine coracana), tef (Eragrostis tef ), proso millet (Panicum miliaceum), kodo millet (Paspalum scrobiculatum), pearl millet (Pennisetum glaucum), foxtail millet (Setaria italica) and browntop millet (Urochloa ramosa). Fonio is a type of millet that includes species of wild or domesticated grains from the genus Digitaria. White fonio or hungry rice (Digitaria exilis) is a minor crop in West Africa. Even though it has small seeds, it is locally an important food crop because it can mature from seed in less than eight weeks, making it useful in areas prone to drought. Black fonio (D. iburua) is a similar crop, and raishan (D. cruciata) is a minor cereal in India. One of the main crops, if not globally dominant, is sugarcane (mainly Saccharum officinarum, but also S. edule and S. barberi are grown). Originally from tropical Asia, it was first domesticated in New Guinea c. 6000 BC but is now cultivated throughout the tropics. The first production of crystalline sugar occurred in northern India. Sugar was one of the major reasons for European colonisation of the tropics, and the rising demand for sugar in Europe fuelled further land grabs and conversion of tropical forests into cane fields, mainly with the help of slave labour. Sugar formed one of the arms in the triangle trade: Caribbean molasses was shipped to Europe and New England where it was sold for rum or sugar; these profits were used to buy manufactured goods that were shipped to West Africa, where they were exchanged for slaves, who were then transported to the Caribbean to work on the plantations. Sugarcane — and the associated

POALES

MONOCOTS

human addiction to sugar — is thus responsible for the displacement of millions of people and the reshaping of the global map. Currently 80% of all raw sugar is produced by sugar cane. Apart from sugar, sugarcane is used for biofuels (bagasse and ethanol production), molasses or other syrups like falernum and alcoholic drinks like cachaça and rum. Fresh canes are also eaten as a treat. Bamboos are extremely useful in many regards. Young bamboo shoots, common in Asian cuisine, are mainly from Bambusa vulgaris and Phyllostachys edulis, but other species can be eaten. They can be peeled and cooked, stir-fried, pickled or fermented. Sap of bamboos can be tapped and fermented into a sweet drink called ulanzi. Tough stems of bamboos are hollow, flexible and sturdy, making them good for construction and scaffolding poles. They are also used for musical instruments (e.g. panpipes and flutes), furniture, basketry, weaponry, fishing rods, bicycles, fireworks, water desalination, and a great range of everyday utensils. Good quality bamboo is stronger than steel. Bamboo can also be compressed and glued into woodblocks that can be turned into various shapes. Bamboo leaves are sometimes used to wrap food. Bamboo fabrics are made through a long chemical process because bamboo fibres are naturally short. There is a long paper-making tradition in East Asia using bamboo (mostly from Bambusa blumeana and Dendrocalamus asper), and it is a good alternative to wood. Because of the longevity and usefulness of bamboos, several Asian cultures believe that humanity emerged from a bamboo shoot. Another useful grass is the common reed (Phragmites australis), which was traditionally used for thatching of roofs and is still harvested and valued for that purpose, especially in western Europe. Although considered a famine food, young stems of reeds can be dried, ground into powder, made into a dough and roasted. Common reed is also used for basketry, mats, quills and spears. Currently research is being done into its use in compressed form as a building material, insulation for houses and a substitute for traditional wood-based paper. Phragmites

is also employed for natural water cleaning treatment. The plants extract nutrients from the water, which is then mown, composted and used in agriculture. Several grass species are used for perfume or as a spice. Most famous is lemongrass (Cymbopogon citratus and C. flexuosus), which is native to tropical Asia and is often used in local cuisines. It has a citrusy flavour and is added to soups, curries and teas. It is also commonly grown and naturalised in other tropical countries. Lemongrass oil has antifungal properties and is used as a pesticide on plants. Citronella grass (C. nardus) is similar but is usually grown for the insect repellent properties of the oil, commonly used to scent candles. Vetiver (Chrysopogon zizanioides) is mostly cultivated for its fragrant essential oils, present in almost all modern perfumes. It is also sometimes used to flavour foods and drinks. As a tropical cash crop, it also finds use in the prevention of erosion in depleted lands due to its deep roots. Since the early 1980s, Miscanthus giganteus has been used for the production × of biofuel in Europe due to its high biomass, low mineral content and rapid growth. In a single growing season it can reach heights of 3.5 m, which is why it sometimes is called ‘elephant grass’. It is made into ethanol. Rosary beads and necklaces can be made from the hard-shelled seeds of Job’s tears (Coix lacryma-jobi). A cultivar with soft shells called ‘ma yuen’ is used as a cereal in China. Several species are important plants for stabilising soils. In the tropics vetiver, Chrysopogon zizaniodes, is often used, but in temperate zones lyme grass, Leymus arenarius, can be grown for that purpose. Marram or beach grass (Ammophila arenaria) is used to stabilise sand dunes in the Netherlands and elsewhere. Grasses are a major crop for the dairy, meat and wool industries. Grazing animals (predominantly sheep and cows) depend on sufficient amounts of grasses to thrive and produce wool, milk and meat. Cattle and sheep and their associated meadows have changed the landscapes of the world. Another common use of grasses is the lawn. Lawns are composed of one or

a mixture of selected grasses that are kept short. Lawns are a common feature of gardens and parks and are used for aesthetics and recreation. In temperate regions, lawns are usually composed of a mixture of species of Agrostis, Festuca, Lolium and Poa, but in warmer regions Bermuda grass (Cynodon dactylon) or species of Axonopus, Eremochloa, Paspalum, Stenotaphrum and Zoysia are used. Grasses can also cause major problems. Pollen allergy (hayfever) is not unique to grasses (other wind-pollinated species such as pines and birches can also cause it), but grass pollen is a major allergen for numerous people worldwide. Darnel (Lolium temulentum) was a serious weed of cereal fields. Although not poisonous itself, contamination of cereal harvests with seeds of this species infected with the ergot fungus led to many cases of human poisoning. Some grasses become invasive outside their native range and can crowd out native vegetation, spark increased bushfires and prevent forests from regenerating. Several grass species are in cultivation as ornamental plants, especially species of bamboo and Andropogon, Bouteloa, Cortaderia, Eragrostis, Imperata, Melica, Miscanthus, Molinia, Pennisetum, Phalaris, Sasa, and Stipa, to name a few. Etymology: Poa is derived from the ancient Greek word πόα (poa, poie or poia), a pasture grass. Trivia: If you want to see a plant grow, your best bet is to watch a bamboo shoot. It has reported growth rates of over two metres per day. Bamboos flower infrequently, usually at intervals of 65 to 130 years. The longest period between flowering recorded is of Phyllostachys bambusoides, in which all plants of the species worldwide (the species is commonly planted as an ornamental) flowered at about the same time. Production of masses of seeds allows animals to feed on it without harming the population, especially due to the unpredictability of the occurrences of flowering. Finally, bamboo comprises about 99% of the diet of the giant panda, an emblematic species of conservation.

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CERATOPHYLLALES

UNPLACED

CERATOPHYLLALES The placement of Ceratophyllales is not yet certain. Most frequently in recent gene studies, they are sister to the eudicots. Fossils intermediate between Ceratophyllales and Chloranthales are known, making the placement of both orders problematic from a morphological persective.

140. CERATOPHYLLACEAE Hornwort family

212

occur on the same plant and are submersed and subtended by a whorl of eight to 15 linear bracts, which are sometimes mistaken for a perianth, which they lack. Male flowers have three to 50 stamens with a prolonged connective and sessile anthers that open by slits. Female flowers have a single, simple pistil with a superior, unilocular ovary and an elongate style. The fruits are indehiscent, spiny nuts with a single seed and a persistent style. Flowers are pollinated by water, and seeds are dispersed by water currents.

These submersed aquatic, perennial herbs lack rhizomes or roots, but may be anchored in the substrate by specialised branchlets. Three to ten whorled leaves arise from branched, hollow stems and lack stipules and petioles. Leaf blades are simple or one to four times forked, and the dissections have finely toothed margins. The unisexual flowers are formed solitarily in the leaf axils. Staminate and pistillate flowers

Phylogeny and evolution: Recent molecular research has indicated that Ceratophyllaceae are an ancient lineage with no close extant relatives. It has been hypothesised that it is a relict lineage of ancient angiosperms

Ceratophyllum demersum, Turbaco, Bolívar, Colombia [140]

Ceratophyllum demersum, fruit, private garden, Kingston upon Thames, Surrey, UK [140]

Christenhusz, Fay & Chase

Distribution: The family occurs in fresh water bodies worldwide.

diverging early from a lineage leading to modern eudicots. There is fossil evidence that the family was already present during the Cretaceous, c. 115 million years ago. Ceratophyllaceae are placed in an order of their own and are not generally included in eudicots. Genera and species: The single genus in this family, Ceratophyllum, consists of four species. Uses: Species are sometimes grown in aquaria or ponds as oxygenating plants and breeding places for fish. They are sometimes invasive and can be considered a nuisance in waterways and hydroelectric plants. Etymology: Ceratophyllum is derived from the Greek κέρας (keras), a horn, and φύλλων ( fyllon), a leaf. Ceratophyllum demersum in flower, private garden, Kingston upon Thames, Surrey, UK [140]

EUDICOTS The recognition of the existence of this group was one of the novel aspects of the advent of DNA phylogenetics in the angiosperms. Tricolpate pollen (and its derivatives), the most obvious synapomorphy of the eudicots, had been known for a long time, but no one had ever attached a great deal of significance to this trait. Many palynologists had stated that development of this type of pollen was significantly different from that of monosulcate pollen, the other major pollen type, but the groups in which tricolpate pollen exists are so diverse in terms of their morphology that no taxonomist seriously considered that these groups constituted a clade. Even with just the plastid gene rbcL (Chase et al. 1993), this clade was clear. The term eudicot was first used in a different context by Doyle & Hotton (1991), but it was picked up quickly and used in the first broad analyses of DNA data for this clade. It has been proposed that it is not entirely an appropriate term (magnoliids are “true” dicots as well), and the term “tricolpates” has been proposed in its place ( Judd & Olmstead 2004); despite this, the term “eudicots” continues to be widely used. What is clear is that “dicot” is no longer a useful reference, in that it simply now refers to every angiosperm that is not a monocot; these do not form a clade and they include the ANA grade, magnoliids plus Chloranthaceae, Ceratophyllaceae and eudicots. Recognition of eudicots as a clade has given researchers using molecular clock approaches one of their clearest calibration points. Appearance of tricolpate pollen in the fossil record was

abrupt, but there can be no doubt when it is detected that one is then presented with a member of the eudicot clade. Tricolpate pollen first appeared in the fossil record c. 125 million years ago, but when we use a number of other seemingly clear calibration points and estimates of the age of the eudicots, the resulting age estimate for this clade is usually slightly older. It does make sense that if there is abundant tricolpate pollen 125 million years ago, then the clade must have originated before this point, although this pollen evidence is yet to be found. Other characters for the eudicots include usually four- or five-merous f lowers and trends towards fusion of or a reduction in f lower parts. However, the clade is diverse and, in particular, the first several successive sister groups to the rest, such as the families of Ranunculales, have often been included among the magnoliids (Cronquist 1981) due to their numerous stamens and irregular number of perianth parts. Other families, such as Trochodendraceae, have vesselless wood, which must be due to secondary loss, and were often placed among magnoliids for that reason, even though other morphological characters did not coincide with this. Before the use of DNA data such families often presented problems because of their preponderant seemingly primitive traits, even though they had tricoplate pollen. DNA analyses have played a major role in readjusting the thinking of plant systematists in major and significant ways. Many botanists forget that this major shift took place only

since the mid 1990s; it now seems so much a part of our basic thinking that we forget how recently this revolution in thinking took place. Etymology: Eudicot is derived from the Greek words eu (ευ), well or good, dio (δύο), two, and kotylidon (κοτυληδών), seedlobe. General references: Byng JW. 2014. The f lowering plants handbook, a guide to families and genera of the world. Plant Gateway, Hertford. Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall MR, Price RA, Hills HG, Qiu YL, Kron KA, Rettig JH, Conti E, Palmer JD, Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD, Hedrén M, Gaut BS, Jansen RK, Kim KJ, Wimpee CF, Smith JF, Furnier GR, Strauss SH, Xiang QY, Plunkett GM, Soltis PS, Swensen SM, Williams SE, Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn GH Jr, Graham SW, Barrett SCH, Dayanandan S, Albert VA. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Annals of the Missouri Botanical Garden 80: 528–580. Cronquist A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York. Doyle JA, Hotton CL. 1991. Diversification of early angiosperm pollen in a cladistic context, pp. 169–195, in: Blackmore S, Barnes SH (eds.) Pollen and spores. Patterns of diversification. Clarendon, Oxford. Judd WS, Olmstead RG. 2004. A survey of tricolpate (eudicot) phylogenetic relationships. American Journal of Botany 91: 1627–1644.

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RANUNCULALES Families 141 to 147 form the order Ranunculales. This order (stem node) is between 140 and 113 million years old. A fossil of 122.6 million years old confirms this age. Most of these families diverged from each other c. 100 million years ago. Eupteleaceae are sister to the other families. Wind pollination and floral nectar spurs have evolved several times independently in Ranunculales. The origin of petals in this order has been discussed extensively. Often the inner whorl of petals of especially Ranunculaceae and Berberidaceae are nectar secreting and derived from stamens, whereas the outer, often sepal-like whorl are considered tepals; where both whorls consist of petallike tepals, gene expression is not well distinguished, whereas in species that have a differentiation between a sepal-like and petal-like whorl the gene expression is similar to that in other eudicots, leading to a discrete calyx and corolla. Because of this in the descriptions below we sometimes refer to sepals and petals when these two whorls are readily distinguishable as such, even though developmentally they may be tepals and stamens.

141. EUPTELEACEAE Asian-elm family

axils of bracts at the base of a leafy shoot. The bisexual flowers lack a perianth; the six to 19 stamens occur in more or less a single whorl, and the eight to 31 free carpels are stalked and also in one whorl. The fruit is composed of several stalked samaras. Distribution: This family occurs in temperate East Asia, northern India, central China and southern Japan.

These deciduous trees and shrubs have spirally arranged leaves without stipules. The leaves are petiolate and undivided, but with a toothed margin and pinnate venation. Inflorescences emerge in spring before the leaves and are composed of six to 12 flowers clustered in

Phylogeny and evolution: Euptelea is sister to the rest of Ranunculales and only shares with other members of the order a few cryptic characters. Previously considered by some authors to be a member of Hamamelidales or to be associated with Trochodendraceae or Cercidiphyllaceae, the simple, wind-pollinated

Euptelea pleiosperma, Royal Botanic Gardens, Kew, UK [141]

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flowers of Euptelea, with no sepals or petals, made systematic placement difficult before the advent of analyses of DNA sequences. This is an old lineage, with a maximum estimated age of c. 135 million years. Genera and species: The single genus in this family is Euptelea, which has two species: E. pleiosperma and E. polyandra. Uses: Both species are sometimes grown as ornamental trees, particularly for their attractive glossy, bright green foliage, which emerges purple and turns red and yellow in autumn. Etymology: Euptelea is Greek, from ευ (eu), good, and πτελέα (ptelea), an elm.

Euptelea polyandra in fruit, Royal Botanic Gardens, Kew, UK [141]

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Pteridophyllum racemosum, Royal Horticultural Soctiety Alpine show, London, UK [142]

Corydalis nobilis, Turku, Finland [142]

Capnoides sempervirens, Helsinki Botanical Garden, Finland [142]

Hunnemannia fumariifolia, private garden, Kingston upon Thames, Surrey, UK [142]

Papaver commutatum, Ani, Turkey [142]

142. PAPAVERACEAE

often present. The stems are herbaceous and usually weak and erect, sometimes vining or almost woody. From latex cells a white, yellow or orange, sometimes watery sap exudes when damaged. Leaves are spirally arranged, rarely appearing opposite or whorled, usually along the stems or, in Hypecoum and Pteridophyllum, in basal rosettes, without stipules. The petioles are often somewhat clasping at the base, and the blades are simple but some lobed to pinnately or ternately divided, usually pinnately, rarely palmately veined. Inflorescences or single flowers are borne at the tip of stems or in axillary bracteate cymes, umbels or racemes; some are scapose thyrses (Pteridophyllum). In Papaveroideae, flower buds are often nodding. Flowers are bisexual, actinomorphic

Poppy family

This family of mostly herbaceous perennials and annuals also includes a few woody genera. Hunnemannia, Dendromecon and Romneya are usually shrubs, and Bocconia species can even be trees. Perennating rhizomes or tubers are

Bocconia arborescens, San Francisco Botanical Garden, USA [142]

or zygomorphic, and the perianth divided into two whorls. The two (or three) sepal-like tepals which are sometimes fused into a cap, soon fall off, often upon opening of the flower. The (two to) four or six petal-like tepals can be all similar or sometimes one or two of them can be extended into a sac or spur, rarely the petals absent (Bocconia, Macleaya) or more (six to 12 in Sanguinaria). Stamens can be two, four, six, eight or numerous, the anthers basifixed, opening by slits, the filaments free or fused, with or without nectaries. The superior ovary is composed of two to 20 fused carpels, topped with a stalked or sessile, variably shaped stigma, which is one of the best characters for species identification. The fruits are usually capsules, sometimes nuts or loments.

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RANUNCULALES Distribution: Papaveraceae are widespread and most diverse in the temperate Northern Hemisphere, but a few species occur in Central and South America and southern Africa. Papaver radicastum can be found at 83ºN latitude, and it and Salix arctica (Salicaceae) are the most northerly distributed vascular plants. Phylogeny and evolution: In some earlier classifications the families Fumariaceae and Pteridophyllaceae were treated separately from Papaveraceae and the three together comprised Papaverales. This order is embedded in Ranunculales, and all members of the former order Papaverales are now treated as a single family with two subfamilies. European Meconopsis cambrica is the type of the genus Meconopsis, but genetically this species falls within Papaver, rather than with Asian Meconopsis species. This and some other findings may result in further taxonomic changes in the circumscription of some genera. The anomalous genus Pteridophyllum is surprisingly most closely related to Hypecoum. The crown group diverged c. 110 million years ago. Genera and species: This family has c. 45 genera and about 775 species divided into two subfamilies: Fumarioideae – Adlumia (1), Capnoides (1), Ceratocapnos (3), Corydalis (c. 400), Cryptocapnos (1), Cysticapnos (4), Dactylicapnos (18), Dicentra (c. 12), Discocapnos (1), Ehrendorferia (2), Fumaria (c. 50), Fumariola (1), Hypecoum (33), Ichtyoselmis (1), Lamprocapnos (1), Plat ycapnos (3), Pseudofumaria (2), Pteridophyllum (1), Rupicapnos (7), Sarcocapnos (4) and Trigonocapnos (1). Papaveroideae – Arctomecon (3), Argemone (23), Bocconia (10), Canbya (2), Cathcartia (6), Chelidonium (1), Dendromecon (2), Dicranostigma (5), Eomecon (1), Eschscholzia (c. 10), Glaucium (c. 25), Hesperomecon (1), Hunnemannia (2), Hylomecon (3), Macleaya (2), Meconella (3), Meconopsis (c. 40), Papaver (c. 80), Platystemon (1), Roemeria (3), Romneya (2), Sanguinaria (1), Stylomecon (1) and Stylophorum (2).

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Uses: Poppy seeds (mostly Papaver somniferum and P. rhoeas) are commonly used as a condiment. A paste of ground poppy seeds and sugar is used in Central and Eastern European patisserie. Poppy seeds are a traditional ingredient of breads, bagels and other baked goods. Opiate drugs derived from the opium poppy (Papaver somniferum), including codeine, heroin and morphine, are widely used in medicine as analgesics. Opium poppies were cultivated in Mesopotamia before written history, and images of them appear on 6,000-year-old Sumerian artefacts. Papaver somniferum was associated with several ancient deities, including Demeter, goddess of fertility and agriculture in Greek mythology. Native to Anatolia, opium gave its name to the Turkish city of Afyonkarahisar (meaning ‘opium black castle’), and until the 1960s that region of Turkey was a major producer of raw opium. Opium became infamous due to the Opium Wars between China and Britain and France in the 19th century, when China wanted to prevent import of the drug into China by western traders. Currently there is a huge illicit trade in opiates, fuelling wars and social unrest in Central Asia, and it has a major social cost resulting from problems related to addiction. Poppy seeds used in baking contain small amounts of opiates. The yellow juice of greater celandine (Chelidonium majus) has been used as a wart remedy. Commercially, bloodroot (Sanguinaria canadensis) is used in products for removing moles and in dental hygiene. The root of Sanguinaria yields a red dye that is popular in Native American handicrafts. Many familiar garden plants belong to the family; e.g. Corydalis (birdin-a-bush), Dicentra (western bleeding hearts), Lamprocapnos (eastern bleeding heart), Pseudofumaria (yellow corydalis), Eschscholzia (Californian poppy), Macleaya (plume-poppy), Meconopsis (blue poppy), Papaver (poppy) and Romneya (matilija or tree poppy). Etymology: Papaver is the Latin name for a poppy or poppy seed, referring to pappa, milk, i.e. the latex exuded from the plant.

143. CIRCAEASTERACEAE Witch’s-star family

This is a family of annual and perennial herbs. In the annual Circaeaster the cotyledons are persistent in mature plants, and in perennial Kingdonia a slender rhizome is formed. Leaves are either simple, rosulate and borne on an elongated hypocotyl (Circaeaster) or single, long-petiolate and palmately arranged (Kingdonia). The veins are flabellate-pinnate, (dichotomously) forking. The bisexual flowers are solitary or in fascicles in leaf axils. The two, three, (four), five or six (rarely seven) tepals are Circaeaster agrestis, China (BX) [143]

Kingdonia uniflora, near Emei Shan, Sichuan, China (CD) [143]

RANUNCULALES

EUDICOTS

persistent or not. Stamens are one, two, or (three to) five to eight, and staminodes, if present, can be eight to 13. The anthers have two pollen sacs. Ovaries are superior and composed of one to nine free carpels, with or without a style. The fruits are indehiscent nutlets. Distribution: These enigmatic plants are found in patchy populations in montane habitats from India (Kumaun) through Nepal to western and central China. Phylogeny and evolution: The two genera differ in many characters, and some taxonomists therefore prefer to recognise each in a separate family. However, molecular results show a close relationship between the two, and the species share characters of venation and anatomy. The genera diverged c. 80 million years ago. Circaeasteraceae are most closely related to Lardizabalaceae. Genera and species: This family includes two genera each with a single species: Circaeaster agrestis and Kingdonia uniflora. Etymology: Circaeaster is derived from the Greek goddess of magic, Κίρκη (Circe), known for her vast knowledge of drugs, herbs and witchcraft, and Latin aster, a star.

144. LARDIZABALACEAE Zabala-fruit family

This family consists mostly of lianas, rarely shrubs. The alternate leaves are palmately compound or (in the case of the shrub Decaisnea) pinnate. The three to many leaflets are stalked. The inflorescences are essentially racemose, but sometimes they appear as cymes or umbels or have sessile solitary flowers. The unisexual flowers are usually combined in mixed inflorescences (with the female flowers

Decaisnea fargesii in fruit, Royal Botanic Gardens, Kew, UK [144]

Akebia longeracemosa, National Botanic Gardens of Ireland, Glasnevin [144]

at the base of the inflorescence), but in South American genera (Boquila, Lardizabala) the male and female flowers occur on different plants. Flowers have three or six petal-like sepals. The six petals are absent or strongly reduced and gland-like, petaloid in Lardizabala. The male flowers are smaller than the female flowers and have six stamens and rudimentary carpels. Female flowers have superior free carpels, often three, but can be irregular in number. The fruits are one to several round or sausage-like berries with a leathery skin and sweet slimy flesh in which the black or brown shiny seeds are embedded. Distribution: This disjunctly distributed family is found in temperate East Asia (Kashmir to northern Vietnam, China, Taiwan, Japan) and southern South America (Chile, Juan Fernández Islands). Phylogeny and evolution: Lardizabalaceae were previously thought to be related to or part of Berberidaceae or Menispermaceae. They are closest to the latter together with Circaeasteraceae. Their relationship to Berberidaceae is more distant. Sargentodoxa is sister to the rest of Lardizabalaceae and sometimes segregated as Sargentodoxaceae, but it has many similarities to Lardizabalaceae and is thus maintained in that family here. The current distribution of Lardizabalaceae is widely disjunct, fossils being known from Europe and North America, but the relationship between the Chilean and East

Stauntonia libera (KWJ12218), Crûg Farm Plants, Wales, UK [144]

Asian species is not old enough to warrant a vicariant hypothesis. The two genera in South America are most likely derived from a more recent long-distance dispersal event. Genera and species: Lardizabalaceae include seven genera and c. 40 species: Akebia (6), Boquila (1), Decaisnea (2), Lardizabala (1), Sargentodoxa (1), Sinofranchetia (1) and Stauntonia (c. 28). Uses: In its native Japan, young shoots of ‘akebi’ (Akebia quinata) are used in vegetable tempura. It is commonly grown in gardens in the temperate zones for its exotic appearance and fragrant flowers and naturalises in North America and New Zealand. Some other species, particularly of the genera Akebia, Decaisnea and Stauntonia, are grown in more specialist horticultural collections. The sweet fruit pulp of all species is edible, but especially Lardizabala (zabala fruit) and Akebia are found in local markets in Chile and China, respectively. Fruits of other species are locally consumed, aiding dispersal when seeds are spat out along the path. Etymology: Lardizabala is named for the philosopher Miguel de Lardizábal y Uribe (1744–1824) from Tlaxcala, Mexico, who studied geology and history at the Botanical Garden of Madrid, Spain. He later became a politician and as Minister to the Indies in 1814 he tried to stop aspirations of independence in the Spanish colonies of America, leading to his imprisonment in 1815.

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145. MENISPERMACEAE Moonseed family

EUDICOTS

to male flowers, but sometimes dimorphic and then female flowers with fewer petals, with or without staminodes. The ovary is composed of one, three, six or more (to 21) free or rarely fused carpels. Fruits are drupes, the seeds often with a curved embryo (hence the common name moonseed). Distribution: These occur mostly in Old and New World tropical lowland rainforests, but some are subtropical or warm temperate in North America and East Asia.

These are predominantly lianas, but some are herbs, shrubs and small trees. They usually grow terrestrially (rarely epiphytically), and stems are vining or erect, usually unarmed, sometimes with spines (Antizoma). Phylloclades are found only in Cocculus balfourii. Leaves have petioles but lack stipules, and the petioles are thickened at both ends, allowing leaves to turn towards the light. Blades are usually simple, sometimes lobed, often peltate at the base, with palmate or pinnate venation. Inflorescences are borne in axils or terminate leafless twigs and are usually many-flowered racemes or panicles, sometimes cymes or flowers solitary. The unisexual (rarely bisexual), actinomorphic or zygomorphic flowers have three to 12 free or fused sepals that are in whorls of three. Petals are one to six, sometimes absent, and free or fused. Male flowers have free or fused stamens that can be three to six, rarely one or two or up to 40. Female flowers are usually similar

Phylogeny and evolution: The family is estimated to have diverged from its sister clade c. 124 million years ago, the two subfamilies diverging during the Late Cretaceous, with Laurasian diversification and distribution c. 70–60 million years ago. Palaeocene fossils are known from South America. Genera and species: Menispermaceae comprise 71 genera and c. 440 species: Abuta (32), Albertisia (17), Anamirta (1), Anisocycla (3), Anomospermum (6), Antizoma (2), Arcangelista (2), Aspidocarya (1), Beirnaertia (1), Borismene (1), Burasaia (5), Calycocarpum (1), Carronia (4), Caryomene (4), Chasmanthera (2), Chlaenandra (1), Chondrodendron (3), Cissampelos (20), Cocculus (8), Coscinium (2), Curarea (4), Cyclea (29), Dialytheca (1), Dioscoreophyllum (3), Diploclisia (2), Disciphania (c. 25), Echinostephia (1), Elephantomene (1), Eleutharrhena (1), Fibraurea

(2), Haematocarpus (2), Hyperbaena (19), Hypserpa (8), Jateorhiza (2), Kolobopetalum (4), Legnephora (5), Leptoterantha (1), Limacia (3), Limaciopsis (1), Macrococculus (1), Menispermum (2), Odontocarya (31), Orthogynium (1), Orthomene (4), Pachygone (9), Parabaena (6), Parapachygone (1), Penianthus (4), Pericampylus (3), Platytinospora (1), Pleogyne (1), Pycnarrhena (9), Rhaptonema (4), Rhigiocarya (3), Sarcolophium (1), Sarcopetalum (1), Sciadotenia (19), Sinomenium (1), Sphenocentrum (1), Spirospermum (1), Stephania (36), Strychnopsis (1), Synclisia (1), Syntriandrium (1), Syrrheonema (3), Telitoxicum (6), Tiliacora (22), Tinomiscium (1), Tinospora (33), Triclisia (10) and Ungulipetalum (1). Uses: The South and Central American genus Chondrodendron is the source of the muscle relaxant and arrow poison curare. Species of Abuta, Curarea, Sciadotenia and Telitoxicum in South America and ipos (Anamirta, Cocculus, Coscinium and Tinospora) in Asia have also been used by local tribes as arrow or fish poison. Picrotoxin, used as a parasiticide in modern medicine, is derived from the Indomalaysian Anamirta. A yellow dye derived from the stems of Fibraurea is used in China and India. Etymology: Menispermum is derived from the Ancient Greek μενε (mene), the moon, and σπέρμα (sperma), seed, in reference to the curved seeds.

Stephania brachyandra, female flowers, Kunming Botanical Garden, China [145]

Cocculus balfourii, Royal Botanic Gardens, Kew, UK [145]

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Chondodendron tomentosum, New York Botanical Garden, USA [145]

Menispermum canadense, Ruissalo Botanical Garden, Turku, Finland [145]

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EUDICOTS

Berberis buxifolia, Royal Botanic Gardens, Kew, UK [146]

Berberis haematocarpa in fruit, Rancho Santa Ana Botanical Garden, California, USA [146]

Diphylleia cymosa, Royal Botanic Gardens, Kew, UK [146]

Nandina domestica, Kunming Botanical Garden, China [146]

Epimedium wushanense, Royal Botanic Gardens, Kew, UK [146]

Sinopodophyllum hexandrum, Royal Botanic Gardens, Kew, UK [146]

146. BERBERIDACEAE

of six to nine, free, petal-like sepals in two or three whorls and six free petals that may be hooded, pouched or spurred. Nectaries are sometimes present. The six stamens have two thecae that open by valves or slits. When touched by an insect, the stamens of some taxa (notably some Berberis species) move quickly towards the style, effecting pollen placement on an insect. The superior ovary has one carpel with a terminal, often persistent style (when a style is present). The fruit is a berry, capsule, follicle or utricle with one to many seeds.

from each other c. 90 million years ago. Three subclades can be recognised in Berberidaceae, represented by three wellsupported subfamilies, but how these are related to each other is not certain. Nandina is morphologically distinctive and has been placed in its own family in the past, but it has some similarities to Caulophyllum. Mahonia is included in Berberis, the distinction between the two genera being vague. Aceranthus is merged with Epimedium, and Vancouveria is sister to this genus. Plagiorhegma is a synonym of Jeffersonia, and Podophyllum has been shown to be polyphyletic, resulting in the acceptance of Dysosma.

Barberry family

Berberidaceae include evergreen and deciduous, perennial herbs, shrubs and sometimes small trees. Stems are erect and woody or wholly or partially underground and rhizomatous or tuberous, with or without prickles. The alternate or opposite leaves are simple or pinnately or ternately compound, sometimes with stipules. Venation is pinnate or palmate. The terminal or axillary flowers are solitary or in fascicles, racemes, spikes, umbels, cymes or panicles. The bisexual flowers are radially symmetrical and sometimes have bracts. The perianth is sometimes absent, but is usually composed

Distribution: Berberidaceae occur in temperate climates worldwide, with woody genera (such as Berberis) more diverse in tropical mountains and the temperate regions of the Southern Hemisphere, and herbaceous genera being more diverse in the northern continents. Several genera (Achlys, Caulophyllum, Diphylleia and Jeffersonia) have a disjunct distribution in East Asia and North America. Phylogeny and evolution: Berberidaceae form a clade with Ranunculaceae, separating

Genera and species: This is a family of 14 genera and c. 700 species in three subfamilies. Berberidoideae – Berberis (c. 600), Ranzania (1); Nandinoideae – Caulophyllum (3), Gymnospermium (9), Leontice (3) and Nandina (1); Podophylloideae – Achlys (3), Bongardia (1), Diphylleia (3), Dysosma (c.10), Epimedium (c. 55), Jeffersonia (2), Podophyllum (2), Sinopodophyllum (1) and Vancouveria (2). Plants of the World

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RANUNCULALES Uses: Fruits of barberry (Berberis vulgaris), American barberry (B. canadensis), holly grape (B. repens), Texas mahonia (B. swaseyi), calafate (B. heterophylla and B. microphylla), michay (B. darwinii) and algerita (B. haematocarpa) are high in vitamin C and sometimes made into preserves or used in cooking. A yellow dye was historically extracted from barberry plants, and the fine wood was used for toothpicks. Berberis species are alternative hosts of black stem rust also affecting wheat and other cereals, and eradication of Berberis has been advocated as a control measure for this economically destructive disease since at least the 17th century in Europe and the 18th century in North America. An extract of Podophyllum is used in some laxative preparations. Several genera are important in horticulture, especially Berberis (barberry and mahonia), Epimedium (fairy wings or horny goat weed) and Nandina (sacred bamboo), many of which are highly valued plants for hedging or the woodland garden. Nandina domestica is believed to be extinct in the wild and to only have survived due to cultivation for centuries in temple gardens in East Asia. Etymology: Berberis is the Latinised form of the Arabic word ‘alburbaris’ (‫ )سيرب رابلا‬for the plant.

147. RANUNCULACEAE Buttercup family

This family includes mostly perennial, annual, sometimes aquatic herbs, woody vines and short shrubs. Leaves are alternate (rarely opposite), simple, three-parted or palmately, pedately or once or twice pinnately compound and usually lack stipules. Flowers can be terminal or axillary, solitary, or in

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racemes, cymes or panicles, usually with bracts. The usually bisexual, regularly symmetrical f lowers (actinomorphic or zygomorphic) have a perianth, with variable numbers of parts (sometimes absent) that is differentiated into sepals and petals or all whorls petal-like, often pentamerous. Segments are usually free, but can be clawed, spurred or hooded. Stamens are usually numerous and arranged in spirals or whorls. The usually free carpels are numerous and whorled. Fruits are usually nuts, follicles or berries, sometimes with elongated plumes that are formed by the persistent styles. Most species are insect-pollinated, but some are pollinated by wind or hummingbirds. Distribution: These plants are found in temperate regions worldwide, extending into the tropics with a few species. They are missing from Antarctica and desert regions of Africa and Australia. Clematis and Ranunculus are among the most widespread genera of flowering plants. Phylogeny and evolution: The crown node has been estimated to be c. 50 million years old. A 125–122 million year old (Early Cretaceous) fossil, Leefructus, was found in China and has been assigned to this clade, which, if correct, may provide different insights on eudicot evolution. Eocaltha, 77 million years old, from the Mexican Cretaceous, and Paleoactaea, 58 million years old, from the Late Palaeocene are similar to modern members of Caltha and Actaea, respectively. The evolution of Delphinium probably occurred around the same time as the diversification of bumblebees, 40–25 million years ago. Members of Circaeasteraceae and Paeoniaceae were previously placed in or near this family. Glaucidium, previously placed in Paeoniaceae or in its own family, is close to Hydrastis; together, they are sister to the rest of Ranunculaceae (or form a grade). These two genera do share morphological similarities, but in some analyses the two are not found in the same clade and thus separate subfamilies have been proposed; the similarities between the two have been interpreted as plesiomorphic for the family.

Molecular studies have resulted in major reorganisations of genera in this family. For instance Anemone has been expanded with Barneoudia, Hepatica, Knowltonia, Miyakea, Oreithales and Pulsatilla, but it was recently found that this interpretation was due to incorrect outgroup sampling and in fact these genera form a grade leading up to Clematis. Thus Anemone needs to be divided, accepting Anemonidium, Eriocapitella, Hepatica, Pulsatilla and an expanded Knowltonia. Beckwithia, Ceratocephala, Coptidium, Ficaria, Krapfia, Kumlienia, Laccopetalum and Myosurus are part of a clade including Ranunculus and these genera are merged, but Oxygraphis, Paroxygraphis and Halerpestes are maintained due to the unexpected position of Trautvettera in this clade (formerly in Thalictroideae). Cimicifuga and Souliea are part of Actaea, and Aconitella and Consolida are embedded in Delphinium. Even though Delphinium is polyphyletic with regard to Aconitum, these genera can be maintained when Staphisagria is accepted. Megaleranthis has been merged with Trollius. Anemonella and Paropyrum are now placed in Thalictrum, and Enemion is better placed in Isopyrum. Genera and species: Ranunculaceae are a family of 43 genera and c. 2,346 species in five subfamilies: Glaucidioideae – Glaucidium (1); Hydrastidoideae – Hydrastis (1); Coptidoideae – Coptis (15) and Xanthorhiza (1); Ranunculoideae – Aconitum (c. 300), Actaea (27), Adonis (26), Anemoclema (1), Anemone (c. 75), Anemonidium (14), Anemonopsis (1), Asteropyrum (2), Beesia (2), Calathodes (3), Callianthemum (14), Caltha (12), Clematis (c. 332), Delphinium (c. 350), Eranthis (9), Eriocapitella (6), Halerpestes (c. 10), Hamadryas (6), Helleborus (21), Hepatica (5), Knowltonia (28), Nigella (22), Oxygraphis (5), Paroxygraphis (1), Pulsatilla (c. 33), Ranunculus (c. 625), Staphisagria (3), Traut vetteria (1) and Trollius (31); Thalictroideae – Aquilegia (80), Dichocarpum (20), Isopyrum (2), Leptopyrum (1), Metanemone (1), Paraquilegia (5), Paropyrum (1), Semiaquilegia (1), Thalictrum (c. 250) and Urophysa (2).

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Clematis alpina, private garden, Kingston upon Thames, Surrey, UK [147]

Adonis annua, Royal Botanic Gardens, Kew, UK [147]

Hydrastis canadensis, Sarah Duke Botanical Nigella papillosa near Ronda, Spain [147] Garden, Durham, North Carolina, USA [147]

Pulsatilla slavica, Helsinki Botanical Garden, Finland [147]

Delphinium consolida, Hengelo, the Netherlands [147]

Ranunculus californicus, Point Bonita, California [147]

Glaucidium palmatum, Royal Botanic Gardens, Kew, UK [147]

Anemone nemorosa near Turku, Finland [147]

Xanthorhiza simplicissima, private Helleborus foetidus, Royal Botanic Gardens, Kew, UK [147] garden, Kingston upon Thames, Surrey, UK [147]

Thalictrum aquilegifolium, Helsinki Botanical Garden, Finland [147]

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Uses: Kalonji or kaljeera, Nigella sativa, is used as a spice in baking, although it is toxic if used to excess. Goldenseal (Hydrastis canadensis) has been used extensively by Native Americans as a general medicine and a dye. The plant has been used to treat various ailments, from cancer to ophthalmia. Currently it is used as a natural antibiotic, but it is not sustainably harvested and is therefore threatened in North America. Its international trade is restricted by CITES. Many species are poisonous, and some are deadly; Aconitum species have been

the cause of accidental and intentional poisonings. Despite these unpleasant properties, Ranunculaceae remain among the most popular of garden plants, including Aconitum (monkshood), Actaea (baneberry), Adonis (pheasant’s eye), Anemone (anemone), Aquilegia (columbine), Caltha (marshmarigold), Clematis (traveller’s joy), Delphinium (larkspur), Eranthis (winter aconite), Eriocapitella (Japanese anemone), Helleborus (hellebore), Hepatica (liverleaf), Nigella (love -i n-a-m ist), Pulsat illa (pasquef lower), Ranunculus (buttercup),

Thalictrum (meadow rue) and Trollius (globeflower). Record: Together with a species of Brassicaceae, Ranunculus lobatus, can be found at an elevation of 7,756 m in the Himalayas, the highest elevation at which any seed plant will grow. Etymology: Ranunculus, a name already in use for the plant in classical times, is a diminutive form of the Latin rana, a frog, probably because many species grow in wet places.

PROTEALES Families 148 to 151 comprise Proteales, an unlikely assembly of aquatic herbs, windpollinated trees and shrubs, vines and trees, often with colourful nectar-producing flowers. Sabiaceae have not been found to be a member of Proteales in all analyses, but they share some characters with Proteaceae, such as the nectariferous hypogynous disc. They differ markedly in numbers of floral parts and many other characters, but considering the diversity in the rest of Proteales, Sabiaceae do not add much additional heterogeneity. Characters that are shared among Proteales are the anther connectives extending beyond the anther locules and frequent occurrence of tubular leaf epidermal waxes.

148. SABIACEAE Pao-hua family

Sabia yunnanensis, Guo Ba Yan, Han Bai Yu Shan, Sichuan, China (CD) [148]

Meliosma arnottiana, Royal Botanic Gardens, Kew, UK [148]

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Meliosma dilleniifolia, Royal Botanic Gardens, Kew, UK [148]

These deciduous and evergreen trees, shrubs and woody vines have alternate, simple or once pinnate leaves without stipules. The petioles or petiolules are often pulvinate, and the blades are frequently covered with red glands. Inf lorescences are axillary or terminal, usually cymes or panicles, sometimes the flowers solitary. The bisexual or rarely polygamous-dioecious plants have actinomorphic or zygomorphic flowers that are generally tiny. The usually five sepals are free or fused at the base and are equal or

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Superhydrophobic leaves of sacred lotus, Nelumbo nucifera, Helsinki Botanical Garden, Finland [149]

Nelumbo nucifera, Brooklyn Botanical Garden, USA [149]

Nelumbo nucifera, fruiting, Royal Botanic Gardens, Kew, UK [149]

unequal in size. The usually five petals are equal or the inner two are much reduced. The five stamens are attached to the petals at the base and are often all fertile, or three of them are staminodial. The filaments are filiform or expanded below the anther to form a collar. The anthers have two thecae on either side of a narrow or thickened cup-shaped connective that forms a central pore through which the style grows. The superior ovary is bi- or rarely trilocular. The fruit is a drupe or indehiscent follicle with a single seed.

Genera and species: This family has three genera with about 66 species: Meliosma (c. 40), Ophiocaryon (7) and Sabia (19).

filaments and anthers that open by slits. The connective has an incurved appendage. The superior ovaries are numerous and individually embedded in the flattened top of a turbinate receptacle. The style is short and the stigma nearly sessile. Fruits are indehiscent nuts that are loose in cavities in the receptacle. They are pollinated by insects, usually beetles.

Etymology: Sabia is derived from the Hindi sab-ya, the Indian name for Sabia lanceolata.

149. NELUMBONACEAE Sacred-lotus family

Distribution: Sabiaceae are distributed in warm temperate and tropical Central and South America and East Asia. Phylogeny and evolution: Sabiaceae are sometimes treated as the only family in the order Sabiales, but have recently been found to be sister to the remaining Proteales and included with them. Fossils of Insitiocarpus from the Middle Cretaceous (c. 90 million years old) are believed to be Sabiaceae and Meliosma and Sabia from the Late Cretaceous are also known. The family is estimated to have appeared c. 120 million years ago (stem node). Pollination of Meliosma is remarkable; the anthers open within the bud, enclosed and held under pressure by the staminodes. The bud opens explosively upon rough contact (for instance by a visiting insect), releasing the pollen, although pollen also collects in the broad connectives from where insects can collect it.

This family of perennial aquatic herbs has branched, slender, underground, creeping rhizomes, of which the terminal portions become tuberous late in the growing season. Vegetative parts of the plant have conspicuous air chambers. Leaves arise alternately from the rhizome and are floating or emergent. The petiole is long and warty, the blade round, peltate, and the margin entire. The surface is covered with a layer of water-repellent wax. The bisexual flowers emerge solitarily from the leaf axils and are held above the water surface on a long peduncle. The free petals are numerous, the outermost somewhat reduced, the inner ones larger and more petal-like. The stamens are numerous and have slender

Distribution: The family occurs in temperate and subtropical North America, tropical and temperate southern and East Asia. One species is naturalised throughout the tropics. Phylogeny and evolution: Nelumbonaceae have frequently been included in Nymphaeaceae s.l., although on the basis of the leaf anatomy and flower morphology it had already been suggested that the family must be distantly related. However, gene expression in the flowers of Nelumbo and Nymphaea is remarkably similar. It has been shown by DNA analyses that Nelumbonaceae are most closely related to Platanaceae and Proteaceae, sharing characters of the seed with Proteaceae. Mid Cretaceous fossils known as Nelumbites share the remarkable f lower morphology but have different leaf venation, so this may or may not be associated with this family. An age of over 100 million years is likely for this lineage. Flowers thermoregulate in order to release a scent to attract insects, and the c. 30 ºC temperature rewards visiting insects with a stable environment for mating.

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PROTEALES Genera and species: The family has a single genus, Nelumbo, with three species: N. komarovii, N. lutea and N. nucifera. The last may have to be subdivided on the basis of anatomical characters. Uses: The tubers formed by mature plants of sacred lotus are edible and cultivated as a food crop, especially in tropical Asia. Their seeds are also widely used in Asian cuisine. Nelumbo nucifera is the sacred flower of Buddhism and Hinduism, and it is the national flower of India and Vietnam. Sacred lotus is frequently cultivated in (sub-)tropical water gardens and ponds. The leaves of lotus are superhydrophobic, and the wax of the leaves is now synthetically imitated as a highly waterrepellent wax used in industry and as a paint for cars etc. Etymology: Nelumbo is derived from nelum, the Sinhalese word for the lotus flower.

150. PLATANACEAE Plane-tree family

Platanus racemosa showing flaking bark, Palm Springs, California, USA [150]

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This is a family composed of large deciduous trees, with erect trunks that have smooth bark exfoliating in thin plates to form a mosaic of colours, becoming furrowed when older. Twigs have buds that are hidden by the base of the swollen petiole and often bear simple or compound hairs, the compound ones with whorls of branches, appearing stellate. The alternate, simple leaves are palmately lobed, with three, five or seven lobes or unlobed. The petioles are flanked by sheathing stipules with entire or serrate margins. The leaf bases are cordate, truncate or cuneate, and the blade surface is tomentose with stellate hairs, becoming glabrous with age. The unisexual inflorescences occur on the same plant (monoecious) and are axillary or solitary, crowded into dense heads, and emerge together with the leaves. Male inflorescences are sessile, with one to five, globose heads that soon fall off. Female inflorescences are sessile or stalked, terminal, with one to seven globose heads in pendulous racemes. Individual flowers are usually tri- or tetramerous and minute, the sepals free or fused at the base and the petals vestigial. Male flowers have stamens that are as many as the numbers of sepals, the connective enlarged to a peltate appendage, and pistillodes are sometimes present. Female flowers have three or four staminodes and three to eight free superior carpels that are placed in two or three whorls, with linear styles that have the stigmatic surface on the inner side. Fruits are achenes that are densely crowded in the heads, often Platanus ×acerifolia (P. occidentalis × P. orientalis), Hyde Park, London, UK [150]

persisting on the tree until spring, surrounded by numerous hairs that are basally attached and simple. Flowers are wind-pollinated. Distribution: The family occurs in North America, from Canada and the Great Lakes to central Mexico and Guatemala, southeastern Europe and warm-temperate Asia, south to Laos. Phylogeny and evolution: The family is well represented in the fossil record of North America and Eurasia and was once more widespread across the Northern Hemisphere. The relationship between Platanaceae and Proteaceae may explain why there are so many platanoid leaf fossils in the Southern Hemisphere. Some Proteaceae have wood that is similar to that of Platanaceae, although in general the wood anatomy of the two families is different, which may be due to the different climatic conditions in which these families are found. The family is estimated to have diverged c. 110 million years ago, and fossils of this approximate age, like Platanocarpus, are known. Genera and species: The family includes only the genus Platanus with eight species. Uses: The wood of sycamore (US) or plane (UK) (Platanus) is resistant to splitting, and it is therefore traditionally used to make buttons and butchers’ blocks, but the wood is Grevillea rhyolitica flowers visited by an eastern spinebill (Acanthorhynchus tenuirostris) in the Australian National Botanic Gardens, Canberra [151]

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difficult to work and of limited commercial value. Several species and hybrids are popular shade or avenue trees especially in urban areas because they are tolerant to pollution and respond well to pruning. However, the hairs formed on the new growths can cause irritation and allergic reactions in humans.

or four glands are usually present around the ovary, and these are scale-like or fleshy, sometimes fused. The superior (sometimes semi-inferior) ovary is composed of a single locule, topped by a single simple, persistent style. The fruit is a dehiscent or indehiscent woody or coriaceous follicle, achene, nut or drupe, fused in cones in some cases. Seeds are sometimes winged.

This is a family of creeping to upright shrubs and trees, sometimes appearing herbaceous, but then with persistent woody roots. Roots are often produced in short clusters (proteoid roots, a unique character). Stems are usually covered with short, three-celled hairs. The alternate, or rarely opposite or whorled, leaves are simple or pinnatifid, pinnately or bipinnately compound, rarely palmately so, usually leathery and stiff. They lack stipules and usually have a pinnate venation. The inflorescences are simple or compound, axillary or terminal, racemes, panicles or condensed heads, umbels, corymbs or cones or with solitary flowers. The usually bisexual flowers are zygomorphic or actinomorphic, usually one or two in the axils of bracts, which are sometimes absent. The four petal-like tepals are free or variously fused, and each petal has a slightly expanded valved limb. The four stamens are usually all fertile and oppose the tepals, with the filaments partly or completely fused with these, often forming colourful ‘pollen presenters’, rarely free, the anthers are often sessile on the perianth. Two

Phylogeny and evolution: Proteaceae are known to have been diverse in the Late Cretaceous and Eocene of Australia, and some genera such as Cardwellia and Gevuina may show a distribution related to continental

Leucadendron discolor, San Francisco Botanical Garden, USA [151]

Macadamia integrifolia, Royal Botanic Garden, Sydney, Australia [151]

Persoonia pinifolia, Royal Botanic Garden, Sydney, Australia [151]

Banksia hookeriana, Western Australia [151]

Protea cynaroides, Royal Horticultural Society Garden, Wisley, UK [151]

Telopea speciosissima ‘Corroboree’, Australian National Botanic Garden, Canberra [151]

Etymology: The name comes from πλάτανος ( platanos), the classical Greek name for Platanus orientalis, possibly related to Greek πλάτος (platos), flat.

151. PROTEACEAE Sugarbush family

Distribution: This family has its greatest diversity in the Southern Hemisphere, especially in Australia and southern Africa. Smaller numbers of species are known from Mexico, Central and South America, tropical Africa, Madagascar, India, East and Southeast Asia, Malesia, New Caledonia, New Zealand and Fiji.

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PROTEALES drift, whereas others such as the genus pair Brabejum and Panopsis are certainly a product of more recent long-distance dispersal. Banksia may be c. 60 million years old, and fire may have spurred diversification in this and other Australian genera. Diversification of Proteaceae in the Cape of South Africa may have been more recent than in Australia and is likely a result of edaphic speciation. Some generic reorganisation may be needed in the future. Dryandra has already been included in Banksia, but Grevillea, for example, may have to be expanded to include Hakea and Finschia, and other recircumscriptions of genera are likely when the phylogenetics of Proteaceae become better understood. Genera and species: Proteaceae are a family of 79 genera and about 1,750 species in five subfamilies: Bellendenoideae – Bellendena (1); Persoonioideae – Acidonia (1), Garnieria (1), Persoonia (c. 100), Placospermum (1) and Toronia (1); Grevilleoideae – Alloxylon (4), Athertonia (1), Austromuellera (2), Banksia (169), Bleasdalea (2), Brabejum (1), Buckinghamia (2), Cardwellia (1), Carnarvonia (1), Catalepidia (1), Darlingia (2), Embothrium (1), Eucarpha (2), Euplassa (20), Finschia (3), Floydia (1), Gevuina (1), Grevillea (362), Hakea (149), Helicia (c. 100), Heliciopsis (14), Hicksbeachia (2), Hollandaea (2), Kermadecia

EUDICOTS

(4), Knightia (1), Lambertia (10), Lomatia (12), Macadamia (9), Malagasia (1), Megahertzia (1), Musgravea (2), Neorites (1), Opisthiolepis (1), Oreocallis (1), Orites (8), Panopsis (25), Roupala (33), Sleumerodendron (1), Sphalmium (1), Stenocarpus (21), Strangea (3), Telopea (5), Triunia (4), Turrillia (3), Virotia (6) and Xylomelum (6); Proteoideae – Adenanthos (33), Aulax (3), Beauprea (13), Beaupreopsis (1), Cenarrhenes (1), Conospermum (53), Diastella (7), Dilobeia (2), Eidothea (2), Faurea (c. 15), Franklandia (2), Isopogon (35), Leucadendron (80), Leucospermum (48), Mimetes (13), Orothamnus (1), Paranomus (19), Petrophile (53), Protea (114), Serruria (51), Sorocephalus (11), Spatalla (20), Stirlingia (7), Synaphea (50) and Vexatorella (4); Symphyonematoideae – Agastachys (1) and Symphionema (2). Uses: Seeds of several species can be eaten. Macadamia nuts are perhaps the best known, and Macadamia integrifolia, M. ternifolia and M. tetraphylla are now grown commercially for their seeds, especially in Australia and Hawaii. The delicious and nutritious seeds are also valued in the cosmetic industry, with the oil being considered a botanical alternative to mink oil. Chilean hazel (Gevuina avellana) is commonly consumed in South America and New Zealand. Other genera that produce edible nuts are Floydia prealta (coohoy nut),

Hicksbeachia pinnatifolia (red boppel nut) and Finschia chloroxantha. Nectar of some species is also harvested in traditional societies. Banksia fruit ‘cones’ are woody and sometimes used for carving. Cross-sections of the fruits make attractive decorations. Several species of Banksia, Leucadendron, Leucospermum, Protea and Telopea are popular in the cut-flower industry, and many species are grown as garden ornamentals especially in areas with Mediterranean climates. Warratah (Telopea speciosissima) is the floral emblem of New South Wales, but as a cut flower it is exported as ‘kiwi rose’ from New Zealand. Oldest clone: Lomatia tasmanica is known from a single clone only. All individuals are triploid and sterile, and the plants spread vegetatively. Based on fossil evidence, these clones formed 43,600 or more years ago, and the remaining 500 plants now cover an area of 1.2 km. Even though each individual part of this plant lives for ‘only’ some 300 years, the clone is one of the oldest plants alive. Etymology: Protea is named for Πρωτεύς (Proteus), a half-god of sea and rivers and the oldest son of Poseidon in ancient Greek mythology. Proteus was capable of appearing in many forms, which members of this family do as well.

TROCHODENDRALES This order consists of the single family Trochodendraceae, which has a fossil history dating back to the Late Cretaceous. It was widespread across the Northern Hemisphere, sometimes at high latitudes. Fossils of both extant genera have been known since the Eocene.

Tetracentron sinense, China (YN) [152]

152. TROCHODENDRACEAE Wheel-tree family

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This family of evergreen trees and shrubs has (near) vessel-less wood. Leaves are spirally arranged in false whorls or on elongate branches, without stipules or with a stipule covering the bud and later fused with the petiole. The blades are simple and pinnately or palmately veined. The terminal racemes or panicles bear bisexual flowers subtended by two to five bracts (Trochodendron) or catkinlike spikes in whorls of four (Tetracentron). Flowers have a broadened receptacle and

BUXALES

EUDICOTS

lack a perianth (Trochodendron) or have four sepals only (Tetracentron). In Trochodendron the 40–70 stamens are placed in several whorls around the (four to) six to 17 carpels in a single, laterally fused whorl. In Tetracentron only four stamens surround the four basally fused carpels. The fruit is composed of the laterally fused follicles that open on the inside with short slits. Seeds are winged or not. Distribution: This family is distributed in temperate and subtropical East Asia, from

Japan south to Taiwan and southern China, Nepal and northwestern India.

the Northern Hemisphere (as Nordenskioldia) from the Late Cretaceous.

Phylogeny and evolution: Trochodendraceae are the sole member of the order Trochodendrales; in some previous classifications, each genus was placed in its own family. Previously the genera were of unclear relationships and were sometimes linked with Cercidiphyllaceae (Saxifragales) or Eupteleaceae (Ranunculales). Now restricted to isolated populations in East Asia, fossils are known from around

Genera and species: The family has two genera, each with a single species: Tetracentron sinensis and Trochodendron aralioides.

Tetracentron sinense, China (YN) [152]

Etymology: Trochodendron is derived from Greek τροχός (trochos), a wheel, and δένδρων (dendron), a tree, in reference to the stamen arrangement.

Trochodendron aralioides, Brooklyn Botanical Garden, USA [152]

BUXALES This order diverged probably around 120 million years ago. It consists of the single family Buxaceae.

153. BUXACEAE Box family

Buxaceae are trees, shrubs and rarely herbs that are usually glabrous, sometimes hairy, the hairs simple. Leaves are opposite or alternate and simple, lack stipules and are pinnately veined, but sometimes have three prominent veins from the base. Inf lorescences are bracteate axillary or terminal spikes, racemes or cymes, often congested. The unisexual flowers have an undifferentiated perianth (tepals) or none. Male flowers usually have four free tepals

(none in Haptanthus) and four to six (to many) usually free stamens (two fused stamens in Haptanthus), the filaments absent to enlarged and showy, the anthers opening by lengthwise slits or valves; a pistillode is sometimes present. Female flowers are usually few and larger than the male flowers, with four to six tepals (none in Haptanthus). The superior ovary is composed of two to three fused carpels that are topped with two or three styles. Fruits are loculicidally Plants of the World

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BUXALES dehiscing capsules or drupes, the styles usually persistent and showy. Distribution: The family has a patchy distribution across the world, from the southern USA through Mexico, the Caribbean and the Andes to the Azores, western and southern Europe, North Africa, the Caucasus, Sub-Saharan Africa, Madagascar, southern India and Sri Lanka, the Himalayas to Southeast and East Asia, the Philippines and western Indonesia. Phylogeny and evolution: Madagascan Didymeles and Honduran Haptanthus are sometimes placed in their own families. Didymeles is probably sister to the other genera; Haptanthus is now known to be sister to just Buxus. When Haptanthus was first

EUDICOTS

described, it could not be assigned to a family, but it is clear now that it belongs to Buxaceae. Fossil Didymeles is known from New Zealand. Crown group Buxaceae are at least 111 million years old, a date corroborated by fossils from the Early Cretaceous. The family was widespread in the Northern Hemisphere during the Tertiary. Genera and species: The family has six genera with c. 65 species: Buxus (c. 40), Didymeles (2), Haptanthus (1), Pachysandra (3), Sarcococca (13) and Styloceras (6). Uses: The fine grain of boxwood (Buxus) is good for wood carving, particularly for decorative storage boxes, hair combs and chess pieces. It was the preferred wood for woodblock book printing, and it still is

commonly used to make musical instruments such as string instruments and bagpipes. Buxus sempervirens is commonly planted for garden hedging and topiairy, Sarcococca is planted for winter scent, and Pachysandra terminalis is a common evergreen ground cover in shady gardens. Box branches as a substitute for palm fronds are blessed with holy water in the Catholic church on Palm Sunday. Etymology: Buxus is the classical Latin name for B. sempervirens, which in turn is derived from Ancient Greek πυξός (puksos), the classical name for a box tree. Formerly more widespread in England, B. sempervirens is commemorated in place names including Box Hill (where it still occurs) and Bix Bottom (where it is now absent).

Buxus sempervirens, Box Hill, UK [153] Buxus obtusifolia, near Malindi, Kenya [153]

Sarcococca hookeriana, Royal Botanic Gardens, Kew, UK [153]

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Haptanthus hazlettii, Honduras (AS) [153]

Pachysandra terminalis, Royal Botanic Gardens, Kew, UK [153]

GUNNERALES

EUDICOTS

GUNNERALES Families 154 and 155 form the order Gunnerales. The two genera in this order have been merged into one family by some authors, but their differences are so great that it is deemed better to maintain them in separate families. They share some wood anatomical characters (hydathodes), stipules, excretion of resin and lack of a perianth, but one is often a giant herb of humid habitats, the other a resurrection shrub of arid places, and the hydathodes and stipules are not of similar ontogeny. The age of this lineage (stem node) has been suggested to be somewhere between 118 and 77 million years.

154. MYROTHAMNACEAE Resurrection-shrub family

These aromatic, glabrous shrubs are unisexual (dioecious). Their narrowly winged main branches produce short lateral vegetative shoots that carry the opposite, sessile leaves, which can survive extensive desiccation. The leaves are obtriangular, broadly sheathing at the base and surrounded by a tooth-like stipule. The leaves are flabellate and plicate with all veins emerging from the sheathing base and at their distal end dentate. The catkin-like inflorescences terminate short shoots. The flowers occur in triads at the base of the

Myrothamnus flabellifolius, South Africa (NH) [154]

inflorescence, solitary above, with the flowers or triads opposite each other and subtended by a bract. The sessile flowers sometimes have two bracteoles and two to four scale-like tepals, sometimes tepals absent, usually present in terminal flowers. The male flowers have three to eight stamens, with short filaments and basifixed anthers that open by lengthwise slits. Female flowers have a superior ovary composed of three to four partially fused carpels and free styles and recurved stigmas. The fruits are capsules that open from the tip exposing the numerous small seeds.

Genera and species: Myrothamnus is the sole genus in this family and has two species: M. flabellifolius and M. moschatus. Etymology: Myrothamnus is composed of the Greek μύρων (myron), a scent, and θάμνος (thamnos), a shrub.

155. GUNNERACEAE Giant-rhubarb family

Distribution: This family is restricted to arid regions of southern and tropical East Africa and western Madagascar. Phylogeny and evolution: Myrothamnus is the only woody resurrection plant. Its leaves dry out completely during the dry season, but turn green in a short time after exposure to water. It is most closely related to Gunneraceae, but there is no fossil record.

This family includes usually perennial, rarely annual, rhizomatous and stoloniferous small to gigantic herbs. They have a close association with the nitrogen-fixing cyanobacterium

Gunnera pilosa, Ecuador [155]

Gunnera prorepens, Beth Chatto Gardens, UK [155]

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GUNNERALES Nostoc in their stems and roots. Their leaves are alternate and placed spirally at the tip of the rhizome, emerging in the axils of stipule-like scale-leaves. The blades have a toothed margin, palmately forking venation that is prominent on the lower surface, and a finer reticulate tertiary venation. Inflorescences are usually branched racemes, simple racemes or spikes or panicles. The numerous actinomorphic, bisexual flowers cover the inflorescences. They have two, sometimes three sepal-like tepals and the same number of petal-like tepals that are usually hooded, or the inner whorl sometimes absent. The usually two (rarely one) stamens are fused with the petals. The inferior ovary is composed of two fused carpels forming one locule. The fruits are nut-like drupes. Distribution: This family occurs in mesic habitats in tropical mountains and temperate areas of the Southern Hemisphere, from Mexico to Patagonia and the Falkland Islands, East African mountains from Ethiopia to the

Tetracera indica, Lazarus Island, Singapore [156]

Tetracera loureiri in fruit (WA) [156]

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Cape, northern Madagascar, the Malesian Archipelago, Tasmania, New Zealand and Hawaii. Phylogeny and evolution: The distinctive pollen has been found in sediments as old as the Early Cretaceous and was known at that time from all southern continents, leading some to conclude that this family may have a distribution related to vicariance events after the separation of Gondwana. However, its present-day occurrence on volcanic islands like Juan Fernández and Hawaii indicates that long-distance dispersal is also common. The annual Gunnera herteri is sister to the rest of the genus. Gunneraceae were in the past often associated with Haloragaceae or Saxifragales. They are certainly inedible and not related to rhubarb (Rheum, Polygonaceae) to which the common name refers. Mutualistic association: Glands on the stem just below the leaves secrete mucilage Dillenia suffruticosa, Lazarus Island, Singapore [156]

that contains various sugars attracting the cyanobacterium Nostoc. The motile hormogonia of Nostoc enter the plant via the glands and once inside lose their mobility and form heterocysts; they often do not photosynthesise. Nostoc fixes nitrogen, and in return Gunnera provides Nostoc with sugars. Genera and species: Gunneraceae include only Gunnera, with 63 species. Uses: Gunnera is frequently planted as a pond margin plant in gardens in temperate climates. Etymology: Gunnera is named in honour of Norwegian bishop and botanist Johan Ernst Gunnerus (1718–1773), who was the first to describe the basking shark and published the Flora Norvegica. Like his contemporary Linnaeus, who named the genus for him, he also made an expedition to Lapland and described the habits of the Saami people in detail. Dillenia philippinensis, Singapore Botanical Garden [156]

Hibbertia acerosa, Mt Benia, Western Australia [156]

SAXIFRAGALES

EUDICOTS

DILLENIALES This order has been difficult to place and consists of the single family Dilleniaceae with an estimated age of 114 million years (stem node).

156. DILLENIACEAE Guineaflower family

This family includes trees, shrubs, woody climbers and a few herbs. The alternate (rarely opposite) spirally arranged leaves are simple, rarely pinnatisect or pinnately compound and petiolate. There are no stipules, and petioles are sometimes winged. Blades are often toothed, and venation is pinnate with strong, parallel secondary veins that end in teeth. Inflorescences are racemes, panicles or cymes, but frequently the flowers are formed singly in the leaf axils. The bisexual, rarely unisexual, flowers are mostly actinomorphic, occasionally slightly zygomorphic (Hibbertia) or strongly zygomorphic (Schumacheria). They have three or four to five, sometimes up

to 18, sepals that are leathery or succulent and persistent in fruit. The two or three to five, sometimes to seven free petals are crumpled in bud. Stamens are usually numerous (or one to ten) and free or fused at the base and then in fascicles. Anthers are basifixed and open by lengthwise slits or apical clefts or pores. Staminodes are also often present. The superior ovary is composed of two to seven (or up to 20) free or partially fused carpels, tipped with free styles terminated by the stigma. Fruits are follicles (sometimes compound), berries, or capsules often enclosed by fleshy or leathery sepals. Seeds are often arillate. Distribution: This pantropical family extends into warm-temperate Asia and Australia. Phylogeny and evolution: This is the sole family in the order Dilleniales, which have proved difficult to place in eudicot classification, even with the aid of whole plastid genome sequences and many low-copy nuclear genes. The stem age may be c. 115 million years old, but the crown group diverged c. 52 million years ago. A subfamilial classification has been proposed,

but not all genera are yet placed so it is therefore not adopted here. Previously, the family was viewed as the type of subclass Dilleniidae, but they have no close relatives based on molecular studies, and as mentioned have defied a wellsupported assignment to any major clade of eudicots, which makes their assignment to their own order appropriate. Genera and species: Dilleniaceae include 12 genera with c. 300 species: Acrotrema (10), Curatella (3), Davilla (18), Didesmandra (1), Dillenia (60), Doliocarpus (40), Hibbertia (c. 120), Neodillenia (3), Pachynema (7), Pinzona (1), Schumacheria (3) and Tetracera (c. 44). Uses: Fruits of Dillenia crenatifolia, D. indica and D. salomonensis are edible. Some species of Dillenia and Hibbertia are grown as garden ornamentals. Etymology: Dillenia was named by Carolus Linnaeus in honour of German botanist Johann Jacob Dillenius (1684–1747), who hosted Linnaeus in Oxford in 1736.

SAXIFRAGALES Families 157 to 171 form the diverse order Saxifragales, which are mostly distributed in the temperate zones and rare in the tropics. Relationships within the order have been problematic, and this may be the result of rapid radiation over a period of 3–6 million years during the Mid Cretaceous.

157. PERIDISCACEAE Ringflower family

Peridiscaceae are small trees and large shrubs with alternate, simple leaves. They have stipules in the leaf axils and petioles with a pulvinus at the tip immediately below entire or crenulate leaves. Flowers are formed in axillary clusters of racemes or spikes, sometimes reduced to one or two racemes. Bisexual flowers are regularly symmetrical with four to seven free sepals. The African genera have five petals, whereas the South American species lack them. Stamens are

numerous and arranged around the nectar disc, or (in Medusandra) only ten stamens are present, of which five are staminodial and the disc absent. The superior ovary can appear half inferior due to the size of the disc. A unilocular ovary consists of three or four carpels. The one-seeded fruit is a berry in American Peridiscus and a capsule in the African genera. Distribution: This family is distributed in the Guiana Shield and in West and Central Africa. Plants of the World

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Whittonia guianensis, holotype specimen collected by B. A. Whitton in Guyana in 1959 (Kew Herbarium) [157]

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Medusandra richardiana, Mokoko Forest Reserve, Cameroon (CD) [157]

Medusandra richardiana, Mokoko Forest Reserve, Cameroon (CD) [157]

Paeonia obovata in fruit, Helsinki Botanical Garden, Finland [158]

Paeonia delavayi, private garden, Hengelo, the Netherlands [158]

Paeonia cambessedesii, Royal Botanic Gardens, Kew, UK [158]

Phylogeny and evolution: Peridiscaceae had been restricted to the American genera Peridiscus and the enigmatic Whittonia, but were expanded in APG III to include the African Medusandra (formerly Medusandraceae) and Soyauxia, which in earlier classifications had been placed in Passifloraceae or the now defunct Flacourtiaceae (Malpighiales). The placement of Peridiscaceae in Saxifragales was surprising, given that this order is dominated by families from temperate regions. Some association with Paeonia has been found in molecular analyses. Whittonia is only known from its type specimen collected near the

Kaieteur Falls in Guiana and has not been recollected. It is suggested that this species may be extinct, or, because it shares many characters with Peridiscus, it may have been confused with that genus in the field.

158. PAEONIACEAE

Christenhusz, Fay & Chase

Peony family

Genera and species: This small family has four genera with 11 species: Medusandra (2), Soyauxia (7), Peridiscus (1) and Whittonia (1, possibly extinct). Etymology: Peridiscus is composed of the Greek περί (peri), around, and δίσκος (diskos), a ring.

This family of perennial herbs and shrubs has simple and branched stems, growing from short

SAXIFRAGALES

EUDICOTS

Phylogeny and evolution: In the past Paeonia was included in Ranunculaceae, especially associated with Glaucidium, or it was thought to be related to Dilleniaceae. Neither of these families made a good morphological fit, and molecular studies now place this family in Saxifragales in which an association with Peridiscaceae in some cases has been found.

woody rhizomes and often has tuberous roots. They produce stipule-like scales that clasp the winter bud on the stem. Leaves are alternate, long-petiolate and pinnate, ternate or twice ternately compound or dissected. Terminal inflorescences are usually composed of solitary flowers, but sometimes there are two or three (up to six). Flowers are bisexual with a spirally arranged perianth composed usually of four to six free sepals (sometimes fewer or more) that are unequal in size and more or less leathery. The five to 13 free petals (sometimes four or up to 25 in some cultivars) are large and showy. Stamens are free, numerous, usually 50–200 or so. Anthers are basifixed, upright and opening by lengthwise slits. Staminodes are not present, but the intrastaminal glands are sometimes interpreted as such. The superior ovary is composed of usually five (sometimes less or more) free carpels. Follicle-like fruits open adaxially exposing the large rounded seeds with an outer integument that can be showy and aril-like.

Uses: Seeds of Paeonia officinalis were reputedly eaten in Mediaeval England during Lent. Several species are of medicinal importance in the Himalayas. Many cultivars of especially P. lactiflora and P. suffruticosa are popular garden plants around the temperate zones. The largest species in the genus, P. ludlowii, is also sometimes grown and can develop into a shrub 3.5 m tall.

Distribution: The genus can be found in temperate climates in western North America south to northwestern Mexico, and it is widespread in Eurasia from the Mediterranean through Anatolia, the Caucasus, north to Siberia and the Kola Peninsula, east to Japan and Korea through China and the Himalayas.

Etymology: Paeonia is derived from the classical Greek name for the peony plant, παιονία (paionia), which in turn means ‘of Paion’. The deity Παιών (Paion or Paean) was physician to the gods in Greek mythology, and the name refers to its medicinal use in classical times.

159. ALTINGIACEAE Sweetgum family

Genera and species: Paeonia is the only genus and has 33 species. This family of deciduous and evergreen trees has alternate leaves that are usually arranged in a plane (distichous). Leaves have a simple or three-, five- or seven-lobed blade and linear stipules at the base of the petiole. They have a pleasant aroma when crushed and are pinnately or palmately veined. Unisexual f lowers lack a perianth and are wind pollinated. Male inflorescences are globular or cylindrical many-flowered heads that are grouped in terminal racemes or panicles, each male flower with one to four basal bracts and four to ten (or many) stamens that have anthers opening by lengthwise slits or valves. Female inflorescences are ball-shaped (condensed panicles) of five to 30 flowers formed subterminally on long peduncles or in the lower parts of bisexual inflorescences. Staminodes are sometimes present in female flowers and resemble needles. The (semi-) inferior bicarpellate ovary is surrounded by rudimentary scales (bracts). The fruit is a woody capsule called a ‘gumball’, the carpels opening to release the narrowly winged seeds. Distribution: The family has a relictual distribution in eastern and southern North America, Mesoamerica, southern Turkey and Greece, India, Southeast Asia, China, Taiwan, Peninsular Malaysia, Sumatra and Java.

Liquidambar styraciflua, Royal Botanic Gardens, Kew, UK [159]

Liquidambar formosana, young fruit, Mt Huangshan, Anhui, China [159]

Phylogeny and evolution: Altingiaceae were widespread across the Northern Hemisphere during the Tertiary but went extinct in many localities during the Pleistocene ice ages. They were formerly often included in Hamamelidaceae, to which they are not closely related, as subfamily Liquidambaroideae. Previously two genera were recognised,

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SAXIFRAGALES but apart from the lobed deciduous leaves Liquidambar is similar to Altingia in most characters, and the genera also hybridise. Phylogenetic studies also indicated that there is a single genus. Genera and species: The family is now considered to have only one genus, Liquidambar, with 15 species.

EUDICOTS

Etymology: Altingia is named for Willem Arnold Alting (1724–1800), Governor-General of the Netherlands East Indies. Altingia has been synonymised with Liquidambar, a name that is composed of the Latin liquidus, liquid, and Arabic ambar, amber or balm, referring to the sweet resinous sap.

160. HAMAMELIDACEAE Witch-hazel family

Uses: Levant storax (Liquidambar orientalis) produces a sweet resin, which was the balm of Gilead in the Bible. Resin of sweetgum (L. styraciflua) is antiseptic and used in skin products. Both of these species produce good timber, and the latter is a popular street tree away from its native range due to its spectacular autumn colour. Ransamala (L. excelsa) also produces good timber. Distylium pingpienense in fruit, Kunming Botanical Garden, China [160]

This family includes evergreen and deciduous shrubs and trees. Their leaves are usually alternate, rarely subopposite or opposite, and distichously or spirally arranged. Petioles are flanked by stipules that are usually paired and free, sometimes solitary with an enclosing bud (Mytilaria) or absent (Rhodoleia). Leaf blades are simple and pinnately veined or palmately (tri-)lobed and veined. Axillary or terminal inf lorescences are usually bracteate spikes or heads, rarely racemes, thyrses or panicles. Bisexual or unisexual flowers are often flanked by bracteoles and usually actinomorphic, rarely zygomorphic (Rhodoleia), the floral cup shallow to urnshaped, sometimes lacking. The four or five (sometimes ten) sepals are usually persistent, but absent in some species. Petals are absent or four or five and usually ribbon-like. The

Loropetalum chinense var. rubrum, Royal Botanic Gardens, Kew, UK [160]

Fothergilla major, Bodnant Estate, North Wales, UK [160]

Corylopsis chinensis, Royal Botanic Gardens, Kew, UK [160]

Molinadendron sinaloense, University of California Botanical Garden, Berkeley, USA [160]

Hamamelis virginiana, near Madison, Wisconsin, USA [160]

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EUDICOTS

four, five or numerous free stamens are usually all fertile, but can be arranged in two whorls with a staminodial inner whorl. The basifixed anthers open by one two valves or a simple lengthwise slit with a protruding connective. Disk scales are sometimes present between stamens and carpels. The inferior to superior ovary is composed of two carpels that are apically free, each topped by a style and stigma. Fruits are septicidally or loculicidally dehiscent, woody or leathery capsules with two or four valves. Seeds are winged or not. Distribution: This family is found in eastern North America, Mesoamerica, northern South America, eastern and southern Africa, the Comoros, Madagascar, Turkey, the southern Caspian region, and from the Himalayas through China to Korea and Honshu and south though Southeast Asia, Malesia, northern Australia (Queensland) and several Pacific islands. Phylogeny and evolution: The crown age of the family has been estimated at 26 million years old, but with a much older fossil record: Allonia decandra was described from the Cretaceous of the USA, and many fossils are known from the Early Tertiary. Four subfamilies (Disanthoideae, Exbucklandioideae, Hamamelidoideae and Mytilarioideae) are sometimes recognised, but several genera are not yet placed because they are poorly known and are in need of critical revision.

Dicoryphe (12), Disanthus (1), Distyliopsis (7), Distylium (10), Embolanthera (2), Eustigma (2), Exbucklandia (2), Fortunearia (1), Fothergilla (2), Hamamelis (4), Loropetalum (3), Maingaya (1), Matudaea (2), Molinadendron (3), Mytilaria (1), Neostearia (1), Noahdendron (1), Ostrearia (1), Parrotia (1), Parrotiopsis (1), Rhodoleia (c. 10), Sinowilsonia (1), Sycopsis (3) and Trichocladus (6). Uses: Distylium and Exbucklandia produce a valuable fine-grained timber used for furniture and turning. Witch hazel (Hamamelis virginana) was used for eye lotions in the past and is still used as a treatment for bruising. Many species are grown as garden ornamentals, especially Corylopsis and Hamamelis; the latter has sweetly scented flowers appearing before the leaves in late winter and spring. Etymology: Hamamelis (αμαμελής) is the classical Greek name for the wych elm, Ulmus glabra (Ulmaceae), but is now used for witch

Distribution: The family is restricted to China and Japan, a relictual distribution.

hazel only.

161. CERCIDIPHYLLACEAE Caramel-tree family

Genera and species: A family with 26 genera and c. 86 species: Chunia (1), Corylopsis (7), Cercidiphyllum magnificum, male flower, Royal Botanic Gardens, Kew, UK [161]

This is a family of deciduous trees with two branch types: long vegetative shoots with opposite leaves and short vegetative or reproductive shoots with a single leaf. Leaves are heart-shaped, petiolate with the latter flanked by two small stipules that fall off quickly. Blades are simple with palmate venation. Inflorescences appear before the leaves and are unisexual fascicles in heads. Each flower is usually subtended by a bract, but a perianth is lacking. Male inflorescences are sessile with four or more flowers that are difficult to separate, the inflorescence with 16–35 stamens, perhaps one to 13 per flower. Anthers are basifixed and red, opening by lengthwise slits. Female inflorescences are shortly stalked with two to eight flowers. The ovary is composed of a single naked carpel, tipped with a long-papillate stigma. The fruit is a follicle that opens away from the inflorescence axis.

Phylogeny and evolution: Cercidiphyllaceae were widely distributed across the Northern Hemisphere during the Tertiary. A great number of fossil Cercidiphyllaceae are known from the Palaeocene and Upper Cretaceous. They have an isolated position, but with Daphniphyllaceae form the sister of Hamamelidaceae. Genera and species: The sole genus in this family, Cercidiphyllum, has two similar species: C. japonicum and the smaller C. magnificum.

Cercidiphyllum japonicum, female flowers, Ruissalo Botanical Garden, Turku, Finland [161]

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SAXIFRAGALES Uses: Katsura (Cercidiphyllum japonicum) is the largest tree species in Japan. Its wood is used to make boards for the Japanese board game go. It is a valuable ornamental, the leaves in autumn turning yellow with a caramel-like scent. Etymology: Cercidiphyllum is composed of classical Greek κέρκης (kerkis), a poplar tree, and φύλλων ( fyllon), a leaf. Cercis is the ancient name used by Theophrastus to describe the Judas tree, Cercis siliquastrum (Fabaceae); the leaves of Cercidiphyllum resemble those of Cercis.

162. DAPHNIPHYLLACEAE Laurel-leaf family

EUDICOTS

bracteate racemes of unisexual flowers. The small calyx is composed of three or six parts and sometimes persistent. Petals are absent. Male flowers have five to 12 (–18) stamens in one whorl, with short filaments and moonshaped anthers that open laterally along their length and have an exserted connective. Female flowers sometimes have five to ten staminodes, but these are sometimes absent. The bilocular, superior ovary has a short, two-branched style. The fruit is an ovoid or ellipsoid, tuberculate or rugose, often glaucous fleshy drupe with a hard stone. Distribution: The family is distributed in southern India, Sri Lanka, East and Southeast Asia, and from the Himalayas throughout Indochina and Malesia to the Solomon Islands. Phylogeny and evolution: Daphniphyllaceae have been difficult to place in the past, and associations have been made with a variety of families such as Balanopaceae, Euphorbiaceae and Hamamelidaceae. The last relationship is now confirmed by molecular data.

These are evergreen, unisexual trees and shrubs with alternate leaves that are often clustered or nearly verticillate at the apex of branches. The petiolate simple leaves lack stipules and are often leathery and glaucous beneath. Inflorescences are axillary, solitary

Daphniphyllum macropodum, male flowers, Royal Botanic Gardens, Kew, UK [162]

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Genera and species: The family has a single genus, Daphniphyllum, which has c. 30 species. Uses: The Ainu people of Japan and Siberia used to dry and smoke the leaves of Daphniphyllum humile like tobacco.

Daphniphyllum humile, female flowers, Crûg Farm Plants, Wales, UK [162]

Etymology: Daphniphyllum is composed of Greek δάφνη (dafne), a laurel bush, and φύλλων ( fyllon), a leaf, from the resemblance of the leaves to Laurus nobilis (Lauraceae).

163. ITEACEAE Sweetspire family

This family is composed of trees and shrubs that are sometimes lax climbers. They are usually evergreen, sometimes deciduous, and their leaves are alternate, simple with minute stipules and distinct petioles. Blades are usually pinnately veined and glandularserrate, rarely entire. Inf lorescences are many-flowered terminal or axillary racemes or panicles that are often in groups of two or three, or few-flowered corymb-like cymes. The bisexual flowers are actinomorphic. The five sepals are fused basally into a short, conical tube around the base of the ovary, forming a hypanthium. Five petals are placed between the sepals on the hypanthium and are persistent, triangular or clawed. The five

Daphniphyllum chartaceum, Kunming Botanical Garden, China [162]

SAXIFRAGALES

EUDICOTS

Itea rhamnoides, Moodie Mountains, South Africa (CD) [163]

Itea yunnanenis, Nichols Arboretum, Ann Arbor, Michigan, USA [163]

Ribes sanguineum, planted in a garden in Giethoorn, the Netherlands [164]

stamens are inserted on an annular nectar disc, with short filiform (Itea) or broad and apically toothed filaments (Pterostemon). The anthers are dorsifixed, open by lengthwise slits, and in Itea have a globular protrusion of the connective at the tip. The nearly superior or mostly inferior ovary is composed of two or five fused carpels and is bi- or pentalocular. It is topped with single fused (Pterostemon) or two fused or free (Itea) styles, with capitate, globular stigmas that are coherent in flower, separating in fruit (Itea) or the stigma shallowly five-lobed (Pterostemon). The often woody capsules have a persistent perianth and open by two or five valves. Distribution: Iteaceae are found in temperate eastern North America, southwestern North America (Oaxaca, Mexico), southeastern Africa and temperate and tropical East and

Itea virginica, Hortus botanicus, Leiden, the Netherlands [163]

Ribes roezlii, Royal Horticultural Society Show, London, UK [164]

Southeast Asia, from the Himalayas to China, Japan, Java and the Philippines.

164. GROSSULARIACEAE Gooseberry family

Phylogeny and evolution: The family is known from c. 90 million years old fossil flowers of the Late Cretaceous of North America and from Tertiary fossil pollen from Europe and North America. Iteaceae have been included in Grossulariaceae or Escalloniaceae, but molecular results place Itea sister to the Mexican genus Pterostemon in core Saxifragales. Genera and species: This bigeneric family has 21 species: Itea (18) and Pterostemon (3). Etymology: Itea (ιτέα) is Greek for willow, given to this genus on account of the rapid growth of some species and the glandular serrate leaf margins, similar to those of willows.

Bisexual, rarely unisexual shrubs, that are usually deciduous, sometimes (semi-) evergreen make up this family. They can be upright, creeping or laxly climbing and are frequently covered in simple or ternate spines. Their alternate, simple leaves are usually petiolate with dry, brown, fimbriate stipules. Leaf blades are usually trilobed or

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SAXIFRAGALES subpalmately lobed or cleft, and normally have toothed margins, rarely subentire. Venation is palmate, usually with three main veins. Inflorescences are terminal or axillary racemes, sometimes corymbs or flowers solitary. Flowers are usually bisexual (unisexual in R. diacanthum), and the perianth is placed on a floral tube that is wholly or partially fused with the ovary at the base. The five (rarely four) sepals are fused at the base and can be green or coloured and petallike. The five (rarely four or absent) petals are distinct, green or coloured. The five (rarely four) stamens are all fertile (rarely all staminodial in female f lowers), with filiform filaments that are free or fused to the hypanthium but distinct from each other. Anthers are basifixed and open by lengthwise slits. A well-developed five-lobed nectar disc surrounds the tip of the ovary. The partly or completely inferior ovary is composed of two fused carpels forming one locule and two free or fused styles on top, with two stigmatic papillose branches. The fruit is a soft berry crowned by a persistent perianth, often covered with hairs, prickles or glandular hairs. Distribution: The family is widely distributed in the temperate zones of the Northern Hemisphere, in North America, south to the mountains in Central America and the Andes in South America, throughout temperate, boreal and montane Europe, the Atlas Mountains in North Africa and temperate and montane northern and East Asia. They jump into the Southern Hemisphere in southern South America. Phylogeny and evolution: Grossulariaceae have in the past been part of a broadly circumscribed and heterogeneous Saxifragaceae or Grossulariaceae s.l. (members of which are now placed in Celastraceae, Escalloniaceae, Iteaceae, Montiniaceae, Phyllonomaceae and Tetracarpaeaceae). A relationship with other woody Saxifragales has been assumed, but with no support phytochemically or embryologically. Molecular analyses place Grossulariaceae as sister to herbaceous Saxifragaceae s.s., which is supported by many morphological characters.

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EUDICOTS

Genera and species: The sole genus in Grossulariaceae is Ribes with c. 150 species. Uses: Many species of Ribes are cultivated for their fruits, especially blackcurrant (R. nigrum), redcurrant (R. rubrum etc.) and gooseberry (R. uva-crispa); superior cultivars and hybrids between these and other Ribes species are now generally grown, rather than the wild types. Sour when fresh, the fruits are widely used in cooking and for juice and jam. Redcurrants are rich in pectin and the juice is sometimes mixed with other fruit to ensure a good set in jam. They can also be made into cordial (e.g. Ribena) and syrups, and are an important ingredient of rødgrød, a kissel-like dessert from Denmark. Minor fruit crops of Ribes include whitecurrant (an albino form of redcurrant), greencurrant (a Finnish form of blackcurrant), American blackcurrant (R. americanum), granite gooseberry (R. curvatum), prickly gooseberry (R. cynosbati), Worcesterberry or Western black gooseberry (R. divaricatum), jostaberry (a complex hybrid between R. divaricatum, R. nigrum and R. uva-crispa) and golden currant (R. odoratum). Some species such as R. grossularoides, R. sanguineum and R. speciosum are solely grown as garden ornamentals and their fruit is inedible. Etymology: A grossulus is a small unripe fig in classical Latin, a word later applied in old French as groseille, although the etymology is also probably tied in with Frankish krusil, Old Dutch kroesel and German kruselbeere, all meaning a wrinkled berry, possibly all from the same Indo-European root. Grossularia is a later synonym of Ribes, which is a Latinised form of Semitic ribas, acid-tasting.

165. SAXIFRAGACEAE Saxifrage family

These are usually perennial herbs, rarely annual or biennial, and often have perennating rhizomes. Leaves are usually formed in alternate basal rosettes, sometimes along the inflorescence stems and then also alternate or sometimes opposite; they are usually simple, rarely pinnately or palmately compound, and the margin can be entire or deeply lobed or cleft, crenate or dentate. Leaves are usually petiolate with stipules. Inflorescences are bracteate racemes or cymes. Bisexual or unisexual f lowers are actinomorphic or sometimes zygomorphic, and the perianth is placed on a hypanthium that can be free or partly fused with the ovary (producing a semi-inferior ovary). The sepals are usually five, sometimes three or up to ten, and fused with the hypanthium. The usually five (sometimes four, six or absent) petals are clawed, sometimes cleft at the tip or finely dissected. The five or ten stamens are free and have basifixed anthers that open by lengthwise slits. The inferior or semi-inferior ovary is composed of two, sometimes three, carpels that are fused at the base, and each carpel is topped with a stylodium and capitate stigma. The capsular or follicular fruit typically has numerous small seeds. Distribution: Saxifragaceae are widespread across the Northern Hemisphere, in North America south to central Mexico, throughout Europe and temperate and subtropical Asia, and North Africa, the Andes and the mountains of Ethiopia, Luzon and New Guinea. Phylogeny and evolution: The age of Saxifragaceae has been estimated to be between 54 and 38 million years old, but fossil data are inconclusive, since pollen records for the family may also refer to the similar Hydrangeaceae or other families now excluded from Saxifragaceae. A 90-million year old Late Cretaceous fossil, Tylerianthus, from North America may be a member of the stem lineage of Saxifragaceae, although the authors of that paper indicated it was related to both Hydrangeaceae and Saxifragaceae, which at that time were still thought to be closely related. Nevertheless, the fossil appears to be related to Saxifragaceae s.s.

SAXIFRAGALES

EUDICOTS

Astilbe thunbergii, Royal Botanic Gardens, Kew, UK [165]

Bergenia crassifolia var. pacifica, Ruissalo Botanical Garden, Turku, Finland [165]

Saxifragaceae used to include a great number of unrelated taxa, such as Parnassia (now in Celastraceae), Escalloniaceae, Hydrangeaceae and Vahliaceae. Mitella has been found to be polyphyletic, representing several distinct lineages and Saxifraga is paraphyletic. The taxonomy has not yet been resolved. Micranthes used to be part of Saxifraga, but it does not fall in the clade that includes typical Saxifraga. Generic recircumscription is greatly needed.

Saxifraga aizoides, above Wasdale Head, Lake District, UK [165]

in the temperate zones, especially Astilbe (false buck’s-beard), Bergenia (elephantears), Darmera (Indian rhubarb), Heuchera (coralbells), Mukdenia, Rodgersia, Saxifraga (saxifrage, londonpride), Tellima (fringecups) and Tiarella; some are houseplants such as the piggyback plant (Tolmiea menziesii) and mother-of-thousands (Saxifraga stolonifera).

Genera and species: The family has 35 genera and c. 640 species: Astilbe (25), Astilboides (1), Bensoniella (1), Bergenia (10), Bolandra (2), Boykinia (6), Brachycaulos (1), Cascadia (1), Chrysosplenium (c. 55), Conimitella (1), Darmera (1), Elmera (1), Heuchera (c. 35), Hieronymusia (1), Jepsonia (3), Leptarrhena (1), Lithophragma (10), Micranthes (c. 70), Mitella (20), Mukdenia (1), Oresitrophe (1), Peltoboykinia (1), Rodgersia (5), Saniculiophyllum (1), Saxifraga (c. 370), Saxifragodes (1), Saxifragopsis (1), Suksdorfia (2), Sullivantia (3), Tanakaea (1), Telesonix (2), Tellima (1), Tetilla (1), Tiarella (3) and Tolmiea (2).

Carnivory: Carnivory of plants with glandular hairs was studied by Charles Darwin, who observed that the glands of meadow saxifrage (Saxifraga rotundifolia) and Pyrenean saxifrage (S. umbrosa) were able to absorb a range of substances. Some saxifrages have glands that are capable of trapping insects, and these same glands have absorptive properties. The exudates from the glands have not been demonstrated to have digestive properties, and it is unlikely that these plants are true carnivores because their habitat preferences do not appear to be those associated with the classic cases of carnivory. However, there is always a chance that an insect is caught and products of its decaying body could be absorbed by the plant via the glands or the roots.

Uses: The leaves of Chrysosplenium oppositifolium (golden saxifrage) and Saxifraga nelsoniana (Nelson’s saxifrage) are eaten fresh as salad greens. The rhizome of Astilboides tabularis has been used to make wine and produces tannin. Several genera include highly valued ornamentals

Etymology: Saxifraga is derived from Latin saxum, a rock, and frango, to break, and two possible reasons for the name have been suggested. Some species grow in rock crevices and appear to break rocks, but alternatively it may refer to the former medicinal use of breaking up kidney stones.

Chrysosplenium davidii, private garden, Kingston upon Thames, Surrey, UK [165]

166. CRASSULACEAE Stonecrop family

These terrestrial, perennial and annual herbs, sometimes shrubs, rarely tree-like, aquatic, scandent or epiphytic, are always more or less succulent. Their stems are succulent, may form rhizomes, underground stems or corms and often have bulbils along the stem or on leaf margins. Many grow easily into new plants from vegetative parts that have fallen off the parent plant. Leaves are alternate or opposite, sessile or petiolate, without stipules, and can be placed along the stem or spirally crowded at the tips of stems, or in a basal rosette. The simple or rarely pinnately divided blades are usually entire or broadly lobed, sometimes dentate or more deeply incised and flat or round in crosssection. Inflorescences are usually terminal and most often many-flowered thyrses composed of cymes, rarely true panicles, racemes or spikes, often many branched and bracteate. The bisexual, rarely unisexual, flowers are actinomorphic or rarely zygomorphic and usually pentamerous, although variation in numbers of flower parts is great. The four to 20 sepals are

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SAXIFRAGALES

EUDICOTS

Sedum caeruleum, Sicily, Italy [166]

Graptopetalum paraguayense, National Botanic Garden of Ireland, Glasnevin [166]

Aichryson laxum, Royal Botanic Gardens, Kew, UK [166]

Kalanchoë fedtschenkoi, The Living Desert, Palm Desert, California, USA [166]

free or fused at the base and sometimes clearly unequal in size. The (3–)4–20(–32) petals are free or fused and sometimes form a tube. Stamens are as many as or twice the number of petals, the filaments free or fused with the corolla tube. Anthers are basifixed and open lengthwise. The superior (to partially inferior) ovary has as many carpels as petals, the carpels usually free or nearly so and each forming a single locule, at the base with a nectary scale. Fruits are follicles that open along the carpel suture and are usually many-seeded. Distribution: The family occurs worldwide, but is absent from some areas. It occurs in North America from the Arctic to tropical Mesoamerica and the Caribbean, in western and southeastern South America, across Eurasia south to the Himalayas and southern China, north and Sub-Saharan Africa,

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Crassula ovata, private garden, San Francisco, USA [166]

Madagascar, southern Arabia, Luzon, southern Australia and New Zealand and on Antarctic islands. There are particular areas with species radiations in the Macaronesian Islands and southern Africa. Phylogeny and evolution: Crassulaceae are estimated to have originated in East Africa between 60 and 100 million years ago, and some authors place the age more specifically around 70 million years. There is a deep split in the family between Crassuloideae and the rest. The taxonomy of Crassulaceae is complicated because morphological characters used to describe genera in the past do not seem to follow clades, and moreover species readily hybridise across genera in the wild and in cultivation, making identification contentious and classification problematic. Sedum is widely polyphyletic even when

Umbilicus rupestris, South Wales, UK [166]

segregate genera are accepted, and perhaps the best solution in the future may be to unite the entire Sempervivoideae into a single genus or at least merge all of tribe Sedeae into Sedum s.l. (including Dudleya, Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Prometheum, Rosularia, Sedella, Thompsonella and Villadia), all Telephiae in Orostachys (including Sinocrassula, Kungia, Meterostachys and Hylotelephium) and expand Aeonium and Sempervivum to include a couple of Sedum species placed there by molecular data. This may, however, cause problems in recognising these genera morphologically and thus a broad Sedum may be a better option, especially because other genera such as Orostachys and Umbilicus are also polyphyletic in the traditional sense. Regardless, a revised generic reclassification and new combinations will be needed.

SAXIFRAGALES

EUDICOTS

Genera and species: Crassulaceae include 7–35 genera (depending on the status of Sedum) and c. 1,400 species in three subfamilies: Crassuloideae – Crassula (c. 195) and Hypagophytum (1); Kalanchoideae – Andromischus (c. 26), Cotyledon (11), Kalanchoë (c. 144) and Tylecodon (46); Sempervivoideae, which are probably better united into the single genus Sedum (then with c. 979 species and to be included in Kalanchoideae), otherwise – Aeonium (c. 36), Afrovivella (1), Aichryson (14), Chiastophyllum (1), Dudleya (c. 47), Echeveria (c. 139), Graptopetalum (18), Hylotelephium (c. 27), Kungia (2), Lenophyllum (7), Meterostachys (1), Monanthes (9), Orostachys (11), Pachyphytum (15), Perrierosedum (1), Petrosedum (7), Phedimus (13), Pistorinia (3), Prometheum (8), Pseudosedum (12), Rhodiola (c. 58), Rosularia (17), Sedella (3), Sedum (c. 420), Sempervivum (c. 63), Sinocrassula (7), Thompsonella (6), Umbilicus (12) and Villadia (21). Uses: Wall pepper, Sedum acre, has a peppery taste and was used in Europe as a spice and is excellent added to salads. Trip-madame, Petrosedum rupestrum, has also been used as a salad green and pot-herb. Many species were used as a cure for burns and probably for a similar reason common houseleek, Sempervivum tectorum, was planted on roofs as a magical protection against thunder and lightning. A tincture of the roots of roseroot, Rhodiola rosea, was used in earlier times to elevate the spirits. It has proved to be medically effective in treating depression and bad moods. Many species of Crassulaceae are cultivated as garden or rockery ornamentals. The jade or money plant (Crassula ovata) and Kalanchoë blossfeldiana are especially popular pot plants around the world and are produced in large quantities. CAM photosynthesis: The family is the source of the name of a particular pathway of photosynthesis, crassulacean acid metabolism (CAM), which was first discovered in this family. CAM photosynthesis evolved independently in many clades (and all members of Crassulaceae have it) as an adaptation to arid environments. In CAM plants the leaf guard

cells remain shut during the day to reduce water loss. At night, they open to absorb carbon dioxide, which is stored as malate and used as a substrate for photosynthesis during the day. Etymology: Crassula is a diminutive form of Latin crassus, thick, referring to the fleshy leaves.

167. APHANOPETALACEAE Gum-vine family

petals and stamens to form a short floral tube (hypanthium). The four petals are minute but can be absent even in the same individual. Eight stamens have nearly basifixed anthers that open by lengthwise slits. The semiinferior ovary is composed of four laterally fused carpels into a tetralocular, four-ribbed pistil topped by a four-lobed style. The fruit is a hard nut with persistent enlarged sepals. Distribution: This family is restricted to Australia; in Queensland and New South Wales it is found in riparian scrub and in Western Australia it is found locally in limestone crevices. Phylogeny and evolution: On morphological grounds the placement of Aphanopetalum has been difficult. It was previously placed in Cunoniaceae, Iteaceae and Saxifragaceae, but does not agree with several characters of those families. Molecular evidence places this family near Crassulaceae in Saxifragales, and the clade is estimated to be c. 60 million years old.

A family of scrambling and climbing shrubs, these plants have stems with prominent lenticels. Leaves are opposite, simple and shortly petiolate. They lack stipules but have minute toothed structures around the internodes. Blades are serrate to almost entire and have pinnate venation. Inflorescences are axillary lax panicles enclosed by two prophylls. The actinomorphic flowers are bisexual and have four persistent white sepals and no or reduced petals. The base of the sepals is fused with the basal portion of

Etymology: Aphanopetalum is composed from Greek αφανώς (afanos), inconspicuous, and πετάλων (petalon), a petal.

Aphanopetalum resinosum, Mt Cordeaux, Queensland, Australia (JC) [167]

Aphanopetalum resinosum, Mt Cordeaux, Queensland, Australia (JC) [167]

Genera and species: The sole genus in this family is Aphanopetalum with two species: A. clematideum and A. resinosum.

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SAXIFRAGALES

Tetracarpaea tasmanica, Mt Field National Park, Tasmania, Australia (CD) [168]

EUDICOTS

Penthorum sedoides, private garden, Kingston upon Thames, Surrey, UK [169]

168. TETRACARPAEACEAE Delicate-laurel family

Haloragodendron glandulosum, Australian National Botanic Garden, Canberra [170]

Phylogeny and evolution: The placement of Tetracarpaeaceae has long been contentious, and this species has been suggested to belong in Cunoniaceae, Escalloniaceae and Saxifragaceae. Molecular evidence has placed it as sister to Haloragaceae and Penthoraceae. Genera and species: The only species in this family is Tetracarpaea tasmanica.

Low, bushy shrubs with upright branches make up this family. Their alternate, petiolate leaves lack stipules, and the blades are simple, crenate or serrate and have pinnate venation with secondary veins terminating near the margin. Inflorescences are terminal, erect, bracteate racemes with small, bisexual, actinomorphic flowers. The four sepals are free and persistent, the four petals are free, spreading, clawed at the base and spathulate. The four or eight stamens have free filaments and basifixed anthers. The superior ovary is composed of four (rarely five) free, stalked carpels with sessile stigmas. Fruits are follicles that open along the carpel suture exposing numerous seeds. Distribution: This family is endemic to Tasmania, where it is restricted to subalpine habitats.

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Etymology: Tetracarpaea is composed of Greek τετρα (tetra), four, and καρπός (carpos), a fruit, in reference to the four-carpellate ovaries.

169. PENTHORACEAE Ditch-stonecrop family

Glischrocaryon aureum, Mt Benia, Western Australia [170]

sharply serrate or doubly serrate margins. Terminal inflorescences are scorpioid racemes with actinomorphic, bisexual flowers that have a hypanthium to which sepals and petals are attached. The usually five (sometimes up to eight) sepals are fused, and one to eight petals are free or absent. The ten stamens are free from each other but are placed on the hypanthium rim, appearing fused to the ovary. The basifixed anthers open by lengthwise slits, and there are no nectaries at the base of the stamens. The superior or semi-inferior ovary is composed of usually five (sometimes four or up to eight) carpels that are fused basally and laterally and are partially fused to the hypanthium, and each carpel is tipped with a distinct short style and capitate stigma. The fruit is a circumscissile capsule that opens at the top, containing up to 400 seeds. Distribution: This family is distributed in eastern North America and temperate East Asia.

This family consists of perennial herbs with simple or branched stems. Cauline leaves are alternate and lack stipules. Petioles are sometimes present, and the simple blades have

Phylogeny and evolution: Placement of Penthorum has been disputed extensively in the past, as it was sometimes considered it to be transitional form between Crassulaceae and Saxifragaceae. Classification of the genus based on morphological, anatomical or embryological traits did not lead to agreement,

SAXIFRAGALES

EUDICOTS

but molecular studies indicated that the genus is sister to Haloragaceae. Even though the stem group of Penthoraceae has been dated to be c. 45 million years old, the split between the North American and East Asian species happened less than 6.5 million years ago. Genera and species: The sole genus is Penthorum with two species, P. chinense and P. sedoides. Uses: All parts of Penthorum are edible after cooking, but overconsumption may function as a laxative or respiratory sedative. Etymology: Penthorum is derived from Greek πέντε (pente), five, in reference to the five-parted fruits.

170. HALORAGACEAE Water-milfoil family

This is a family composed of small trees, shrubs and perennial and annual terrestrial and aquatic herbs. Stems are upright or creeping and rooting at the lower nodes. Stems can be condensed to form turions for overwintering in aquatic species, and sometimes filiform appendages are found in leaf axils and elsewhere along the stem. Leaves are opposite or in whorls (verticillate), without stipules and sessile or petiolate. Blades are simple or deeply pinnately dissected, the margin entire or toothed, and heterophylly occurs in the aquatic, partially submerged taxa. Inflorescences are compound thyrses, panicles or spikes, or the flowers can be solitary in the leaf axils. The bisexual or unisexual flowers are actinomorphic and dimerous. The two or four sepals are fused to the ovary (absent in female flowers of Myriophyllum), and the usually two or four, rarely eight, petals are free,

keeled or hooded (absent in female flowers of some taxa). There are two, four or eight stamens with short filaments and basifixed anthers that open by slits. The inferior ovary is composed of four carpels, and each is topped with a free style and a capitate, sometimes fimbriate stigma. The fruit is usually a one- to four-seeded nutlet or capsule that splits into two or four parts, the outside of which is often winged, ribbed or tuberculate. Distribution: The family has a nearly global distribution and occurs in North America, most of South America, Sub-Saharan Africa, Eurasia, Malesia, Australia, New Zealand and the Pacific. Phylogeny and evolution: Previously thought to be part of Myrtales, where they do not fit well morphologically, molecular analyses have now placed them in Saxifragales, sister to Penthoraceae. The earliest fossils, known from the Upper Cretaceous, are fossil fruits from Mexico and fossil pollen from Europe. Tertiary pollen is common in Europe, North America and New Zealand. Proserpinaca macro-fossils are known from Pliocene Europe and Asia where it is no longer found. Molecular results also show that Gonocarpus and Haloragis are not monophyletic in their traditional circumscription, resulting in the acceptance of Trihaloragis and Meionectes, respectively.

Cynomorium coccineum, Morocco (HR) [171]

Genera and species: This is a family of 11 genera and c. 145 species: Glischrocaryon (5), Gonocarpus (40), Haloragis (26), Haloragodendron (5), Laurembergia (4), Meionectes (3), Meziella (1), Myriophyllum (c. 60), Proserpinaca (2), Vinkia (1) and Trihaloragis (1). Uses: Some species of Myriophyllum are cultivated as aquarium or pond plants and may cause havoc when they escape and naturalise in watercourses. Etymology: Haloragis is the Latinised form of the Greek άλας (halas), salt, and ρόγες (rhoges) grapes or berries.

171. CYNOMORIACEAE Tarthuth family

These perennial, bisexual herbs are parasites on other plants and lack chlorophyll. The entire plant has a reddish-brown colour. The roots are fleshy, forming a rhizome that is attached to the host, and the stem is usually simple,

Cynomorium coccineum, with fly pollinator, Morocco (HR) [171]

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SAXIFRAGALES rarely branched. The scale-like brown leaves are arranged spirally along the stem, which is topped with a terminal spadix of flowers. The four to six (sometimes up to eight) tepals are usually fused basally, rarely free. Flowers are unisexual, and male flowers have a single stamen fused to the perianth with the ovary reduced to a nectary. Female flowers are composed of an inferior uniloculate ovary, topped by a single long, channelled style. Fruits are nutlets. Plants parasitise the roots of various hosts, mainly Amaranthaceae, Cistaceae, Nitrariaceae and Tamaricaceae. Distribution: Cynomoriaceae can be found in deserts and arid regions around the Mediterranean region, from Lanzarote in the Canary Islands and Mauritania in North Africa through to Egypt, south into

EUDICOTS

Phylogeny and evolution: Previously Cynomoriaceae were thought to belong to Balanophoraceae, another family of holoparasites, but there are several morphological differences. Molecular analyses place this family in Saxifragales, albeit without a well-supported placement among those families. Other analyses have placed them in other orders (e.g. Rosales), so the correct relationships and taxonomic placement is still a matter of debate.

Uses: Cynomorium coccineum, known in Arabic as tarthuth, or in English as Maltese mushroom, grows in the desert after rains and has been harvested for thousands of years for food and medicine and as a dye for fabric. Bedouins clean the freshly picked spikes, peel off the outer skin and eat the flavourful white interior, which is sweet tasting. It is edible raw, with a pleasant crisp, succulent texture. In the ‘doctrine of signatures’, it was believed that the phallic shape of the inflorescences would mean that it is a cure for erectile dysfunction and other sexual health issues. It has been used for these purposes throughout its range, probably to no avail.

Genera and species: The sole genus is Cynomorium with two species: C. coccineum and C. songaricum.

Etymology: Cynomorium is composed of classical Greek κυνός (kynos), a dog, and μοριων (morion), a penis.

the Arabian Peninsula to Oman and north to Malta, Greece and Iraq, and in Central Asia to Mongolia and north-central China.

VITALES Relationships of this order have long been uncertain, but it is now known that they are sister to all rosids. They include a single family.

172. VITACEAE Grapevine family

Vitaceae are a family of bisexual and unisexual woody climbers, vines, shrubs and small trees. Stems often have lenticels, and the bark may be shedding, with branches often swollen at the nodes, usually unarmed or with rows of prickles. Tendrils are present or not, but can be simple, forked or multiplebranched, sometimes with adhesive discs

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at the tips of each branch (Parthenocissus), usually placed opposite the leaves. Stipules are present, normally two and interpetiolar or sheathing the petiole margins with stipular wings (Leea). Leaves are petiolate, simple, lobed or palmately compound, trifoliate or one to three times pinnate, alternate, but usually opposite tendrils or inflorescences, often with structures called ‘pearl glands’ on the leaf margins or blade surfaces. Inflorescences are panicles, corymbs or rarely spikes, usually composed of cymes, usually leaf-opposed, or axillary (Cayratia, Tetrastigma). The actinomorphic, unisexual or bisexual flowers have a fused often cupulate calyx with four or five (or up to seven) teeth or lobes, glandtipped in Leea. The four, five or up to seven (Rhoicissus) petals are free or basally fused or sometimes fused to form a calyptrum (Vitis). Stamens are as many as and placed opposite

the petals, and anthers are nearly basifixed and open by lengthwise slits. The floral nectary disc is placed inside the stamens and is ring-shaped, cupular, tubular or glandular. The superior ovary is composed of two or three fused carpels making two or four to six locules, which are usually topped with a short or elongate style and a capitate, discoid or four-lobed stigma. The fruit is a one- to four-seeded, juicy or dryish berry. Distribution: Vitaceae are distributed across the warm temperate and tropical regions of the world. They are found in North America north to the Great Lakes and New England south through Central and South America to Tierra del Fuego. They occur from the Sahel region to southern Africa, in Madagascar, Yemen, Italy and southeastern Europe, the Caucasus, northern Iran to Central Asia, India, Southeast

VITALES

EUDICOTS

Leea coccinea, Brooklyn Botanic Garden, New York, USA [172]

Parthenocissus vitacea, New Mexico, USA (DZ)

Clematicissus angustissima, Western Australia [172]

Vitis vinifera, Transylvania, Romania [172]

and East Asia, north to Sachalin and the Russian Far East, and south to Australia and northern New Zealand.

Eocene and Oligocene from the Neotropics, where the genus is now absent.

Vitis species, such as bush grape (V. acerifolia), canyon grape (V. arizonica), skunk grape (V. labrusca), mountain grape (V. monticola), cat grape (V. palmata), river grape (V. riparia), muscadine (V. rotundifolia), sand grape (V. rupestris), frost grape (V. vulpina) etc., many of which are locally eaten or made into local wines, liqueurs or jellies. Several wild species have been used for hybridising with common grapes, for different flavours, vigour, climatic and disease tolerance. Cream of tartar (potassium hydrogen tartrate) crystallises out of wine on the sides of barrels and bottles. It is a component of most baking powder. Leaves of some Cissus species are edible, and C. gongylodes is commonly cultivated and harvested by local tribes in South America. Cayratia geniculata is used to make ropes. Some species are cultivated as ornamentals, such as Ampelopsis, Cissus, Leea, Parthenocissus, Rhoicissus and Vitis coignetiae. Tetrastigma is the host plant for the endoparasitic Rafflesia (Rafflesiaceae), the genus with the largest flowers.

[172]

Phylogeny and evolution: Vitaceae are the sole family in Vitales, an order often positioned as sister to all other rosids. Relationships of Vitaceae have long been considered uncertain, and the family was often associated with Rhamnaceae or even Proteaceae because all have stamens opposite the petals. Leeaceae have in all analyses been found to be sister to Vitaceae s.s. and because the two share numerous characters they are now merged. Generic delimitation in Vitaceae will need some attention by future taxonomists because Cissus appears to be polyphyletic and Tetrastigma is embedded in Cayratia. Pterisanthes has nomenclatural priority over Ampelocissus, but combinations have not been made. Late Cretaceous fossil leaves have been assigned to Vitaceae. The distinctive seeds of Vitaceae are found frequently in Tertiary deposits in North America and Europe. Ampelocissus fossils are known from Mesoamerican and Peruvian Oligocene sediments (c. 30–28 million years old). Leea fossils are known from the Late

Genera and species: Vitaceae include 14 genera and about 910 species in two subfamilies; Leeoideae – Leea (34); Vitoideae – Acareosperma (1), Ampelocissus (c. 100), Ampelopsis (25), Cayratia (c. 65), Cissus (c. 350), Clematicissus (1), Cyphostemma (c. 150), Nothocissus (4), Parthenocissus (10), Pterocissus (1), Rhoicissus (10), Tetrastigma (c. 95) and Vitis (c. 65). Uses: Grapes (Vitis vinifera) can be eaten fresh or dried (raisins, currants), made into jellies or fermented into wine. The leaves are also edible, often stuffed with meat, rice and vegetables. Traditionally a Mediterranean crop, grapes have been widely cultivated, especially in France, Spain, Germany, Austrlia, Greece, Crimea, California, Chile, South Africa, Australia, New Zealand and China, and recent vineyards can be found as far north as Michigan, England and Sweden, this hardiness probably due to hybridisation of French varieties with Vitis riparia. Many other species produce edible fruits, especially Ampelocissus, Cayratia, Tetrastigma harmandii and other

Etymology: Vitis is Latin for a grape vine. It is derived from Proto-Indo-European weytis, referring to something that twines, winds, bends or turns.

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ZYGOPHYLLALES

EUDICOTS

ZYGOPHYLLALES Families 173 and 174 form the order Zygophyllales, a relatively isolated order of two closely related families. The lineage may have evolved c. 108 million years ago, whereas the families diverged somewhere between 70 and 64 million years ago. Zygophyllales are sister to the rest of the rosid I (fabid) clade.

173. KRAMERIACEAE Ratany family

This is a family of parasitic shrubs and perennial herbs with rhizomes and haustorial roots that parasitise a variety of other flowering plants. They have green, alternate, spirally arranged leaves without stipules. Leaf blades can be sessile or petiolate and simple or trifoliate with an entire margin and reticulate venation. Inflorescences are terminal racemes, panicles or axillary single flowers. The bisexual zygomorphic flowers have (four or) five free, showy, colourful sepals. There are (four or) five petals of which two are reduced to glandular structures secreting oil, and the other three variously clawed or lanceolate, clustered on one side of the ovary. The four (rarely three) stamens

have stout filaments that are fused at the base. Anthers are tetralocular, tubular and the same diameter as the filaments and open by membranous pores. The superior ovary is composed of two carpels, of which one is suppressed to form a single locule topped by a style and small stigma. Fruits are globose or slightly flattened, indehiscent, spiny capsules with a single globose seed. Distribution: This family can be found in North America, from the southwestern North America to Florida, through Mesoamerica, the Lesser Antilles and dry regions in South America. Phylogeny and evolution: The highly specialised genus Krameria is the only member of the family. Its affinities have historically been uncertain, and it has been placed variously in Fabaceae and Polygalaceae; more recently it has been accorded familial status, and it was demonstrated that it is the sister to Zygophyllaceae, with which it shares few obvious synapomorphies. Krameriaceae grow in the same habitats as many Zygophyllaceae and have the ability to withstand similarly dry conditions. However, they are unlike

Krameria bicolor, fruit, Andreas Canyon, Palm Springs, California, USA [173]

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Zygophyllaceae in many other respects, notably in having extremely zygomorphic flowers and being root parasites, including some that parasitise species of Larrea (Zygophyllaceae). All species are pollinated by only female oil-collecting solitary bees of the genus Centris (Apidae). Genera and species: The sole genus in this family is Krameria with 18 species. Uses: Due to its astringent properties, ratany (Krameria triandra) has minor economic importance as a medicinal plant, especially for enteritis and angina, but treatment has been proven to be ineffective. It is most commonly used in dental products such as mouthwash and herbal toothpaste, helping to heal gum irritations and minor infection. It is occasionally used as a dye and as an ingredient in cosmetics. Etymology: Krameria is named for Wilhelm Heinrich Kramer (died 1765), a German naturalist and physician, who described the plants and animals of Lower Austria in 1756, which was one of the first works to apply the Linnaean binomial system.

Krameria bicolor, Andreas Canyon, Palm Springs, California, USA [173]

ZYGOPHYLLALES

EUDICOTS

174. ZYGOPHYLLACEAE Twinleaf family

This family of trees, shrubs, perennial and annual herbs often have jointed branches with swollen nodes that are sometimes armed with axillary or stipular thorns. Opposite, rarely alternate, leaves have stipules that are free or fused across the node. Blades are usually bifoliate, trifoliate or pinnate, rarely simple, and leaflets are usually asymmetrical and have an entire margin, usually flat, sometimes terete, often fleshy. Inflorescences are few-flowered cymes that are usually reduced to solitary or paired flowers. The bisexual, actinomorphic to zygomorphic flowers have four to six free or basally fused sepals. The four to six (rarely absent) petals are free and are often clawed. A nectary disc is often present between petals and stamens. Stamens are as many as or twice as many as petals. Filaments sometimes have scales or appendages at their base, anthers are tetralocular, dorsifixed and dehiscent via a Kallstroemia grandiflora, New Mexico, USA (DZ) [174]

Zygophyllum fabago, Ani, Turkey [174]

lengthwise slit. The superior ovary is usually tetra- to pentalocular, or sometimes has fewer (two or three) or up to 12 locules. The style is apical or gynobasic (Zygophyllum), filiform or subulate, and stigmas are capitate, clavate or somewhat lobed or ridged. Fruits are capsules, splitting into mericarps that may be winged, lobed, angled, spined, tuberculate or, in Balanites, a drupe. Distribution: Zygophyllaceae can be found in dry and warm-temperate, tropical to temperate areas of the Americas (Minnesota to Patagonia), Mediterranean Europe, Africa, Madagascar, western, southern, central and eastern Asia, Malesia and Australia, mostly in arid or saline habitats. Phylogeny and evolution: Nitraria, Peganum and Tetradiclis were originally included in Zygophyllaceae, but molecular studies have revealed the relationships of these genera to be among other eurosid families. They are now placed in their own family Nitrariaceae (Sapindales). Balanites has a distinctive morphology with alternate leaves and a calyx that envelops the flower when young. It was previously placed in its own family, but molecular evidence places it close to Zygophyllum. Five subfamilies are sometimes recognised: Larreoideae, Morkilioideae, Seetzenioideae, Tribuloideae and Zygophylloideae. Larrea tridentata, Andreas Canyon, Palm Springs, California, USA [174]

Genera and species: This is a family of 22 genera and c. 220 species: Augea (1), Balanites (9), Bulnesia (8), Fagonia (c. 30), Guaiacum (6), Kallstroemia (17), Kelleronia (3), Larrea (6), Metharme (1), Morkillia (2), Neoluederitzia (1), Pintoa (1), Plectrocarpa (3), Porlieria (6), Seetzenia (1), Sericodes (1), Sisyndite (1), Tetraena (40), Tribulopsis (5), Tribulus (c. 25), Viscainoa (1) and Zygophyllum (c. 50). Uses: Lignum vitae (Guaiacum) is the hardest wood of all. Resin from Guaiacum sanctum has been used medicinally, especially in the 16th century to treat syphilis, and its populations were nearly exterminated for that purpose. This species is now listed in the CITES appendices. It is currently used as an antioxidant food additive (E314). Oil of guaiac is derived from Bulnesia sarmientoi, which is a fragrance used in soap. Retamo wax, harvested from B. retama, is used in cosmetics and polishing. Balanites fruits have high nutritional value (called ‘desert dates’), and their oil is also made into soap. Zygophyllum and Larrea flower buds have been used as caper substitutes. Etymology: Zygophyllum is derived from the Greek ζυγόν (zygon), yoke, and φύλλων ( fyllon), leaf, in reference to each leaf having two leaflets. Balanites aegyptiaca, Kenya [174]

Zygophyllum fruticulosum, Western Australia [174]

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FABALES Families 175 to 178 comprise the order Fabales, which includes nearly ten percent of eudicot diversity. The bulk of this is made up of Fabaceae, a dominant family in many ecosystems around the world. The order is well supported, although relationships among the families in the order are still disputed. The crown group is dated between 87 and 72 million years old.

175. QUILLAJACEAE Soapbark-tree family

These evergreen trees have saponaceous bark. Alternate, simple leaves are leathery and petiolate. Stipules are small and fall off early, and leaf blades are entire with pinnate venation. Inf lorescences are terminal or axillary clusters of a few flowers, the terminal ones bisexual, the lateral ones male; pedicels have prophylls. The five sepals are free and valvate, and the five petals are spathulate and white or cream. The nectary disc is thick and showy; it lines the receptacle and has five lobes fused to the sepals. The ten stamens

are in two whorls, one inserted near the tips of the disc lobes on the sepals, the other around the base of the ovary. Filaments are subulate, and the anthers have two thecae that open by lengthwise slits. The superior ovary is composed of five carpels that are basally fused, each with a terminal stylodium. Fruits are star-shaped follicles that open on the upper side, each carpel into two lobes exposing longwinged seeds. Distribution: This family is almost entirely restricted to the northern Andes in Chile with a disjunct population in northern Argentina, Uruguay and southeastern Brazil. Phylogeny and evolution: Despite uncertain relationships among families in Fabales, Quillajaceae are generally found to be sister to the rest. Until recently Quillaja was included in Rosaceae without question because it shares many characters with certain genera in that family (e.g. Kageneckia). Apart from molecular evidence showing that it is not

Quillaja saponaria, University of Talca Botanical Garden, Chile (CD) [175]

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related to Rosaceae, it has a base chromosome number of 14 and scalariform perforation plates, which do not occur in Rosaceae. Genera and species: The sole genus in the family, Quillaja, includes three species. Uses: The inner bark of the soapbark tree (Quillaja saponaria) contains saponins, which can be used as soap. The Andean peoples used it also for treatment of various chest infections, the saponin helping to produce more mucus in the airways and eliciting coughing, thus removing phlegm. It is also used as a humectant in various processed foods and foaming agent in fizzy drinks, whipping cream and fire extinguishers. It can also be used for the preparation of veterinary vaccines. Etymology: Quillaja is a Latinised form of the Araucani name for the plant, quillai (or killai). The scientific name is thus to be pronounced as ‘killaya’ in both English and Spanish.

Quillaja saponaria, fruit, San Francisco Botanical Garden, USA [175]

FABALES

EUDICOTS

are actinomorphic or slightly or completely zygomorphic. They usually have five (sometimes three or six) sepals that are free or fused into a tube, sometimes bilabiate, rarely reduced or vestigial. Petals are generally as many as the sepals (usually five), seldom fewer or none, and are distinct and often (Faboideae) highly differentiated into a papilionaceous corolla: an outermost upper petal (standard); innermost in Cercis and related to former Caesalpinoideae, two lateral petals that are more or less parallel to each other (wings) and the lower two innermost petals usually fused by their lower margins and forming a keel. Stamens are mostly ten, sometimes fewer or more numerous (especially in Mimosoideae), free or often fused by their filaments to form a closed or open sheath. The bilocular anthers open lengthwise or by pores and are uniform or dimorphic and then alternately basifixed and dorsifixed. The superior ovary is nearly always composed of a solitary carpel (rarely two or more free carpels), and it is unilocular or chambered (septate). The fruit is a pod (legume) that opens by one or both sutures, sometimes indehiscent or jointed and breaking up into single-seeded segments, sometimes winged.

176. FABACEAE Pea family

This large family consists of trees, shrubs, vines and perennial and annual herbs. Stems can be self-supporting and erect, with or without spines, and climbing species can be twining or have tips of branches forming tendrils (Bauhinia). Roots often bear nodules that harbour nitrogen-fixing bacteria. Their leaves are usually alternate, rarely opposite or whorled, and stipules are usually present, sometimes developed into spines or absent. Leaves are usually pinnate or bipinnate, sometimes palmately compound or trifoliolate, seldom unifoliolate or simple, and in some cases they are modified into narrow phyllodes or the terminal leaflets are converted into tendrils; petioles are usually present. Inflorescences are usually racemes, corymbs, spikes, heads or panicles, rarely flowers solitary or clustered in the leaf axils. The bisexual or rarely unisexual flowers

Distribution: This is a more or less globally distributed family, with woody genera more prominent in the tropics, subtropics and

Terminology of papilionaceous flower. Caragana pygmaea [176]

pedicel

Southern Hemisphere, and herbaceous genera more diverse in temperate regions, especially in mild winter-rain and dry-summer climates. Phylogeny and evolution: The crown group of Fabaceae is c. 75 million years old. This is a species-rich group, and diversification may in some clades be related to genome duplication early-on in the evolution of these lineages. This genome duplication is also thought to be related to the evolution of N-fixing bacterial nodules in their roots. Biogeography of the family involves a combination of vicariance and long-distance dispersal. Genera such as Hymenaea (resin of which made amber deposits) probably crossed the Atlantic during the Tertiary, and most other bicontinental genera have divergence times that are too young to be explained by continental drift. The classic division into the three traditional subfamilies (or families) does not hold up, due to the polyphyly of Caesalpinioideae. A recent classification of the family now recognises six subfamilies: Duparquetioideae, Cercidoideae, Detarioideae, Dialioideae, Faboideae and Caesalpinioideae. Not all species have been sequenced and further progress is still to be made. Faboideae are frequently herbs or vines and often have once compound leaves and

Vachellia drepanolobium, Kenya [176]

calyx

standard

wings

stamens style keel

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249

FABALES usually zygomorphic, usually papilionaceous flowers (as described above), although there are also genera that are trees with regular flowers included in this subfamily. Stamens are usually fused, and fruits are often explosively dehiscent, with the two valves twisting. This clade is estimated to have diversified c. 30 million years ago. Mimosoids (fomerly subfamily Mimosoideae, now part of Caesalpinioideae), are mostly woody plants with usually bipinnate leaves and nectaries on their petioles. They have actinomorphic flowers that have long, coloured filaments, and sepals and petals are fused. Seeds have a u-shaped line on both sides. Early divergence of this group occurred in Africa some 21 million years ago, but older ages have been suggested due to recent fossil finds.

Genera and species: This is a large family consisting of about 745 genera and c. 16,020 species, with over a quarter of all species in five genera (Acacia, Astragalus, Crotalaria, Indigofera and Mimosa). A formal classification has recently been published, with the following six subfamilies mentioned above: Duparquetioideae (1 genus) – Duparquetia (1); Cercidoideae (13 genera;

313 species) – Adenolobus (2), Barklya (1), Bauhinia (c. 125), Brenierea (1), Cercis (10), Gigasiphon (5), Griffonia (4), Lasiobema (c. 17), Lysiphyllum (c. 9), Phanera (c. 85), Piliostigma (c. 4), Schnella (45) and Tylosema (5); Detarioideae (84 genera, 755 species) – Afzelia (11), Amherstia (1), Annea (2), Anthonotha (c. 30), Aphanocalyx (14), Augouardia (1), Baik iaea (c. 6), Barnebydendron (1), Berlinia (21), Bikinia (10), Brachycylix (1), Brachystegia (26), Brandzeia (1), Brodriguesia (1), Brownea (12), Browneopsis (6), Colophospermum (1), Copaifera (c. 35), Crudia (c. 53), Cryptosepalum (c. 11), Cynometra (c. 85), Daniellia (c. 9), Detarium (3), Dicymbe (c. 18), Didelotia (c. 11), Ecuadendron (1), Elizabetha (c. 11), Endertia (1), Englerodendron (1), Eperua (14), Eurypetalum (3), Gabonius (1),

Paubrasilia echinata, Recife, Pernambuco, Brazil [176]

Delonix regia, Réunion [176]

Parkinsonia aculeata, Isabela, Galápagos Islands [176]

Dichrostachys cinerea, cultivated in Guadeloupe [176]

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EUDICOTS

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The rest (previously called “Caesalpinioideae”) are woody plants that have slightly zygomorphic f lowers (to more strongly zygomorphic and papilionaceous when Cercis and relatives are included). These often have a well-developed hypanthium, but do not form a natural unit. A number of genera await revision, and generic delimitations of some are not clear.

Mimosa pudica, Guadeloupe [176]

FABALES

EUDICOTS

Acacia spectabilis, Durran Durra, New South Wales, Australia [176]

Cercis siliquastrum, Lebanon [176]

Gilbertiodendron (c. 27), Gilletiodendron (5), Goniorrhachis (1), Gossweilerodendron (2), Guibourtia (c. 14), Hardwickia (1), Heterostemon (7), Humboldtia (6), Hylodendron (1), Hymenaea (14), Hymenostegia (c. 15), Icuria (1), Intsia (3), Isoberlinia (5), Julbernardia (c. 11), Kingiodendron (6), Lebruniodendron (1), Leonardoxa (1), Leucostegane (2), Librevillea (1), Loesenera (4), Lysidice (2), Macrolobium (c. 75), Maniltoa (c. 23), Michelsonia (1), Micklethwaitia (1), Microberlinia (2), Neoapaloxylon (3), Neochevalierodendron (1), Normandiodendron (2), Oddoniodendron (3), Oxystigma (5), Paloue (4), Paloveopsis (1), Paramacrolobium (1), Pellegrinodendron (1), Peltogyne (c. 25), Plagiosiphon (5), Polystemonanthus (1), Prioria (1), Pseudomacrolobium (1), Saraca (11), Schotia (4), Scorodophloeus (3), Sindora (c. 19), Sindoropsis (1), Stemonocoleus (1), Talbotiella (3), Tamarindus (1), Tessmannia (c. 12), Tetraberlinia (7) and Zenkerella (5); Dialioideae (20 genera, 714 species) – Androcalymma (1), Apuleia (3), Baudouinia (6), Cassia (c. 30), Chamaecrista (c. 330), Dialium (c. 28), Dicorynia (2), Distemonanthus (1), Eligmocarpus (1), Kalappia (1), Koompassia (3), Labichea (14), Martiodendron (4), Mendoravia (1), Petalostylis (2), Poeppigia (1), Senna (c. 300), Storckelia (4), Uittienia (1) and Zenia (1); Faboideae (484 genera, 13,483 species) – Abrus (c. 17), Acmispon (8), Acosmium (1), Adenocarpus (15),

Adenodolichos (c. 17), Adesmia (c. 240), Aenictophyton (1), Aeschynomene (c. 177), Afgekia (3), Aganope (7), Airyantha (2), Akschindlium (1), Aldina (c. 14), Alexa (9), Alhagi (3), Alistilus (3), Almaleea (5), Alysicarpus (c. 27), Amburana (3), Amicia (c. 7), Ammodendron (4), Ammopiptanthus (1), Ammothamnus (2), Amorpha (c. 15), Amphicarpaea (5), Amphimas (4), Amphithalea (42), Anagyris (2), Anarthrophyllum (15), Andira (29), Angylocalyx (7), Antheroporum (4), Anthyllis (22), Antopetitia (1), Aotus (c. 17), Aphyllodium (7), Apios (c. 7), Apoplanesia (1), Apurimacia (2), Arachis (69), Argyrocytisus (1), Argyrolobium (c. 80), Arthroclianthus (c. 30), Aspalathus (278), Astragalus (c. 2,400), Ateleia (20), Austrodolichos (1), Austrosteenisia (4), Baphia (47), Baphiastrum (1), Baphiopsis (1), Baptisia (c. 16), Barbieria (1), Barnebyella (1), Behaimia (1), Bergeronia (1), Biserrula (1), Bituminaria (2), Bobgunnia (2), Bocoa (3), Bolusafra (1), Bolusanthus (1), Bolusia (5), Bossiaea (c. 60), Bowdichia (2), Bowringia (4), Brongniartia (c. 63), Brya (4), Bryaspis (2), Burkilliodendron (1), Butea (2), Cadia (7), Cajanus (34), Calia (4), Calicotome (3), Callerya (c. 20), Callistachys (1), Calobota (16), Calophaca (c. 7), Calopogonium (5), Calpurnia (7), Camoensia (2), Camptosema (10), Campylotropis (c. 37), Canavalia (c. 60), Candolleodendron (1), Caragana (c. 75), Carmichaelia (23), Carrissoa (1), Cascaronia (1), Castanospermum (1), Centrolobium (7),

Bauhinia tomentosa, Kenya [176]

Centrosema (c. 36), Chadsia (9), Chaetocalyx (13), Chapmannia (7), Chesneya (c. 30), Chorizema (27), Christia (c. 10), Chrysoscias (4), Cicer (43), Cladrastis (7), Clathrotropis (6), Cleobulia (4), Clianthus (2), Clitoria (c. 62), Clitoriopsis (1), Cochlianthus (2), Codariocalyx (2), Collaea (7), Cologania (c. 12), Colutea (c. 28), Cordyla (c. 7), Corethrodendron (4), Coronilla (9), Coursetia (c. 35), Craibia (10), Cranocarpus (3), Craspedolobium (1), Cratylia (c. 7), Cristonia (1), Crotalaria (c. 690), Cruddasia (2), Cullen (c. 34), Cyamopsis (4), Cyathostegia (1), Cyclocarpa (1), Cyclolobium (1), Cyclopia (23), Cymbosema (1), Cytisophyllum (1), Cytisopsis (2), Cytisus (c. 65), Dahlstedtia (2), Dalbergia (c. 250), Dalbergiella (3), Dalea (c. 165), Dalhousiea (3), Daviesia (135), Decorsea (6), Deguelia (c. 17), Dendrolobium (18), Derris (c. 57), Desmodiastrum (4), Desmodium (c. 275), Dewevrea (1), Dichilus (5), Dicraeopetalum (3), Dillwynia (c. 40), Dioclea (c. 40), Diphyllarium (1), Diphysa (15), Diplotropis (12), Dipogon (1), Dipteryx (c. 12), Discolobium (8), Disynstemon (1), Dolichopsis (1), Dolichos (c. 60), Dorycnium (8), Dorycnopsis (2), Droogmansia (5), Dumasia (c. 10), Dunbaria (20), Dussia (9), Dysolobium (4), Ebenus (c. 20), Echinospartum (5), Eleiotis (2), Eminia (4), Endosamara (2), Eremosparton (3), Erichsenia (1), Erinacea (1), Eriosema (c. 150), Erophaca (1), Errazurizia (4), Erythrina (c. 120), Etaballia (1), Euchilopsis (1), Euchresta (4), Eutaxia Plants of the World

251

FABALES

Swainsona formosana, South Australia [176]

(10), Eversmannia (4), Eysenhardtia (c. 14), Exostyles (4), Fiebrigiella (1), Fissicalyx (1), Flemingia (c. 33), Fordia (18), Galactia (c. 57), Galega (6), Gastrolobium (109), Geissaspis (2), Genista (c. 90), Genistidium (1), Geoffroea (2), Gliricidia (5), Glycine (19), Glycyrrhiza (c. 20), Gompholobium (44), Gonocytisus (3), Goodia (2), Grazielodendron (1), Gueldenstaedtia (c. 10), Guianodendron (1), Halimodendron (1), Hammatolobium (2), Hanslia (2), Haplormosia (1), Hardenbergia (3), Harleyodendron (1), Harpalyce (24), Hebestigma (1), Hedysarum (c. 160), Hegnera (1), Herpyza (1), Hesperolaburnum (1), Hesperothamnus (5), Hippocrepis (c. 34), Hoita (3), Holocalyx (1), Hosackia (11), Hovea (37), Humularia (c. 35), Hylodesmum (14), Hymenocarpos (1), Hymenolobium (c. 17), Hypocalyptus (3), Indigastrum (c. 8), Indigofera (c. 700), Inocarpus (3), Isotropis (10), Jacksonia (74), Kebirita (1), Kennedia (c. 15), Kotschya (31), Kummerowia (2), Kunstleria (8), Lablab (1), Laburnum (2), Lackeya (1), Ladeania (2), Lamprolobium (2), Lathyrus (c. 160), Latrobea (6), Lebeckia (15), Lecointea (4), Lembotropis (2), Lennea (3), Lens (5), Leptoderris (c. 20), Leptodesmia (3), Leptolobium (10), Leptosema (13), Lespedeza (c. 35), Lessertia (c. 50), Leucomphalos (1), Limadendron (2), Liparia (20), Lonchocarpus (c. 120), Lotononis (c. 150), Lotus (c. 125), Luetzelburgia (8), Lupinus (c. 225), Luzonia (1), Maackia (c. 8), Machaerium (c. 130), Macropsychanthus (2), Macroptilium (c. 17), 252

Christenhusz, Fay & Chase

EUDICOTS

Cadia purpurea, a legume with regular flowers, Nairobi, Kenya [176]

Macrot yloma (24), Maraniona (1), Margaritolobium (1), Marina (38), Mastersia (2), Mecopus (1), Medicago (83), Meizotropis (2), Melilotus (c. 20), Melliniella (1), Melolobium (15), Microcharis (36), Mildbraediodendron (1), Milletia (c. 150), Mirbelia (32), Monarthrocarpus (1), Monopteryx (4), Montigena (1), Mucuna (c. 105), Muellera (2), Muelleranthus (3), Mundulea (12), Myrocar p u s (5), Myrospermum (3), Myroxylon (2), Mysanthus (1), Neocollettia (1), Neoharmsia (2), Neonotonia (2), Neorautanenia (3), Neorudolphia (1), Nephrodesmus (6), Nesphostylis (4), Nissolia (13), Nogra (3), Oberholzeria (1), Ohwia (2), Olneya (1), Onobrychis (c. 130), Ononis (c. 75), Ophiocarpus (1), Ophrestia (c. 16), Orbexilum (8), Oreophysa (1), Ormocarpopsis (6), Ormocarpum (c. 18), Ormosia (c. 130), Ornithopus (5), Oryxis (1), Ostryocarpus (2), Otholobium (61), Otion (c. 8), Otoptera (2), Ottleya (13), Ougeinia (1), Oxylobium (6), Oxyrhynchus (4), Oxytropis (c. 350), Pachyrhizus (5), Panurea (2), Paracalyx (6), Paraderris (13), Paramachaerium (5), Paratephrosia (1), Parochetus (2), Parryella (1), Pearsonia (13), Pediomelum (22), Peltiera (2), Periandra (6), Pericopsis (4), Petaladenium (1), Peteria (4), Petteria (1), Phaseolus (c. 63), Philenoptera (12), Phylacium (2), Phyllodium (8), Phyllota (11), Phylloxylon (7), Physostigma (4), Pickeringia (1), Pictetia (8), Piptanthus (2), Piscidia (c. 7),

Pisum (c. 3), Plagiocarpus (1), Platycelyphium (1), Platycyamus (2), Platylobium (4), Platymiscium (19), Platypodium (2), Platysepalum (8), Podalyria (19), Podocytisus (1), Podolobium (6), Podolotus (1), Poecilanthe (c. 11), Poiretia (11), Poissonia (5), Poitea (12), Polhillia (7), Pongamiopsis (3), Pseudarthria (4), Pseudeminia (4), Pseudoeriosema (4), Pseudolotus (1), Pseudovigna (2), Psophocarpus (c. 10), Psoralea (c. 50), Psoralidium (3), Psorothamnus (9), Pterocarpus (c. 37), Pterodon (3), Ptycholobium (3), Ptychosema (2), Pueraria (c. 18), Pultenaea (104), Pycnospora (1), Pyranthus (6), Rafnia (19), Ramirezella (7), Ramorinoa (1), Requienia (3), Retama (4), Rhodopis (2), Rhynchosia (c. 230), Rhynchotropis (2), Riedeliella (3), Robinia (4), Robynsiophyton (1), Rothia (2), Rupertia (3), Sakoanala (2), Salweenia (1), Sarcodum (3), Sartoria (1), Schefflerodendron (4), Scorpiurus (2), Securigera (13), Sellocharis (1), Sesbania (c. 60), Shuteria (5), Sinodolichos (2), Smirnowia (1), Smithia (c. 20), Soemmeringia (1), Sophora (c. 50), Spartidium (1), Spartium (1), Spathionema (1), Spatholobus (29), Sphaerolobium (22), Sphaerophysa (2), Sphenostylis (7), Sphinctospermum (1), Spirotropis (3), Spongiocarpella (c. 7), Staminodianthus (3), Stauracanthus (3), Stirtonanthus (3), Stonesiella (1), Streblorrhiza (1, possibly extinct), Strongylodon (12), Strophostyles (3), Stylosanthes (c. 25), Styphnolobium (9), Sulla (7), Sutherlandia (5),

FABALES

EUDICOTS

Swainsona (84), Swartzia (c. 180), Sweetia (1), Sylvichadsia (4), Syrmatium (14), Tadehagi (c. 6), Taralea (c. 7), Taverniera (15), Templetonia (10), Tephrosia (c. 350), Teramnus (9), Tetragonolobus (6), Teyleria (3), Thermopsis (c. 23), Thinicola (1), Tibetia (4), Tipuana (1), Trifidacanthus (1), Trifolium (c. 250), Trigonella (c. 55), Tripodion (1), Trischidium (5), Uleanthus (1), Ulex (c. 15), Uraria (c. 20), Uribea (1), Urodon (4), Vandasina (1), Vatairea (8), Vataireopsis (4), Vatovaea (1), Vaughania (11), Vavilovia (1), Vicia (c. 160), Vigna (c. 104), Viminaria (1), Virgilia (2), Wajira (5), Weberbauerella (2), Wiborgia (9), Wiborgiella (9), Wisteria (6), Xanthocercis (3), Xeroderris (1), Xiphotheca (9), Zollernia (10), Zornia (c. 75) and Zygocarpum (6); Caesalpinioideae (139 genera, c. 3,725 species) – Arapatiella (2), Acrocarpus (1), Aroa (1), Balsamocarpon (1), Batesia (1), Burkea (1), Bussea (7), Caesalpinia (c. 25), Campsiandra (19), Cenostigma (3), Ceratonia (2), Chidlowia (1), Colvillea (1), Conzattia (1), Cordeauxia (1), Coulteria (10), Delonix (11), Dimorphandra (26), Diptychandra (1), Erythrophleum (10), Erythrostemon (12), Gleditsia (c. 14), Guilandina (c. 7), Gymnocladus (6), Haematoxylum (3), Hoffmannseggia (24), Jacqueshuberia (7), Lemuropisum (1), Libidibia (7), Lophocarpinia (1), Melanoxylon (1), Mezoneuron (c. 26), Moldenhawera (9), Clitoria ternatea, Royal Botanic Gardens, Kew, UK [176]

Mora (6), Moullava (1), Orphanodendron (2), Pachyelasma (1), Parkinsonia (12), Paubrasilia (1), Peltophorum (c. 6), Poincianella (c. 35), Pomaria (16), Pterogyne (1), Pterolobium (11), Recordoxylon (3), Schizolobium (1), Stachyothyrsus (2), Stahlia (1), Stenodrepanum (1), Stuhlmannia (1), Sympetalandra (5), Tachigali (c. 65), Tetrapterocarpon (2), Tara (3), Umtiza (1), Vouacapoua (3) and Zuccagnia (1); mimosoid clade (now a subclade under Caesalpinioideae) (82 genera, 3,301 species) – Abarema (46), Acacia (c. 1,075), Acaciella (15), Adenanthera (13), Adenopodia (c. 7), Alantsilodendron (10), Albizia (c. 130), Amblygonocarpus (1), Anadenanthera (2), Archidendron (94), Archidendropsis (14), Aubrevillea (2), Blanchetiodendron (1), Calliandra (c. 135), Calliandropsis (1), Calpocalyx (11), Cathormion (1), Cedrelinga (1), Chloroleucon (10), Cojoba (12), Cylicodiscus (1), Desmanthus (c. 24), Dichrostachys (14), Dinizia (1), Ebenopsis (3), Elephantorrhiza (9), Entada (c. 28), Enterolobium (11), Faidherbia (1), Falcataria (3), Fillaeopsis (1), Gagnebina (8), Guinetia (1), Havardia (5), Hesperalbizia (1), Hydrochorea (3), Indopiptadenia (1), Inga (c. 300), Kanaloa (1), Lemurodendron (1), Leucaena (22), L e u c o chloron (5), Lysilo m a (9), Macrosamanea (11), Marmaroxylon (c. 11), Microlobius (1), Mimosa (c. 500), Trifolium pratense, Turku, Finland [176]

Mimozyganthus (1), Neptunia (12), Newtonia (15), Painteria (3), Parachidendron (1), Parapiptadenia (6), Paraserianthes (1), Parkia (c. 34), Pentaclethra (3), Piptadenia (c. 21), Piptadeniastrum (1), Piptadeniopsis (1), Pithecellobium (18), Pityrocarpa (3), Plathymenia (1), Prosopidastrum (c. 5), Prosopis (c. 44), Pseudopiptadenia (11), Pseudoprosopis (7), Pseudosamanea (2), Samanea (3), Sanjappa (1), Schleinitzia (4), Senegalia (c. 230), Serianthes (c. 18), Sphinga (3), Stryphnodendron (c. 30), Tetrapleura (2), Thailentadopsis (3), Vachellia (c. 160), Viguieranthus (c. 23), Wallaceodendron (1), Xerocladia (1), Xylia (9), Zapoteca (20) and Zygia (c. 47). Uses: Fabaceae have a wide range of uses, including food, aphrodisiacs, agroforestry, beads, biofuel, boats, boxes, bridges, brooms, buttons, cabinetry, canoes, carpentry, char coal, climate mitigation, construction, coppicing, cosmetics, crates, dental floss, docks, dyes, ecological restoration, fertiliser, fibre, firewood, flooring, fodder, furniture, garden ornamentals, glue, gums, honey, insecticides, lacquers, lubricants, medicines, musical instruments, paper, perfumes, poisons, reforestation, rope, roof shingles, shade trees, shore protection, soap, soil stabilisation, sorcery, sewing needles, spices, tanning, utensils, veneers, Abrus precatorius, pods and seeds, Seychelles [176]

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Saraca indica, Hong Kong, China [176]

weaponry, witchcraft etc. Here, we have selected economically important and some remarkable usages and grouped the species accordingly. Pulse and vegetable crops: soybean (Glycine max) was an important crop in China before written records and was domesticated from wild G. soja in East Asia, the oldest records being from 2,000-year-old archaeological sites in Korea. It is now the most economically important legume crop in the world. The USA, Brazil and Argentina are the biggest soybean producers, but only a small proportion of the crop is directly consumed by humans, most being used as animal feed usually on an industrial scale. However, soya is used in a great number of processed foods (soy lecithin as a stabiliser), and of course it forms the basis of soy sauce, soy milk, tofu, miso, natto and tempeh, foods of increasing importance in vegetarian diets. The marama bean (Tylosema esculentum) from southern Africa is also eaten and has oil-rich seeds that have a similar amount of protein as soybeans. Being perennial, droughtand poor-soil-tolerant, this species has great potential for cultivation in semi-arid lands as a sustainable, erosion-preventing, high-yield crop. When roasted the seeds taste like cashew or chestnut. Second to the soybean, the most important legume is the peanut or groundnut (Arachis hypogea). It was domesticated from the Neotropical A. monticola and was already cultivated in pre-Colombian times. Until 254

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Tamarindus indica, China (HS) [176]

1930, it was mostly used as animal fodder, after which use for human consumption was promoted in Europe and North America. Introduced into China in the 17th century, it is now a major part of Chinese cuisine, and China is the largest producer of the crop, followed by India and the USA. Eaten roasted or boiled, often ground into flour, as peanut butter (an Aztec invention popularised in the late 19th century), or pressed into oil, it finds a great variety of applications in cooking, soaps, cosmetics, polishes, paints and lubricants. Of course peanuts are also commonly used in snacks and confectionery such as candy bars, peanut butter cups and peanut brittle. The common or garden pea (Pisum sativum) is a major pulse and vegetable crop. Originally collected from the wild in Neolithic times, it was soon domesticated and became a staple of the vegetable garden. The dried seeds could be kept for a long time and were ground into flour or boiled into pea soup, keeping famine at bay during Mediaeval times. Fresh garden peas and immature pods (mange-tout or snow peas) were a luxury food in early modern Europe (15th to 18th centuries), and pea shoots are a recent culinary novelty. The village of Peasenhall in Suffolk, England, celebrates an annual Pea Festival in honour of this famous legume. The type species of the family, the broad or fava bean (Vicia faba), is native to the southern Mediterranean and southwestern and southern Asia but is now cultivated

Amherstia nobilis, cultivated in Singapore (WA) [176]

worldwide. It was an important part of the Mediterranean diet c. 6,000 years ago, together with lentils, peas and chickpeas. There are many varieties, those with larger beans being eaten when young and tender, and those with smaller harder seeds (often called field or horse beans) being used as animal food or in some Middle Eastern dishes. They can also be deep fried as a savoury, crunchy snack, known in Latin America as ‘habas’. Broad beans are an important part of local cuisine in many parts of the world, but the main broad bean producers are China, Ethiopia and England. Of American origin, Phaseolus species were already widely cultivated before Columbus reached the hemisphere. Most important is the common bean (Phaseolus vulgaris), which includes varieties that produce brown, white or yellow beans, kidney beans, black turtle beans, borlotti beans, flageolet beans, haricots, string beans, pea beans and pinto beans. Some are eaten dried, whereas others are eaten while the pods are still young as a vegetable (e.g. haricots verts). They are often canned or made into soups or sauces and are important in many traditional cuisines in the New and Old World. Other commonly cultivated Phaseolus species of which seeds and/or pods are eaten are the scarlet runner bean (P. coccineus), lima or butter bean (P. lunatus), year bean (P. dumosus) and tepary bean (P. acutifolius). The mainly Old World genus Vigna includes a number of bean species. Most

FABALES

EUDICOTS

important are the black-eyed pea (Vigna unguiculata var. unguiculata) and yardlong bean (V. unguiculata subsp. sesquipedalis), which are important in African and Asian cuisines. Other more locally used beans are the moth bean (V. aconitifolia), adzuki bean (V. angularis), urd bean (V. mungo), mung bean (V. radiata) and rice bean (V. umbellata). High in protein, the groundnut or bambara bean (Vigna subterranea) is a staple food in West Africa and is currently also frequently grown in Spain, Pakistan, China and Australia. Lentils (Lens culinaris) have been widely cultivated since Neolithic times and are still an important pulse crop. Seeds are eaten cooked or ground into flour or dhal. The largest producers are Canada, India and Australia. Lentils come in a number of varieties and colours including green, yellow, orange-red, brown and black. Chickpeas (Cicer arietinum) are a major crop also cultivated since the Neolithic (c. 3500 BC), and wild chickpeas have been carbon-dated to c. 6800 BC after being found in a Mesolithic cave in France. Chickpeas were widely grown in Ancient Greece and throughout the Roman Empire and have been popular ever since. Seeds can be eaten freshly cooked or made into flour or paste and are a key ingredient of hummus, falafel and other Middle Eastern, Iberian, Jewish, Burmese and Indian recipes. Yeheb nuts (Cordeauxia edulis), from Cassia fistula, Seychelles [176]

the Horn of Africa, taste like sweet chestnuts and are a good arid-land food crop. Other pulse and/or vegetable crops eaten locally are pigeon bean (Cajanus cajan), jack bean (Canavalia ensiformis), Moreton Bay chestnut (Castanospermum australe), Tahitian chestnut (Inocarpus fagifer), bonavist bean (Lablab purpureus), chickling vetch (Lathyrus sativus), lupine bean (Lupinus albus and L. luteus), pearl lupine (L. mutabilis), Kersting’s groundnut (Macrotyloma geocarpum), yam bean (Pachyrhizus erosus), Goa bean (Psophocarpus tetragononolobus), African yam bean (Sphenostylis stenocarpa) and asparagus pea (Tetragonolobus purpureus). The seeds of carob (Ceratonia siliqua), Kentucky coffee tree (Gymnocladus dioicus) and Bauhinia petersiana can be roasted and used as a coffee substitute. Seeds of “flamboyant” (Delonix) can be crushed and eaten as a snack, and seeds of the South African boer-bean (Schotia) can be eaten after cooking. Young sataw beans (Parkia speciosa) and néré (P. biglobosa) are frequently eaten in the Old World tropics as a vegetable. Seedlings of Medicago sativa are eaten as alfalfa sprouts, and the bean sprouts common in Asian cuisine are from Phaseolus aureus and other species. Edible pulp: many legumes produce pods that contain edible pulp surrounding the seed. Tamarind (Tamarindus indica) is native to East Africa, but was introduced into Asia in ancient times. The pods have an acid-sweet

pulp around the seeds that can be eaten raw but is frequently used in confectionery and as an ingredient of Asian and African cuisine, especially curries, sauces and pickles. The pulp is also fermented into drinks, and seeds are also edible, raw, cooked or pressed into oil. The bark and leaves can be used to produce a dye, it produces good wood and is a good bee fodder. Tannin yields are also high. It truly is a plant with many uses. The fruit of velvet tamarind (Dialium spp.) is similar to tamarind and used in chutneys. The icecream bean (Inga edulis) has an edible white pulp that is often eaten in the tropics and used to flavour desserts. Guayamochil (Pithecellobium dulce) and rain tree (Samanea saman) are frequently planted for shade and their edible pulp, which can be made into drinks. Fruits and seeds of Chilean palo verde or kumbaru (Geoffroea decorticans) are sweet and replace molasses in local cuisines. Beans of mesquite (Prosopis spp.) can be ground into flour and added to bread or made into jelly or wine. Carob (Ceratonia siliqua) is a cultigen originating in Arabia (derived from C. oreothauma) that has pods containing sugary pulp and gum, which were formerly sold as sweets and now as an alternative for chocolate; they are used as a food additive and in cosmetics, medicines, photographic film, adhesives, paints, ink and polishes. The pods are commonly used as fodder and pet food (for hamsters, rabbits etc.) and sometimes made into a flour for

Labichea lanceolata, Australian National Botanic Garden, Canberra [176]

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255

FABALES

Genista tinctoria, Ruissalo Botanical Garden, Turku, Finland [176]

EUDICOTS

Phaseolus coccineus, private allotment Kingston upon Thames, Surrey, UK [176]

diabetics. The seeds were highly valued and were the locust beans of the Bible, used as a measure of weight and the origin of the carat (200 mg) of jewellers, due to their regular size and weight. The fruit pulp of honey locust (Gleditsia triacanthos) is edible and used in locally brewed beers, and the edible leaves and sweet pulp of datach (Detarium senegalense) are eaten locally in Sub-Saharan Africa. Edible roots: tubers of jicama (Pachyrhizus erosus) are popular in Mexican cuisine and delicious eaten fresh or cooked. Tubers of potato bean (Apios americana), hog potato (Hoffmannseggia glauca) and prairie turnip (Pediomelum esculentum) are roasted and eaten in North America, and tubers of some Lathyrus species are also edible (e.g. L. montanus, L. ochrus, L. tuberosus). Kudzu (Pueraria spp.) has starchy edible tubers that are popular in Japan, but elsewhere it is often invasive, for example causing ecological havoc in southeastern North America. Spices and f lavourings: fenugreek (Trigonella foenum-graecum) is an important condiment for flavouring food, especially Indian curries. Liquorice roots (Glycyrrhiza glabra) have a somewhat sweet flavour and are used in confectionery, drinks and tobacco. Liquorice root was often sold by herbalists and was chewed to release the sweet juice. In candy, liquorice extract is commonly mixed with aniseed oil and sometimes with 256

Christenhusz, Fay & Chase

Pisum sativum, private allotment, Kingston upon Thames, Surrey, UK [176]

salmiak (ammonium chloride), especially in the Netherlands (drop), Denmark (lakrids) and Finland (lakritsi, salmiakki). In England, liquorice was first mixed with sugar to make Pontefract cakes. In its natural form, liquorice is frequently used in Italy and Spain as a mounth freshener and made into ice-cream and liqueurs. Bark and leaves of bumba (Scorodophloeus zenkeri) are used as a garlic substitute in Africa. Teas: grown on a commercial scale in South Africa, the chopped and fermented stems and leaves of redbush or rooibos (Aspalathus linearis) are used as an infusion (to calm the stomach, especially of babies) and sold as a herbal tea. Similarly, honeybush or heuningbos (Cyclopia intermedia and C. subternata) is marketed in health food shops as an alternative to tea. Edible flowers: leaves, flowers and pods of the orchid tree (Bauhinia variegata) are edible. It is a sacred plant of Buddhists and the floral emblem of Hong Kong. Flowers of redbud (Cercis canadensis) can be eaten fresh in salads or pickled. Flowers and young leaves of the pride of Burma (Amherstia nobilis) are also edible. This plant has only been found in the wild twice and is possibly extinct, but it is a highly valued ornamental tree and planted throughout the tropics. Gums: several species can be tapped for their resin or gum, most notably gum

arabic, the sap of Senegalia senegal, used in sweets, lozenges, thickeners, adhesives, inks, watercolour paints etc. Other species of Senegalia (e.g. Amritsar gum, S. modesta) and species of Acacia (A. dealbata, A. pycnantha), manna (Alhagi graecorum), Burkea africana, guar gum (Cyamopsis tetragonoloba) and Vachellia (V. gummifera, V. horrida, V. nilotica) also produce edible gums, often used as food additives. Copal (Copaifera sp.) and gum copal (Daniellia oliveri) produce a highquality resin used in medicine, varnishes, paint and lacquer and as biodiesel. Gum tragacanth (Astragalus gummifer) is used in a wide range of industries. Wallaba oil (from Eperua oleifera) and other oleo-resins from, for instance Gossweilerodendron, Guibourtia, Kingiodendron, Myrospermum, Myroxylon and Prioria, can be used to make incense. Seeds of Adenanthera pavonina produce an industrial lubricant. Soap and perfume: seeds of Gleditsia sinensis and Gymnocladus chinensis contain saponins and are used in Asia to make soap and for washing clothes. Flowers of needle bush, Vachellia farnesiana, produce a perfume called ‘cassie’, highly valued in the perfume industry. Flowers of a number of broom species (Genista, Cytisus and relatives) are also used to produce perfume. The fragrant seeds of cumaru or tonka bean (Dipteryx odorata), are high in coumarin and used as a flavouring.

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Dyes: indigo (derived from Indigofera arrecta, I. articulata, I. suffruticosa and I. tinctoria) is probably the best known vegetable dye. It was domesticated originally in India from where it was exported as a luxury item to the Roman Empire. It was used to dye silk and in early book printing. Indigo remained rare due to the availability of woad (Isatis tinctoria, Brassicaceae), which gives a similar blue. A traditional dye for tuareg robes, batik and other fabrics, it was taken to the Carolina Settlement in North America, where it was used to dye clothes, and from where it has been exported since the 18th century. Blue jeans, invented in the USA in the 1870s, were originally dyed with natural indigo. Dyer’s broom (Genista tinctoria) was formerly used in Europe to produce a bright yellow dye. Brazilwood, Paubrasilia (Caesalpinia) echinata, was harvested on a large scale giving the name to the country Brazil (a name given in reference to the colour of glowing coal). Sappanwood (Caesalpinia sappan), also sometimes called brazilwood, is also used as a dye, but originates in tropical Asia. Logwood

(Haematoxylum campechianum) contains haematoxylin, a source of a dark violet dye used in dying cotton, fur, leather, silk, wool and synthetic fibres; it is used additionally to stain nuclei in biological microscope slides. Peachwood (H. brasiletto) yields brasilin, a red dye. Many species yield dyes that are used locally, such as jeheb nut (Cordeauxia edulis, purple), yellow flamboyant (Peltophorum pterocarpum, yellow), Zuccagnia punctata (yellow), Griffonia spp. (black), sangre de drago (Pterocarpus tinctorius, red) etc. Tanning: a good number of species produce tanniferous bark, but especially divi-divi (Libidibia coriaria) and guayacán (L. paraguariensis) are used on an industrial scale for tanning leather. Other species used for tanning are Balsamocarpon brevifolium, Bauhinia variegata, Burkea africana, Dimorphandra mollis, Elephantorrhiza spp., Melanoxylon brauna, Piliostigma thonningii, Tachigali tinctoria, Tara spinosa, etc. Fodder, forage and green manure: due to the ability to fix nitrogen in their root nodules, many Fabaceae are employed to fertilise the

soil. Commercially, a number of species are available as green manure, plants grown and then ploughed into the soil as natural fertiliser. Well-known species for this purpose are bird’s-foot trefoil (Lotus corniculatus), yellow lupine (Lupinus luteus), lucerne (Medicago sativa), serradella (Ornithopus sativus) and clover (Trifolium spp.). Clover and lucerne (called alfalfa in North America) are also important fodder and forage plants, being cultivated to produce high-protein greens to feed to animals. Other popular fodder plants are calopo (Calopogonium mucunoides), creeping indigo (Lotononis bainesii), jigal tree (Lysiphyllum cunninghamii), sainfoin (Onobrychis viciifolia), Brazilian lucerne (Stylosanthes guianensis), sulla clover or Italian sainfoin (Sulla coronaria), bitter vetch (Vicia ervilla), common vetch (V. sativa), fodder vetch (V. villosa) etc. Some species of forage legumes can cause the disease known as bloat if animals are allowed to eat them in large quantities, particularly in spring. In this disease, proteins in the forage can cause the formation of a stable foam in the

Plants at war — Augustin Pyrame de Candolle (1778–1841) In 1824 Swiss botanist Augustin de Candolle started a world f lora called the Prodromus systemtatis naturae regni vegetabilis. Although it was never finished, 161 families and 58,000 species were described, and it was later continued by his son Alphonse and his grandson Casimir Pyrame de Candolle. Both were famous botanists in their own right and they also passed this tradition on to a fourth generation, as his great-grandson Richard Emile Augustin de Candolle became a botanist too. Augustin de Candolle believed that there were sharp discontinuities between natural catagories, contrary to some contemporary researchers like Jussieu, and he also advocated the struggle between plant species for nutrients and light, which he called ’nature’s war’. This later influenced Charles Darwin in developing his principle of natural selection as

the driving force behind evolution. He also discovered that several species could develop similar characteristics that were not present in their ancestors, a term later called analogy. This was of great importance in his classification of plants as he distinguished between morphological and physiological characteristics, allowing him to come to a better understanding of relationships. He also studied movement of plants, suggesting that an internal biological clock exists, a finding many scientists at the time were sceptical about, but which was proven to be correct nearly a century later. De Candolle described over 10,000 new species, more than any botanist. At least six times a genus Candollea was described during the 19th century, the oldest having been based on a species of Pyrrosia (Polypodiaceae), so Candollea is now a synonym of that genus. Candolleodendron brachystachyum is used for a species of Fabaceae. Candollea is also

Engraving of Augustin Pyramus de Candolle by Jules Pizzetta, Galerie des Naturalistes de Jules Pizzetta, ed. Hennuyer, 1893 (public domain)

the name of the botanical journal of the Botanical Garden of the city of Geneva, in honour of the important contributions of this botanical dynasty.

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Vicia faba, private allotment, Kingston upon Thames, Surrey, UK [176]

Hoffmannseggia glauca, New Mexico, USA (DZ) [176]

Glycine mas, private allotment, Kingston upon Thames, Surrey, UK [176]

rumen, stopping the animal being able to release the gases formed by fermentation/ digestion. If caught in time, the gas can be released by puncturing the rumen (as seen in the film Far from the madding crowd, based on the book by Thomas Hardy). In severe cases, the rumen can burst, leading to death of the animal. Species containing relatively high concentrations of condensed tannins (e.g. Onobrychis sativa and some Trifolium species) are sometimes advocated as a way of avoiding bloat, as the tannins precipitate the soluble proteins, but they also give a bitter taste to the forage making them less attractive to the animals. Agroforestry: many Fabaceae are important shade trees, especially for coffee plantations and other shade-loving crops. A special type of agroforestry is alley cropping, for which species of Inga (e.g. I. edulis) are especially suitable in Latin America, and trials with other genera such as Milletia and Lysiloma are under way in Africa. Alley cropping, in which Inga (or other Fabaceae) tree seedlings are planted in a series of hedgerows, is a sustainable alternative to slash and burn cultivation. The fallen Inga leaves create a thick layer of mulch, shading out weeds and exotic grasses, and the nitrogen-fixing nodules on the roots enrich the soil. After two years, the Inga hedgrows are coppiced, the leaves used as mulch, the branches for firewood near the homestead (tackling further deforestation for that purpose), and a crop can then be planted in the alleys between the coppiced hedgerows. Once the crop is harvested, the

Inga plants grow tall until the next planting season, hence recycling the nutrients and preventing erosion at the same time. For more information, see the webpages of the Inga Foundation: www.ingafoundation.org. Brooms and fibre: Cytisus scoparius, or common broom, is grown as an ornamental and soil stabiliser and for perfume and dyes, but it received its name from the broom-like branches that can be easily made into brooms and were used frequently for that that purpose in the past. Other species of Cytisus, Genista, Spartium and Ulex have also been used as brooms. Sunn hemp (Crotalaria juncea) yields a high-quality bast fibre for cordage and fine paper, and there are numerous other species that are locally important fibre crops. Wood: as many species are dominant members of tropical forests, the family is an important source of wood for local communities, used for construction, turning, furniture, firewood and charcoal. Many species are cultivated commercially for their hard wood, the species being too numerous to list here. Plantations of Robinia pseudoacacia in the temperate zones supply these regions with durable hardwood to reduce the global demand for tropical hardwood timber and hopefully indirectly preserve some rainforests. However, some tropical species are still extensively harvested for their hard and durable wood. Truncheons of London policemen were once made of West Indian ebony (Brya ebenus) because it takes a smooth, high, black polish. The oily wood of umthiza, Umtiza listerana, could

be turned into self-lubricating propeller shafts. Rosewood (Dalbergia) is harvested extensively from the wild for its scented wood valued by the perfume industry. Plantations have now been established, but threats to wild populations remain. International trade of D. nigra, the most threatened species, is now controlled by CITES. Beads: several species have attractive seeds, often bright red or red with a black spot, and are used to make beads and buttons. Most popular species are the red bead tree (Adenanthera pavonina), the prayer bead vine (Abrus precatorius), chamfuta (Afzelia quanzensis), sea hearts (Entada spp.), the lucky bean tree (Erythrina spp.), the nicker bean (Guilandina bonduc), jumby bead (Ormosia spp.) and the rosary bean (Rhynchosia spp.). Some of these are extremely poisonous (e.g. Abrus precatorius), so care should be taken if jewellery made with these is handled. Hallucinogens, medicine and poison: despite the large number of species with edible seeds, Fabaceae include many species containing complex compounds, many of which are medicinal, hallicinogenic or poisonous. Seeds of cohoba (Anadenanthera peregrina), and to a lesser extent Dimorphandra parviflora, are crushed to make an intoxicating snuff in Latin America, frequently used in shamanic rituals. The seeds of Madagascan Lemuropisum edule are edible when young, but become poisonous when ripe. Sweet peas (Lathyrus odoratus) are grown for their flowers, but the seeds are poisonous and should not be confused with edible peas.

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EUDICOTS

Kennedia, Lablab, Laburnum, Lathyrus, Lespedeza, Lotus, Lupinus, Maackia, Mimosa, Neptunia, Parkinsonia, Robinia, Samanea, Saraca, Senna, Sophora, Strongylodon, Styphnolobium, Swainsona, Thermopsis, Virgilia and, of course, Wisteria are cultivated as ornamentals. Acacia dealbata is sold as a cut flower called ‘florist’s mimosa’.

Pachyelasma tessmannii and Erythrophleum suaveolens were used as fish poison and the latter also as a penal punishment in the past. Roots of Abrus precatorius (crab’s eye or prayer beads) can be used as a drink sweetener or alternative to liquorice but are mildly toxic. Additionally, many species contain insecticidal alkaloids, such as Derris elliptica and Gymnocladus dioica, and are used to protect crops and kill mosquitoes. Several species produce medicinally important compounds, useful in treatments of many diseases, including cancer, diabetes and malaria, but these are too numerous to list here. Ornamentals: many species of Fabaceae are grown as ornamentals, in perennial borders and as street and shade trees. Best known for its scented flowers (but poisonous seeds!) is perhaps the sweet pea (Lathyrus odoratus). Most commonly, members of the genera Acacia, Albizia, Amherstia, Amorpha, Anthyllis, Argyrocytisus, Baptisia, Bauhinia, Brownea, Caesalpinia, Calliandra, Caragana, Carmichaelia, Cassia, Ceratonia, Cercis, Chorizema, Cladrastis, Clianthus, Clitoria, Coronilla, Cytisus, Dalea, Delonix, Dorycnium, Erythrina, Erythrostemon, Galega, Gleditsia, Gliricidia, Gymnocladus, Hardenbergia, Hippocrepis, Hovea,

Giant beans: There are record holders among Fabaceae. Tualang (Koompassia excelsa) is, at 88 metres, one of the tallest eudicots, only surpassed by Eucalyptus regnans (Myrtaceae). The monkey ladder (Entada gigas) has pods that can grow up to two metres long and 12 cm across, holding up to 15 seeds, each c. 6 cm across. However, these are not the largest seeds, giant nato (Mora megistosperma) has the largest seed of any eudicot, measuring c. 12 by 18 cm. The seeds of oil nato (Mora oleifera) have the largest embryo of any eudicot seed. It can be up to a kilogram in weight and is as such similar to that of the monocot coco-de-mer (Lodoicea maldivica, Arecaceae). Remarkably, the large seeds of Mora and Entada are distributed by water currents. They float due to an air-filled cavity between the cotyledons; in contrast, the largest but heavy seeds of coco-de-mer (Lodoicea, Arecaceae) sink.

Lupinus nootkatensis, Helsinki Botanical Garden, Finland [176]

Ceratonia siliqua, Gibraltar [176]

Plant movement: Many species of Fabaceae have ‘sleeping’ leaves that fold and droop when night falls and revive in the morning. Not many plants move quickly enough to enable plant movement to be observed by the human eye, but the sensitive plant or touch-me-not (Mimosa pudica) has leaves that do this quickly in response to being touched or shaken, reopening minutes later. This movement is caused by an electric current that causes the turgor in cells of the petioles to change so that the leaflets fold inwards. It is not exactly known why this has evolved, but it is thought to be a response to predation by herbivores. Water mimosa (Neptunia oleracea) is an aquatic legume that also has leaves that close after touching. Another species with rapid movement is the semaphore plant (Codariocalyx motorius), but the leaves of this species move in response to the intensity of sunlight, not touch. Etymology: Faba is the Latin form of the Proto-Indo-European bhabh, a bean. Faba vulgaris is a later synonym of Vicia faba, the currently accepted name for broad bean. An alternative name for this family is Leguminosae, based on the type of fruit called a legume or bean pod. Indigofera ammoxylum, Réunion Botanical Garden [176]

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FABALES

EUDICOTS

Suriana maritima, Yucatán, Mexico [177]

Guilfoylia monostylis in fruit, Royal Botanic Gardens, Melbourne, Australia (CD) [177]

Suriana maritima in fruit, Guadeloupe [177]

Stylobasium spathulatum, Karijini National Park, Western Australia [177]

177. SURIANACEAE

clawed. The ten stamens are all fertile or the inner whorl of five sterile staminodes often reduced in number. Filaments are free, and the basifixed anthers open by a lengthwise slit. The superior ovary is sessile or stalked (in Recchia) and composed of one to five distinct carpels, each with a gynobasic style topped with a clavate to capitate stigma. Fruits are composed of one to five drupe- or nut-like monocarps (drupetum or achenetum).

well. Molecular analysis revealed that they are sister to Polygalaceae in Fabales; they share an apocarpous gynoecium with Quillajaceae, which is reduced to a single carpel in Fabaceae. A single fossil of Suriana from the Eocene of Wyoming is known, but molecular dating estimates that the family is c. 40 million years old. Stylobasium is windpollinated and was previously placed in its own family due to lack of synapomorphies with any taxon. Molecular analyses have placed it here.

Bay-cedar family

These are trees and shrubs with simple or pinnately compound alternate leaves that are petiolate or sessile. Stipules are small or absent. Inflorescences are terminal or axillary cymes or panicules, sometimes solitary in the leaf axils or borne on branches (cauliflorous), usually with bracts and bracteoles. The actinomorphic flowers are bisexual, sometimes unisexual, and have usually five (to seven) free, persistent sepals. The usually five petals (absent in Stylobasium) are free and sometimes 260

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Distribution: Due to the wide distribution of Suriana maritima along tropical coasts, the family occurs in the Caribbean, Indian Ocean islands, Malesian Archipelago and Pacific islands. Recchia is endemic to Mexico, and the remaining genera are only found in Australia. Phylogeny and evolution: Previously assigned to Simaroubaceae or Chrysobalanaceae, they were found to have numerous characters that did not match these families

Genera and species: Surianaceae include five genera with a total of eight species: Cadellia (1), Guilfoylia (1), Recchia (3), Stylobasium (2) and Suriana (1). Etymology: Suriana is named for French physician and apothecary Joseph Donat Surian (died 1691), who collected plants with Michel Bégon and Charles Plumier during an expedition to Haiti and Martinique.

FABALES

EUDICOTS

This is a family of perennial and annual herbs, shrubs, vines, trees (up to 50 m

in Xanthophyllum) and achlorophyllous mycoheterotrophic herbs (Epirixanthes). Stems are mostly round, but sometimes angular or winged, and branches can be tipped with spines. Leaves are alternate, opposite or in whorls, simple, petiolate or not, and blades have pinnate venation and entire margins. They are rarely reduced and scale-like (Epirixanthes). Stipules are absent, rarely spine- or scale-like. Inflorescences are terminal or axillary, simple or compound racemes or panicles (rarely f lowers can be solitary), with bracts and bracteoles subtending flowers. The zygomorphic flowers

Polygala vulgaris, Ireland [178]

Comesperma calymega, Mt Lesueur, Western Australia [178]

178. POLYGALACEAE Milkwort family

Polygala ehlersii, Taita Hills, Kenya [178]

are bisexual, pedicellate or sessile. The five sepals are free, partly fused at the base or fused into a tube, the outer three small, inner two (wings) large, petal-like, or all five nearly equal and petal-like. The three or five petals are free or basally fused, the lower one boat-shaped (keel), entire or trilobed, sometimes with an apical crest or fimbriate/ lamellate appendages. Stamens are usually five to eight (sometimes as few as two or upto ten), with free or variously united filaments and then forming a sheath open on the upper side, fused to the corolla lobes. The basifixed anthers usually open by a single apical pore.

Monnina crassifolia, Ecuador [178]

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FABALES A nectary disk is usually absent, but if present it is annular or glandular. The superior ovary is composed of two to eight carpels fused into one or two locules. The single style is erect or curved with one or two capitate stigmas. Fruits are capsules that open by valves, unilocular samaras or a fleshy drupes. Distribution: This family has a nearly worldwide distribution (absent from the Sahara and higher latitudes). They are especially diverse in tropical and subtropical regions of both hemispheres. Phylogeny and evolution: Previously placed either in their own order or in Malpighiales, the latter now known not to be correct. Molecular studies placed this family close to Surianaceae and Fabaceae in Fabales, with which they share few morphological characters, apart maybe from floral development. The family diversified during the Triassic, probably in response to the evolution of elaiosomes, which allow mutualism with ants for seed

EUDICOTS

dispersal. The genera are not all well defined. Bredemeyera and Polygala are polyphyletic and are in the process of being recircumscribed. Xanthophyllum, in the past often placed in its own family, is sister to the rest of Polygalaceae. Epirixanthes is mycoheterotrophic, otherwise unknown in the family and order; mycoheterotrophy is rare in eudicots in general. Genera and species: Polygalaceae include c. 26 genera (possibly more) and about 900 species: Acanthocladus (8), Asemeia (28), Atroxima (2), Badiera (c. 25), Balgoya (1), Barnhartia (1), Bredemeyera (c. 60), Caamembeca (15), Carpolobia (4), Comesperma (40), Diclidanthera (4), Epirixanthes (5), Eriandra (1), Gymnospora (2), Hebecarpa (9), Heterosamara (17), Monnina (c. 150), Moutabea (8), Muraltia (119), Phlebotaenia (3), Polygala (c. 275), Polygaloides (1), Rhinotropis (17), Salomonia (3), Securidaca (80) and Xanthophyllum (94).

Uses: The family is of little economic importance, apart from use in local herbal medicine. The beni seeds of Polygala butyracea, after heating and frying, can be made into malukang butter added to soups and meat dishes. Fibre of the bark of this species is made into thread used for weaving and basketry. Seeds of Securidaca longipedunculata are ground and used in soup in West Africa. Roots of Polygala arillata are fermented into an alcoholic drink in Nepal. Drupes of Carpolobia and Muraltia are sometimes eaten in Africa, and berries of Monnina have antifungal properties useful as a natural anti-dandruff shampoo and can be used for dyeing. Xanthophyllum produces a fine wood. Etymology: Polygala (πολυγαλα) is the classical Greek name for the plant. It is derived from πολύς ( polys), much, and γάλα (gala), milk, because it was assumed that animals that ate this plant would give more milk.

ROSALES Families 179 to 187 make up Rosales, the sister clade to Fagales and Cucurbitales. They are dated to have diversified c. 76 million years ago, but older estimates (up to 103 million years) are also published. It is a large order containing c. 2% of eudicot diversity. The order shows a clear development toward the loss of petals and wind pollination, occuring independently in several lineages. Most species have associations with ectomycorrhizal fungi and many with nitrogen-fixing bacteria.

179. ROSACEAE Rose family

These are annual and perennial herbs, shrubs and trees, often with spiny branches. Leaves are usually alternate, sometimes in

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basal rosettes, or distichous, rarely opposite (Lyonothamnus, Rhodotypos). Blades are entire to pinnately or palmately compound, often toothed or lobed along the margin with pinnate or palmate venation. Stipules are usually present and often fused with the petiole. Inf lorescences are usually terminal compound racemes but can be variously cymes, corymbs, false umbels or solitary flowers in leaf axils. The usually bisexual (sometime unisexual) flowers are actinomorphic, with a receptacle forming a flat, concave or tubular hypanthium bearing sepals, petals and stamens on the outer or upper margin. An epicalyx formed of fused

bracts is often present. The usually five (sometimes four to six) sepals are free or partly fused to the ovary. Petals are as many as sepals or absent in some genera. Stamens are usually numerous, rarely reduced to one or two, the filaments are normally free and anthers open by lengthwise slits. The usually semi-inferior, sometimes inferior or superior, ovary is usually adnate to the calyx tube and composed of one to many (often five), fused or free carpels forming one to several locules. The styles are free or fused into a single branched style. Fruits are variable, fleshy or dry, of one or more achenes, drupes, pomes, hips, follicles or capsules, the hypanthium

ROSALES

EUDICOTS

sometimes coloured and fleshy, sometimes the dry fruits bearing a feathery, persistent style for wind dispersal. Distribution: This cosmopolitan family is only absent from permanently frozen regions and dry habitats in Africa and Australia. They are most diverse in the temperate zones and less diverse in the lowland tropics. Phylogeny and evolution: The family is c. 76 million years old. Eocene Prunus seeds are known from China; the stem group of Pyrinae has been dated to c. 60 million years, and there was substantial divergence in that group during the Eocene and Oligocene. The taxonomy of the family has been problematic due to apomixis and hybridisation (difficult species concepts) and excessive splitting at the generic level of temperate genera of economic importance. Many genera such as Sorbus and Potentilla are polyphyletic in their traditional sense, due to the separation of economic genera such as Malus, Pyrus and Fragaria. Hybrids occur often between genera, making these difficult to keep separate. Genera are therefore in great need of recircumscription. A broad generic concept would be the easiest solution to the convoluted nomenclature in these groups, as in the formerly highly divided genus Prunus (now including the former genera Amygdalus, Armeniaca,

Rosa ‘Centifolia’, Royal Botanic Gardens, Kew, UK [179]

Cerasus, Laurocerasus, Maddenia, Padus, Persica and Pygeum). Mespilus is embedded in Crataegus, with which it makes fertile hybrids, and Neillia includes Stephanandra. The status of Poterium as separate from Sanguisorba and acceptance of Bencomia, Marcetella, Dendriopoterium and Sarcopoterium are not at all clear. Geum includes Acomastylis, Coluria, Novosieversia, Oncostylus, Orthurus, Taihangia and Waldsteinia. Potentilla should include Alchemilla, Aphanes, Comarella, Comarum, Dasiphora, Dr ymocallis, Duchesnea, Fragaria, Horkelia, Horkeliella, Ivesia, Lachemilla, Pentaphylloides, Potaninia, Purpusia, Schistophyllidium, Sibbaldia, Sibbaldianthe, Sibbaldiopsis, Stellariopsis, Tormentilla and Zygalchemilla, which would make this one of the larger genera of vascular plants (c. 1,350 species). Also Pyrus should be expanded to comprise c. 465 species and include Aria, Aronia, Chaenomeles, Chamaemelum, Chamaemespilus, Cormus, Cotoneaster, Cydonia, Dichotomanthus, Docynia, Docyniopsis, Eriobot r ya, Eriolobus, Hesperomeles, Heteromeles, Malus, Macromeles, Micromeles, Osteomeles, Photinia, Pseudocydonia, Pyracantha, Rhaphiolepis, Stranvaesia, Torminaria and Sorbus (to avoid a polyphyletic Sorbus), but these nomenclatural changes are highly unpopular due to the common application

Potentilla (Fragaria) virginiana, Helsinki Botanical Garden, Finland [179]

of many of these names. Taxonomic combinations have often not yet been made. Genera and species: Rosaceae include about 54–75 (to c. 90, depending on delimitation) genera with about 2,950 species (excluding the thousands of apomictic species in some genera). Following molecular phylogenetics, they are divided into three subfamilies: Rosoideae – Acaena (c. 100), Agrimonia (c. 13), Alchemilla (c. 1,000), Aremonia (1), Bencomia (7), Chamaerhodos (5), Cliffortia (120), Fallugia (1), Filipendula (15), Geum (c. 55), Hagenia (1), Leucosidea (1), Margyricarpus (8), Parageum (6), Polylepis (c. 20), Potaninia (1), Potentilla (c. 330), Rosa (c. 125), Rubus (c. 250), Sanguisorba (c. 15), Sarcopoterium (1), Sieversia (1) and Spenceria (1); Dryadoideae – Cercocarpus (5), Chamaebatia (2), Dryas (2) and Purshia (7); Spiraeoideae – Adenostoma (2), Amelanchier (c. 20), Aronia (c. 15), Aruncus (1), Chaenomeles (3), Chamaebatiaria (1), Chamaemeles (1), Coleogyne (1), Cotoneaster (c. 260), Crataegus (142), Cydonia (1), Dichotomanthes (1), Docynia (1), Eriobotrya (c. 17), Eriogynia (1), Eriolobus (1), Exochorda (1), Gillenia (2), Hesperomeles (11), Heteromeles (1), Holodiscus (1), Kageneckia (3), Kelseya (1), Kerria (1), Lindleya (2), Lyonothamnus (1), Macromeles (4), Malus (c. 40), Neillia (15), Neviusia (1), Oemleria

Potentilla erecta, Lake District, England, UK [179]

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ROSALES

EUDICOTS

(1), Osteomeles (1), Petrophytum (3), Photinia (c. 50), Physocarpus (10), Prinsepia (4), Prunus (c. 200), Pseudocydonia (1), Pyracantha (3), Pyrus (c. 15), Rhaphiolepis (5), Rhodotypos (1), Sibiraea (4), Sorbaria (4), Sorbus (c. 135), Spiraea (c. 80), Spiraeanthus (1), Vauquelinia (3) and Xerospiraea (1). Uses: The economic importance of this family lies mainly in the fruits of many species. Apples, peaches, pears and cherries are among the most commonly grown tree fruits and are popular around the world. Almonds (Prunus dulcis) have sweet or bitter seeds that have been cultivated in the eastern Mediterranean since ancient times, domesticated almonds appearing in Early Bronze Age archaeological sites (c. 2,500 years old). Sweet seeds (var. dulcis) are used raw or toasted for confectionery, desserts, snacks, baked goods, marzipan, macaroons, almond milk, almond butter etc. Cheap processed food that includes almonds often have this adulterated with the more bitter seeds of apricot or peach. Sweet almonds are the most commonly grown nut for human consumption, the largest production being in California, the Mediterranean and Iran. Bitter almond (P. dulcis var. amara) is used medicinally, for eardrops and in hair lotion, but this has led to some accidental cases of poisoning. Almond oil is made from either Rubus phoenicolasius in fruit, Manhattan, New York, USA [179]

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sweet or bitter almonds and used for cooking, a massage oil and medicinally. The seeds (like many other species of Rosaceae) contain the glycoside amygdalin, which after crushing or chewing of the seeds is transformed into the deadly prussic acid (hydrogen cyanide). This compound is found in low concentrations in modern sweet almond cultivars, but in high concentrations in bitter almonds, which can cause cyanide poisoning when eaten. Edible seeds of wild almond (Prunus webbii) were used in Greece before sweet almonds were introduced. Prunus also includes a number of commonly grown fruits. Best known is the peach (P. persica), which is one of the most widely grown tree fruits. It is a cultigen probably derived in China from P. davidiana, and there are many cultivars, including P. persica ‘Saturn’, the doughnut peach or paragueiro. Similar to almonds, the seeds are ground and used in Italian sweets and liqueur (amaretto). Nectarine (P. persica var. nucipersica) originated as a spontaneous mutant of peach trees, producing fruits with smooth skins. The apricot (P. armeniaca) has been cultivated in northern China since 2000 BC, although this has been disputed and a less likely Indian or Armenian origin has also been suggested. The fruit is rich in vitamin A, and it is eaten fresh, tinned or dried and

Rosa canina, Wavrons, France [179]

is made into jams and cakes, or used as flavouring for liqueur or brandy. The oil of the seed is commonly used in the cosmetics industry. Similarly used is the scented oil of Briançon apricot (P. brigantina). The plum (P. domestica) is also a cultigen of complex hybrid origin, probably involving cherry plum (P. cerasifera) and sloe (P. spinosa), but it is often hybridised in America and Asia with local native species such as P. salicina. Damson or bullace (Prunus insititia) is often considered a subspecies or variety of P. domestica, but sweet forms are sometimes called mirabelles and should not be confused with the cherry plum. Mirabelles (or myrobalan plum = ‘fragrant acorn’) or cherry plum (P. cerasifera) are native to Eurasia and also frequently planted and locally eaten, either unripe or ripe, often cooked or stewed into jams or preserves. Greengage (P. domestica subsp. italica), a delicious fruit on its own, is hybridised with apricots to create plumcots or pluots. The sweet cherry (P. avium) is usually eaten fresh and is commonly available. Sour cherry or morello (P. cerasus) is the cooking cherry, usually made into jams, pies, drinks and juices and used in certain Belgian beers (kriek). The species is of unknown origin, but it is hybridised with sweet cherries to give Duke cherries (P. ×gondouinii), which are sweeter but cook better. Traverse City (Michigan, USA) is nicknamed the ‘Cherry Rosa amblyotis in fruit, Helsinki Botanical Garden, Finland [179]

ROSALES

EUDICOTS

Sarcopoterium spinosum, Lebanon [179]

Cydonia oblonga, Ioannina, Greece [179]

Capital’, due to the large numbers of cherry orchards in that region. Minor fruits of Prunus include American plum (P. americana), chicasaw plum (P. angustifolia), Oregon cherry (P. emarginata), hortulan plum (P. hortulana), beach plum (P. maritima), ume or Japanese apricot (P. mume), wild goose plum (P. munsoniana), Canada plum (P. nigra), Chinese plum (P. salicifolia), rum cherry (P. serotina), apricot plum (P. simonii), sloe (P. spinosa), Nanking cherry (P. tomentosa) and chokecherry (P. virginiana). Sloes (P. spinosa) are still widely collected in Britain and made into sloe gin. Ume fruit is often salted in East Asia. The leaves and flowers of several cherry species are pickled in salt in Japan or are used to flavour sweets or tea. Many wild apples are known from Neolithic times, and they were often stored dried as food during the winter. The orchard apple (Malus pumila), now one of the most widely grown fruit trees, is derived from various wild species. There are currently some 2,500 cultivars of apples, of which ‘Braeburn’, ‘Bramley’, ‘Cox’s Orange’, ‘Elstar’, ‘Fuji’,

Petrophytum cinerascens, Royal Botanic Gardens, Kew, UK [179]

‘Gala’, ‘Golden Delicious’, ‘Granny Smith’, ‘Jonagold’, ‘McIntosh’ and ‘Red Delicious’, are among the most widely grown. Asian crab apple, Malus baccata, is sometimes used in hybridisation to enhance cold-hardiness of apples. Apples have a variety of uses: they are eaten fresh or cooked in baked goods and confectionery, boiled into syrup, made into calvados, cider or vinegar or dried. Malic acid and pectin, derived from apples, are common food additives. Pears (Pyrus communis) are also a widely grown fruit with over 1,000 cultivars. They are native to Eurasia, especially the Caucasus, and can be distinguished from apples (Malus) by having stone cells in their flesh, making the flesh gritty. Pears are often eaten fresh as a table fruit and are sometimes dried or cooked into purée, juice or syrup. Perry, the equivalent of apple cider made from pears, was previously popular in Britain. Vinegar is produced from pears in Western Australia. The Asian or nashi pear (P. pyrifolia) is a species from subtropical China often grown in the tropics and subtropics.

Sorbus species are more commonly used for their wood, but several species have edible fruits, most notably the Whitty pear or true service tree (S. domestica) and the wild service tree (S. torminalis), which have fruit edible after bletting (exposure to frost), whitebeam (S. aria), the fruits of which are used as a flavouring for brandy, and rowan or mountain ash (S. aucuparia), the berries of which can be cooked into jellies, although these can be too astringent for most people. Rowan jelly is a traditional accompaniment for venison. The common strawberry is Potentilla ×ananassa (formerly Fragaria), an octaploid hybrid of P. chiloensis from western North and South America and P. virginiana from the east. Many species are hybridised with other species of Potentilla to create numerous cultivars with different flavours. The garden strawberry in Europe before discovery of the Americas was the hexaploid musk strawberry (P. moschata), frequently depicted on Mediaeval tapestries. The woodland strawberry (P. vesca) is rarely cultivated but Plants of the World

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ROSALES

EUDICOTS

is commonly collected in northern Europe. Hybrids with the garden strawberry have resulted in large fruited cultivars with the wild taste (P. ×vescana). The Indian strawberry (P. indica, Duchesnea indica) produces insipid berries but is grown for ornament. Roots and leaves of silverweed (Potentilla anserina) were formerly used cosmetically to reduce skin spots and in suntan lotion; leaves were placed in shoes to reduce sweating. Rubus is another important fruit genus, which includes raspberries (R. idaeus), blackberries (including R. fruticosus, R. corylifolius, R. plicatus, R. ulmifolius and their apomictic derivatives) and cultivars, many of which are hybrids with other species (e.g. boysenberry, loganberry, silvaberry, sunberry, tayberry, tummelberry and veitchberry). Minor fruits usually collected in the wild include Alleghany blackberry (R. alleghaniensis), Arctic raspberry (R. arcticus), European dewberry (R. caesius), cloudberry (R. chamaemorus), cheeseberry (R. ellipticus), American dewberry (R. flagellaris), Andes berry (R. glaucus), Mysore raspberry (R. niveus), black raspberry (R. occidentalis), wineberry (R. phoenicolasius), roseleaf bramble (R. rosifolius), salmonberry (R. spectabilis) and Pacific dewberry (R. vitifolius). Fruits are eaten fresh or used to make jams, juices, confectionery, baked goods, flavourings and liqueurs. Cercocarpus traskiae, Santa Catalina Island, California, USA [179]

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The quince (Cydonia oblonga) is native to Anatolia and has been cultivated since ancient times. It makes a good jelly because of its high pectin content. Quinces can also be eaten after bletting. In antiquity the quince was considered a love-token and probably is the source of the golden apples of the Hesperides in Greek mythology, used in the judgment of Paris and causing the Trojan wars. The name marmalade was originally used for quince jam, derived from the Portuguese name for this fruit (marmelo). It is also fermented into a sweet, alcoholic wine and is the main ingredient for ‘mostarda di fruta’ in Italy and ‘membrillo’ in Spain. The astringent fruit of Chinese quince (Pseudocydonia sinensis) can be used like European quince. Fruit of other species including Japanese quince (Chaenomeles speciosa) and C. cathayensis can be made into jellies or added to apple sauce. Medlar (Crataegus germanica, formerly Mespilus germanica) can be eaten after frost when it is slightly rotted (bletted). Fruit of azarole (C. azarolus) tastes like apple, Chinese hawthorn (C. pentagyna) is grown as a fruit tree in China and black hawthorn (C. douglasii) is used for jelly in North America. Japanese medlar, loquat or nispero (Eriobotrya japonica) has fruit that is often eaten fresh or made into preserves. Other minor fruits are serviceberry, juneberry

Dryas octopetala, Royal Botanic Gardens, Kew, UK [179]

or saskatoon (Amelanchier spp.) and black chokeberry (Aronia melanocarpa), which are often eaten fresh or dried. Oso-berry (Oemleria cerasiformis) also has edible fruit. Redbead cotoneaster (Cotoneaster racemif lorus) is the source of a sweet substitute for manna that is high in dextrose and used in Iran and India. Inflorescences of burnet (Sanguisorba officinalis) were formerly used to make wine in England. Kageneckia lanceolata leaves yield a black dye, and the bark of Rhaphiolepis umbellata is a source of brown dye. Some species produce good wood, e.g. Cercocarpus, Crataegus, Eriobotrya, Malus, Polylepis, Prunus, Pyrus and Sorbus. Sarcopoterium spinosum is a spiny bush common around Jerusalem and was possibly used to make the crown of thorns worn by Jesus Christ. A sugar alcohol, sorbitol (E420), is found in many Rosaceae and only slowly metabolises in the human body, making it a good alternative to sugar. It is commonly found in apples, pears etc., but is now commercially made from corn syrup and commonly used in diet foods and sugar-free products. Another well-known compound originally extracted from Rosaceae (meadowsweet or swampspiraea, Filipendula ulmaria) is acetylsalicylic acid, better known as aspirin (“a” from acetyl and “spir” from Spiraea). Prunus persica, in fruit, Santa Barbara, California, USA [179]

ROSALES

EUDICOTS

Filipendula ulmaria, Lake District, UK [179]

Gillenia trifoliata, private garden, Kingston upon Thames, Surrey, UK [179]

Essential oils of roses are used for perfume and scent-making, and a jam is made of the flowers, especially R. ×centifolia. Musk-rose (Rosa moschata) is used for rose water in Iran and R. ×odorata (R. chinensis × R. gigantea) is used in scented teas. Roses became symbols of nationalism, as in the War of the Roses, the red rose of Lancaster (R. gallica ‘Officinalis’) and the white rose of York (R. ×alba ‘Semiplena’). The burnet rose (R. spinosissima) is a symbol of midsummer in the Nordic countries. Rose hips used in jam, tea and syrup are harvested from dog rose (R. canina) and eglantine (R. rubiginosa), although care must be taken to remove the seeds as these are covered in irritating hairs. Locally, fruits of other rose species are eaten as well, notably the hips of R. woodsii and Japanese rose (R. rugosa). Roses are well known as ornamental plants, and a great number of cultivars have been produced, varying from bushy and creeping to tall and climbing varieties, some scented, some not, some short stemmed for gardens, some long-stemmed and important in the cutflower industry (currently mostly concentrated in the Netherlands, Kenya, Ethiopia, Colombia and Ecuador). Roses were early symbols of secrecy, mystery and virginity. They have been in cultivation since at least 484 BC, when Herodotus discussed double-flowered

forms, but they were already grown in ancient China and are figured on Minoan murals at Knossos (R. ×richardii). The thousands of cultivars are of such complex origins that a formal classification of roses is difficult, if not impossible. Rosa aside, many other rosaceous genera are popular garden plants, some with numerous horticultural cultivars, especially the genera Acaena, Alchemilla, Aronia, Aruncus, Chaenomeles, Cotoneaster, Crataegus, Dryas, Exochorda, Geum, Gillenia, Holodiscus, Kerria, Malus, Neillia, Photinia, Potentilla, Prunus, Pyrus, Rhaphiolepis, Rubus, Sanguisorba, Sorbaria, Sorbus and Spiraea. Flowering cherries (Prunus ×yedoensis and P. ‘Sato-Zakura Group’) are the national flower of Japan, and many additional cultivars are derived from P. serrulata. Laurel cherries (P. laurocerasus and P. lusitanica) resemble bay (Laurus nobilis, Lauraceae) and are often planted to the same effect; they are so similar to the leaves of laurel that they are even used in “laurel” wreaths, but they are poisonous and should not be used in cooking. Record: At 5,100 m in the Bolivian Andes, trees of Polylepis tarapacana can be found, which is the highest elevation for any tree to grow.

Pyrus amygdaliformis, Royal Botanic Gardens, Kew, UK [179]

Carnivory: Many members of Rosaceae are covered with glandular, often sticky hairs or extraf loral nectaries. Insects often become entrapped in these, and it has widely been assumed that these glands have a defensive capability. However, members of the Potentilla arguta complex (sometimes placed i n Dr ymocallis, including P. glandulosa and P. rupestris) appear capable of not only trapping insects but also of digesting them and absorbing the released nutrients. These species occur in habitats that support other carnivores, especially Drosera, and fit the syndrome of conditions associated with carnivory, but surface microbes as the source of the detected digestive enzymes cannot be ruled out. Rosehip cruelty: Until at least the 1960s, school children in northern England used to torment each other by putting seeds from rosehips down the backs of their shirts, taking cruel advantage of the irritating hairs. Etymology: Rosa is the Latin name for a rose, possibly derived from the Greek rhodon (ρόδον), a rose, but how this was derived remains a mystery. It possibly has an ancient Indo-European root. Plants of the World

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Barbeya oleoides, female flower, Saudi Arabia (TM) [180]

EUDICOTS

Barbeya oleoides, male flowers, Barbeya oleoides in fruit, Saudi Saudi Arabia (TM) [180] Arabia (TM) [180]

180. BARBEYACEAE Elm-olive family

With greyish opposite lanceolate leaves, it has a superficial resemblance to the cultivated olive (Olea europaea, Oleaceae), but it is not related. The flowers are wind-pollinated. Genera and species: This family includes the single species, Barbeya oleoides.

These are unisexual trees with opposite petiolate leaves without stipules. Blades are oblong-lanceolate and white-hairy below. The unisexual flowers are actinomorphic and have three to four nearly free tepals. Male flowers have six to 12 stamens with short free filaments and basifixed anthers that open lengthwise. Female flowers have a superior ovary that is sometimes shortly stalked and is composed of a single carpel or of two or three fused carpels. The apical style is simple with a linear papillate stigma. Fruits are dry nutlets enclosed in the persistent tepals. Distribution: The family is found in dry forested slopes on both sides of the Gulf of Aden in Yemen, Somalia, Eritrea and Ethiopia. Phylogeny and evolution: In the past, the single genus Barbeya was placed in Urticales due to similarities with Ulmaceae, but molecular evidence placed it close to Dirachmaceae.

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Etymology: Barbeya is named for Swiss botanist and politician Wiliam Barbey (1842– 1914). He botanised in the Mediterranean region and was founder of the journal Bulletin de l’Herbier Boissier.

181. DIRACHMACEAE Rachman family

Dirachma socotrana, illustration based on herbarium specimens [181]

hypanthium, together with the same number of free, white petals that bear appendages at the base inside the flower. Stamens are the same number as petals and basally fused to them. Anthers are basifixed and open by lengthwise slits. The semi-inferior ovary has as many locules as sepals and is densely hairy and radially lobed. The style is glabrous at the top and has a club-shaped to cylindrical stigma. The fruit is a septicidal capsule with long hairs inside and smooth seeds. Distribution: This family is restricted to Somalia and the island of Socotra, where it often grows on limestone soil. Both species are rare and highly endangered. Phylogeny and evolution: The placement of Dirachma has been considered puzzling on morphological grounds; Malvaceae, Flacourtiaceae (Samydaceae) and Geraniaceae have previously been proposed as related. Molecular phylogenetics placed Dirachma as sister to the similarly puzzling Barbeya in Rosales, each in their own family.

This is a family of shrubs and trees with simple, glandular-peltate hairs. They have simple, alternate, petiolate leaves with lineartriangular stipules. Blades have serrate to lobed margins, and venation is pinnate. The bisexual, actinomorphic flowers are borne singly in the leaf axils on a bracteate pedicel. The five, six or eight sepals are free, but placed on a

Genera and species: The single genus Dirachma contains two species: D. socotrana and D. somalensis. Etymology: Dirachma is possibly derived from rachman, a Socotran vernacular name for this scented shrub, although no derivation was given when the genus was described.

ROSALES

EUDICOTS

basal part of the hypanthium and therefore seemingly inferior, with a single locule. The single style terminates the ovary, and the stigma is on one side of the style. Fruits are achenes that are enveloped by the persistent hypanthium, which becomes fleshy or mealy and often has a hard inner layer, resembling a drupe or berry with a single seed.

182. ELAEAGNACEAE Oleaster family

These are deciduous and evergreen shrubs and small trees that are sometimes laxly climbing and usually have nitrogen-fixing root nodules. Most parts of these plants have distinctive silvery or coppery peltate scales and/or stellate hairs, and branches are sometimes spine-tipped. The petiolate simple leaves are alternate or opposite (rarely whorled) and lack stipules. Blades have entire margins and are densely covered in peltate hairs below with pinnate venation. Inflorescences are small racemes or fascicles, or the flowers are solitary in the leaf axils. The actinomorphic flowers are usually bisexual, sometimes unisexual and then the plants are dioecious. The hypanthium is constricted above the ovary. Two to six (to eight) sepals are fused to the hypanthium and often petal-like, the petals being absent. Four to eight stamens are inserted in the hypanthium tube, in bisexual flowers as many as the lobes, in male flowers twice as many. Filaments are free, and anthers are dorsifixed or basifixed, opening by lengthwise slits. The ovary is superior but tightly enclosed by the

Hippophaë rhamnoides in fruit, Royal Botanic Gardens, Kew, UK [182]

Distribution: This family of temperate and Arctic North America and temperate Europe and Asia extends into tropical Malesia, Sri Lanka, the Mascarenes and northern Australia. Phylogeny and evolution: Previously this family was placed in Rhamnales, but this order is now part of an expanded Rosales. They are indeed closely related to Rhamnaceae. The oldest fossil pollen attributed to Elaeagnaceae is known from the Eocene of Central Asia. Several Oligocene and Miocene fossils from Europe and North America are known.

or oleaster (Elaeagnus angustifolia), goumi (E. multiflora) and buffaloberry (Shepherdia argentea). Several species are also grown as ornamental garden shrubs. Their roots are able to fix atmospheric nitrogen using Frankia bacteria, making it possible for plants to grow well on poor soils. Therefore some species have been used for land reclamation and soil stabilisation, and they easily naturalise, although due to their beneficial qualities they are not often considered problematic. Etymology: Elaeagnos (ελαίαγνος) is a Greek name for a type of willow tree and is in turn derived from ελώδες (helodes), growing in swamps, and αγνός (agnos), pure, in reference to the white fruits of a willow. The name was later applied to the oleaster (E. angustifolia), which superficially resembles a willow.

183. RHAMNACEAE Buckthorn family

Genera and species: This is a family of three genera and c. 50 species: Elaeagnus (c. 40), Hippophaë (5) and Shepherdia (3). Uses: Sea buckthorn (Hippophaë rhamnoides) fruits are rich in vitamin A, C and E and one of the highest valued for health foods. They have been used for centuries in Europe and Asia to make jams, jellies, juices and drinks. Fruits of most other species are edible, although often astringent. Other species that are sometimes eaten are Russian olive

Elaeagnus umbellata, Royal Botanic Gardens, Kew, UK [182]

A family of deciduous (mostly) and evergreen trees and shrubs, they are rarely herbs (Crumenaria) or climbers, sometimes twining or tendrillate. Stems are sometimes spiny, or

Shepherdia canadensis, Helsinki Botanical Garden, Finland [182]

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ROSALES

EUDICOTS

Colletia armata, Royal Botanic Gardens, Kew, UK [183]

Zizyphus jujuba, Bucharest Botanical Garden, Romania [183]

Alphitonia neocaledonica, New Caledonia [183]

Rhamnus alaternus in fruit, Sicily, Italy [183]

short branches can be spine-tipped. Leaves are simple, usually alternate, sometimes opposite, and stipules are usually free, sometimes fused across the petiole or between petioles or transformed into spines, absent in most Phylica species. Tendrils are circinnate when present. Inflorescences are usually axillary cymes, sometimes compound panicles or reduced to fascicles or single flowers. The bisexual or unisexual flowers are actinomorphic and often small (< 6 mm). A hypanthium is usually present, cup-shaped or tubular. The four or five sepals are fused with the hypanthium rim. The four or five free petals are inserted in the mouth of the hypanthium, usually smaller than the sepals and sometimes absent, often forming a hood over the stamens. The four

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Ceanothus cyaneus, Rancho Santa Ana, California, USA [183]

Phylica pubescens, Adelaide Botanic Garden, South Australia [183]

or five stamens have thin filaments fused to the base of the petals with small anthers that open by lengthwise slits. A fleshy disc is usually present between the stamens on the hypanthium. The superior to inferior ovary is bi- to tetra-loculate and tipped with a shortly lobed style. Fruits are usually drupes, sometimes capsules with wings. Distribution: The family has a nearly cosmopolitan distribution, but is absent from the great deserts and boreal zones. Phylogeny and evolution: Because rhamnaceous pollen is known from c. 90 million-year-old deposits, the assumption can be made that the family was already

cosmopolitan before the continents were widely separated, which may explain some of the wide geographical disjunctions in some clades. The existence of Phylica species on several volcanic islands of different ages in the Atlantic and Indian Oceans allowed the radiation in this group to be dated using one of the early well-documented examples of a calibrated molecular clock. It also shows that long-distance dispersal is not uncommon in this family. Genera and species: Rhamnaceae include 55 genera and c. 1,040 species: Adolphia (1), Alphitonia (c. 13), Alvimiantha (1), Ampelozizyphus (1), Auerodendron (7), Bathiorhamnus (2), Berchemia (c. 20), Berchemiella (2),

ROSALES

EUDICOTS

Blackallia (1), Ceanothus (55), Colletia (5), Colubrina (33), Condalia (18), Crumenaria (6), Cryptandra (c. 55), Dallachya (1), Discaria (6), Doerpfeldia (1), Emmenosperma (3), Gouania (c. 50), Granitites (1), Helinus (5), Hovenia (7), Jaffrea (2), Karwinskia (16), Kentrothamnus (1), Krugiodendron (2), Lasiodiscus (7), Maesopsis (1), Nesiota (1, extinct), Noltea (1), Paliurus (5), Phylica (188), Pleuranthodes (2), Pomaderris (c. 75), Pseudoziziphus (2), Reissekia (1), Retanilla (4), Reynosia (c. 15), Rhamnella (10), Rhamnidium (12), Rhamnus (c. 120), Sageretia (35), Sarcomphalus (32), Schistocarpaea (1), Scutia (4), Siegfriedia (1), Smythea (10), Spyridium (45), Stenanthemum (c. 35), Trevoa (1), Trichocephalus (1), Trymalium (13), Ventilago (c. 40) and Ziziphus (c. 66). Uses: Several species produce edible fruit. Chinese jujube or zǎo (Ziziphus jujuba) and Indian jujube or ber (Z. mauritiana) are grown commercially and usually sold dried, resembling dates. Ziziphus lotus is the lotus fruit of the ancients, eaten by the ‘lotophagi’ mentioned in the Odyssey and ‘sidr’ (‫ةردس‬ ‫ )هتنملا‬or lote-tree of the Quran. Other species of Ziziphus, Condalia, Reynosia and Sageretia and Karwinskia humboldtiana also have edible fruits. Mistol fruits (Ziziphus mistol) are brewed into an alcoholic drink in the Andes. The raisin tree (Hovenia dulcis) has swollen fruit stalks that turn red after frost and are edible. Leaves of Chinese sweet plum (Sageretia thea) are used as a tea in Vietnam. Ventilago calyculata seeds can be pressed for cooking oil. Several species have saponins; leaves of the Australian soap tree (Alphitonia

Ulmus laevis fruits, Turku, Finland [184]

excelsa) are used as a soap substitute, and some Gouania species are an alternative to commercial shampoo. Ampelozizyphus, Colletia and Colubrina also yield soap. Dyes are also common in this family. Several Rhamnus species yield yellow or green dyes, traditionally used in Europe and East Asia. Ventilago madraspatana bark yields a red dye, and kauila (Alphitonia ponderosa) bark yields a bluish dye. Wood of the latter was also used for carving and weapons in Hawaii. Some species have extremely hard and heavy wood. Others produce fine timber for construction, woodworking etc. Charcoal of alder buckthorn (Rhamnus frangula) was traditionally used when making gunpowder. Ziziphus spinacristi is one of the contenders for the tree from which Jesus’ crown of thorns could have been made. The shoots of the rattan vine (Berchemia scandens) are sometimes used for wicker work in the United States, and several species of Ceanothus, Colletia, Noltea and Pomaderris are grown as ornamental garden shrubs. Phylica arborea (Tristan da Cunha and Amsterdam Island) is the only woody native species on Tristan da Cunha (South Atlantic Ocean) and the other islands in the Archipelago, and it is known as the island tree and has been used for firewood etc. Its dispersal between the islands in the archipelago and to Ile Amsterdam (southern Indian Ocean) represent amazing repeated inter-island dispersals, the more so because the seeds do not survive in sea water. Etymology: Rhamnos (ραμνος) is the Greek name for a prickly shrub, such as a buckthorn.

184. ULMACEAE Elm family

These trees and shrubs have watery sap and alternate and opposite leaves that are arranged in a plane along the stem (distichous). Stipules grow in pairs on either side of the petioles, and leaf blades are simple, strongly serrate and basally asymmetrical. Venation is pinnate and, in all but Ampelocera, secondary veins terminate in teeth. Inflorescences are axillary cymes or fasciculate aggregates of flowers in the leaf axils. The unisexual or bisexual flowers are green or brown, the perianth spirally arranged with (two to) four to eight (or nine) fused or free tepals. Stamens are the same number as the tepals (rarely more) with dorsifixed, often versatile anthers that open by lengthwise slits. The superior ovary is composed of two fused carpels forming a unilocular pistil topped with two linear styles that can be simple or bifurcate and a stigma on the inner surface. Fruits are flattened nuts, drupes or samaras. Distribution: Ulmaceae are primarily found in temperate areas in the Northern Hemisphere, but species extend into the

Ulmus laciniata, Helsinki Botanical Garden, Finland [184]

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ROSALES American and African tropics and subtropical East Asia to Sumatra, Sulawesi and the Sunda Islands. Phylogeny and evolution: Fossilised fruits and leaves of Ulmus are known from 50 million-year-old Eocene deposits in northeastern China, suggesting a greater age of the family than initially assumed. The family has two clades, one predominantly tropical and the other mostly temperate. Some genera of the former subfamily Celtidoideae are now part of Cannabaceae. Genera and species: Ulmaceae include seven genera and c. 45 species: Ampelocera (10), Chaetachme (1), Hemiptelea (1), Phyllostylon (2), Planera (1), Ulmus (c. 25) and Zelkova (4). Uses: Elm wood (Ulmus spp.) is rot-resistant and therefore was often used in waterworks, wells, coffins etc. Due to Dutch elm disease (caused by the Asian fungus Ophiostoma ulmi, dispersed by the elm bark beetle Scolytus multistriatus), large plantations are not viable, and elm trees have largely disappeared from the landscape and as street trees in Europe and North America in the latter part of the 20th century. Resistant strains have, however, now been found but are much more expensive to propagate. Keaki (Zelkova

EUDICOTS

serrata) is also cultivated for ornament and wood. The wood of Chaetachme aristata is sometimes made into musical instruments, and the fine-grained wood of baitoa (Phyllostylon brasiliensis) is sometimes used for turning. Etymology: Ulmus is the classical Latin name for an elm tree. It originated from the Akkadian u’ulum, to bind, or from ProtoIndo-European el, brown.

185. CANNABACEAE Hemp family

This is a family of trees, shrubs, perennial and annual herbs and vines with watery sap. Plants are unisexual, bisexual or polygamous. The petiolate leaves are alternate or opposite, usually in two ranks, simple, deeply lobed or palmately compound with paired stipules. Venation is palmately pinnate. The terminal

or axillary inflorescences are cymes, panicles or fascicles, or sometimes the flowers solitary. The brown or green bisexual or unisexual actinomorphic flowers usually have four to eight (sometimes fewer or more) fused tepals. Stamens are usually as many as tepal lobes. Anthers are basifixed or dorsifixed and open by lengthwise slits. The superior ovary is composed of two fused carpels topped with two linear, simple or bifurcating styles with the stigmatic area along their inner surfaces. Fruits are nuts, drupes, achenes and samaras. Distribution: This worldwide family is found throughout the Americas, Europe and North Africa, Sub-Saharan Africa, Madagascar, Yemen, Turkey, Caucasus, Central Asia throughout tropical Asia, Australia and Pacific islands. Phylogeny and evolution: Cannabaceae have always been associated with Moraceae and Urticaceae, with which they share many characters. Molecular results have shown that Ulmaceae in the wider sense (based on an interpretation of their morphology) were polyphyletic, resulting in several former genera of Ulmaceae (mainly from subfamily Celtidoideae) being placed here. The oldest fossil Cannabaceae are known from Tertiary deposits.

Cannabis sativa, legal medicinal marijuana plantation in Portland, Humulus lupulus, male, Canbury Gardens, Humulus lupulus, female, Canbury Oregon, USA (DH) [185] Kingston upon Thames, UK [185] Gardens, Kingston upon Thames, UK [185]

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Lozanella enantiophylla, San Francisco Botanical Garden, USA [185]

ROSALES

EUDICOTS

Genera and species: Cannabaceae include 9 genera and c. 100 species: Aphananthe (3), Cannabis (1), Celtis (c. 60), Gironniera (6), Humulus (3), Lozanella (2), Parasponia (5), Pteroceltis (1) and Trema (c. 15). Uses: Use of cannabis (Cannabis sativa and its many varieties) as a recreational drug, due to the psychoactive resin tetrahydrocannabinol (THC), can be dated back to at least 440 BC. Herodotus, for example, mentioned its use: ‘The Scythians ... take some of this hempseed and ... throw it upon the red-hot stones; immediately it smokes and gives out such a vapour as no Grecian steam-bath can exceed. The Scyths delightedly shouted for joy.’ However, hemp has been cultivated for 8,500 years for its fibre, used in fabric and rope, and oil in its seeds, for food, varnish, soap and fuel, although because of legislative restrictions it is much less frequently cultivated for these purposes nowadays. Cannabis is currently the largest cash crop in the United States, although mostly grown illegally. Nevertheless, legalisation of cannabis is progressing (several countries and states have now legalised the drug) and this has proven to help lift the cultivation, trade and use of mild drugs out of the criminal circuit. Taxes collected from this trade can then be used to alleviate addictions to this and more damaging addictive “soft” drugs (like sugar, alcohol and tobacco). Hops (Humulus lupulus) were already used to flavour beer in Europe in the 8th century, gradually replacing the use of other plants such as bog myrtle (Myrica gale, Myricaceae); hops have been in cultivation since the 1200s. Hops contain lupulin, a pseudo-oestrogen causing the infamous beer belly. The young shoots of hops were Silk moth caterpillars eating mulberry (Morus alba) leaves, market in Shanghai, China [186]

harvested and eaten like asparagus since Roman times, and this is still done in Mediterranean cuisine. Celtis produces strong wood (e.g. beaver wood) that is used for furniture, and the rough leaves of Aphananthe aspera can be used as sand paper. Quickly growing guacimilla (Trema micrantha) was used for pre-Columbian bark cloth and is now harvested for export and tourism. Etymology: Cannabis is a Latinised form of the Greek name of the plant, κάνναβης (kannabis), which originally came from Scythian into Greek. It is possibly a compound of a Fenno-Ugric word for hemp, kene, and the Proto-Germanic pish, to burn. Its IndoEuropean roots (qunabu, hanapiz) may be the source of the English words canvas and hemp.

186. MORACEAE Mulberry family

leaving a scar on the stem when dehiscing. Inflorescences are bisexual or unisexual axillary racemes or cymes, often fused into heads or cup- or urn-shaped structures with the flowers on the inside, sometimes subtended by bracts; flowers sometimes solitary. The unisexual flowers have a simple perianth with usually four (sometimes up to ten) fused tepals or perianth absent. Male flowers have (one to) four (to six) stamens with straight or curved, free or fused filaments. Anthers open explosively when the stamens are curved outward, or the anthers are fused into peltate structures with a circumscissile opening. Pistillodes are often present in male flowers. Female flowers are without staminodes, the superior or inferior ovary is topped with one or two styles and stigmas. The fruit is a drupe, rarely a dry achene, usually enveloped in a fleshy perianth and often immersed in the flashy receptacle, sometimes the inflorescence fused (e.g. a syncarp in Artocarpus, Ficus). Distribution: This widespread family occurs worldwide, but is absent from northern and northwestern North America, northern Eurasia, the Sahara, southern Australia, New Zealand and Antarctica.

These are terrestrial and epiphytic trees, shrubs, climbers and perennial herbs, usually with milky sap. The usually alternate (rarely opposite or whorled) leaves are simple, sometimes palmately or pinnately compound, and blades are entire, toothed or deeply pinnately or palmately lobed; venation is pinnate, palmate or three-veined. Stipules are usually present and fused around the bud,

Phylogeny and evolution: Moraceae are closely related to Urticaceae, and indeed some genera such as Cecropia have been moved between the two families. Together with Cannabaceae and Ulmaceae, they form the ‘urticalean’ clade in Rosales. The heterogeneous genus Ficus has diversified together with its specialised pollinator, the fig wasp (superfamily Chalcidoidea), with many fig species having specialised wasps only found in association with their host.

Dorstenia christenhuszii, Royal Botanic Gardens, Kew, UK [186]

Broussonetia papyrifera, Royal Botanic Gardens, Kew, UK [186]

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ROSALES

Artocarpus altilis, Moorea, French Polynesia [186]

Ficus austrocaledonicus, New Caledonia [186]

Ficus benghalensis, a strangler fig, Tahiti, French Polynesia [186]

Genera and species: Moraceae include 37 genera and c. 1,180 species: Antiaris (1), Antiaropsis (2), Artocarpus (45), Bagassa (1), Batocarpus (3), Bleekrodea (3), Bosqueiopsis (1), Brosimum (15), Broussonetia (8), Castilla (3), Clarisia (3), Dorstenia (107), Fatoua (2), Ficus (c. 850), Helianthostylis (2), Helicostylis (7), Hullettia (2), Maclura (11), Maquira (4), Mesogyne (1), Milicia (2), Morus (13), Naucleopsis (22), Parartocarpus (2), Perebea (9), Poulsenia (2), Prainea (2), Pseudolmedia (9), Scyphosyce (2), Sorocea (14), Sparattosyce (2), Streblus (14), Treculia (3), Trilepisium (1), Trophis (9), Trymatococcus (2) and Utsetela (2).

Captain William Bligh was sent on the H.M.S. Bounty to bring plants from Tahiti to Jamaica to establish a plantation there. Crew members married Tahitian women and were upset when they were forced to sail for the West Indies, resulting in the infamous mutiny on the Bounty. Captain Bligh managed to get to Indonesia in a longboat, whereas the mutineers retrieved their wives from Tahiti and settled on Pitcairn Island. On a second voyage, the breadfruit finally made it to the West Indies, where the slaves refused to eat the fruit. It is still commonly cultivated in Jamaica. The related jackfruit, Artocarpus heterophyllus, is commonly cultivated around the tropics, and the sweet pulp is sometimes canned. Chempedak (A. integer) is used like breadfruit in Malaysia. Seeds of breadnut (Brosimum alicastrum) are eaten in tropical America, and elephants distribute African breadfruit (Treculia africana), which has large seeds that are eaten locally by people. Mulberries are also commonly grown for their fruit, especially southern mulberry (Morus australis) and black mulberry (M. nigra), both cultivated for millennia. Broussonetia luzonica inflorescences are used in Philippine cuisine. Fruit of Clarisia is also eaten, and the fruit of Maclura tricuspidata is valued in China and Korea. Delicate paper is prepared from the paper mulberry, B. papyrifera, in East Asia, and the fibre is used to make cloth in many Pacific islands. Kozo, B. kazinoki, is used in Japan for paper making, often erroneously called ‘rice paper’. Its fruits and leaves can be eaten after cooking. The white mulberry (Morus alba), which also has edible fruit, is the foodplant for farmed caterpillars of the silk moth (Bombyx mori), and the bark of this

species was used to make paper money during the Ming Dynasty in China. Many species yield valuable timber, especially ipoh (Antiaris toxicaria), jak (Artocarpus heterophyllus), letterwood (Brosimum guianense), satiné (B. rubescens), iroko (Milicia excelsa) and osage orange (Maclura pomifera). Ulé rubber (Castilla elastica) and the rubber fig (Ficus elastica) were formerly important rubber sources. Buddha had his true insights beneath a peepul tree, Ficus religiosa, which is now sacred to Buddhists and Hindus, and its leaves are used for miniature paintings. Bark of Helicostylis is hallucinogenic and used for witchcraft in the Guianas. The motile stamens and tepals of Morus alba move at half the speed of sound, releasing the pollen in the wind, being the most quickly moving parts of any plant. Many species are cultivated as ornamentals, especially Ficus benjamina, which is the most common leafy houseplant. Many species of figs are planted as shade trees in the tropics. The oldest known planted tree is a sacred fig (Ficus religiosa) in Sri Lanka, called Jaya Sri Maha Bodhi, which was planted in 288 BC. Several Dorstenia species are grown in specialist collections for their intriguing inflorescences, resembling figs turned inside out.

Uses: The common fig, Ficus carica, is frequently grown as a fruit tree. It was probably already cultivated in pre-pottery Neolithic times in the Levant, c. 11,300 years ago. It naturalised early around the Mediterranean, where it commonly grows on old masonry and in archaeological sites. The sycamore of the Bible, or mulberry fig, Ficus sycomorus, had been domesticated for its fruit in Ancient Egypt (and is believed to be the sycamore that Zacchaeus climbed to get a better view of Jesus, as reported in the New Testament). Figs are now cultivated in many parts of the world, and in the tropics the fruits of many other Ficus species are also eaten, sometimes on a commercial scale. The common breadfruit, Artocarpus altilis, is an important food source in tropical Asia and the Pacific. The value of the fruit in the Pacific was first observed by Captain James Cook in Tahiti in 1769, and in England it was thought to be a useful crop for feeding slaves in the British colonies in the West Indies. 274

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Etymology: Morus is the Latin name for the black mulberry tree, M. nigra. It is derived from Ancient Greek μορον (moron), a mulberry (Morus) or blackberry (Rubus, Rosaceae). The Spanish mora is used for both fruits and can cause confusion where both kinds of fruit occur.

ROSALES

EUDICOTS

187. URTICACEAE Nettle family

This family includes terrestrial and epiphytic, bisexual and unisexual, annual and perennial herbs, shrubs, lianas and trees, sometimes with arial and stilt roots. Opposite or alternate, spirally arranged leaves have free and lateral or interpetiolar and fused, often amplexicaul, stipules that fall when the leaf develops. Blades are simple or pinnately or palmately lobed, the margins entire and venation pinnate or palmate. Petioles can be basally or centrally attached and then the blade peltate, or the petiole absent and the blade sessile. Stinging hairs are present in some species. Inflorescences are often

Obetia ficifolia, Réunion [187]

Cecropia hispidissima, Ecuador [187]

unisexual in pairs in the leaf axils and can be cymes, racemes, heads, false umbels or spikes, clustered and sometimes subtended by a bract. The unisexual, rarely bisexual, f lowers are actinomorphic or (especially female flowers) zygomorphic. Male flowers have (one or) two to five free or fused tepals and one to five stamens, the filaments curved, but suddenly reflexing and ejecting pollen. Anthers are basifixed and open by lengthwise slits. Female flowers have two to five tepals that are free or fused into a tube, often unequal, rarely absent, and staminodes are sometimes present and can be scalelike, often important for fruit dispersal. The superior ovary is composed of a single carpel and topped with a single variably shaped sessile or stalked stigma. Fruits are achenes that are usually free, sometimes fused with a fleshy perianth that develops at fruit maturity. Distribution: The family has a global distribution apart from dry or permafrost areas. They are most numerous in the wet tropics but can also be common in temperate regions.

Pilea microphylla, Guadeloupe [187]

Urtica dioica, near Turku, Finland [187]

Phylogeny and evolution: Fossil leaves of Urticaceae have been reported from Upper Cretaceous deposits, and a stamen and tepal of a putative member of Urticaceae have been found in Eocene Baltic amber. Cecropiaceae have traditionally been associated with Moraceae or placed in that family in the past on the basis of the female structures, but they were already included on morphological grounds in Urticaceae in the 1960s. Cecropiaceae were considered a morphological link between Urticaceae and Moraceae, but molecular studies have placed Cecropia and its relatives in Urticaceae. The latter are polyphyletic if Cecropiaceae are excluded. Genera and species: Urticaceae include 53 genera and c. 1,250 species: Aboriella (1), Archiboehmeria (1), Astrothalamus (1), Australina (1), Boehmeria (c. 80), Cecropia (c. 75), Chamabaina (2), Coussapoa (50), Cypholophus (15), Debregeasia (4), Dendrocnide (37), Didymodoxa (2), Discocnide (1), Droguetia (7), Elatostema (c. 300), Forsskaolea (6), Gesnouinia (2), Forsskaolea angustifolia, Bergius Botanical Garden, Stockholm, Sweden [187]

Laportea canadensis, Wisconsin, USA [187]

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ROSALES Gibbsia (2), Girardinia (2), Gonostegia (5), Gyrotaenia (6), Hemistylus (4), Hesperocnide (2), Laportea (21), Lecanthus (1), Leucosyke (35), Maoutia (15), Meniscogyne (2), Musanga (2), Myrianthus (7), Myriocarpa (18), Nanocnide (2), Neodistemon (1), Neraudia (5), Nothocnide (5), Obetia (8), Oreocnide (15), Parietaria (c. 10), Petelotiella (1), Phenax (12), Pilea (c. 200), Pipturus (30), Poikilospermum (20), Pourouma (25), Pouzolzia (70), Procris (16), Rousselia (3), Sarcochlamys (1), Sarcopilea (1), Soleirolia (1), Touchardia (1), Urera (35) and Urtica (80). Uses: The boiled young shoots of Girardinia, Laportea and Urtica are eaten as vegetables and in soup. Common nettle (Urtica dioica) is used as flavouring in Dutch nettle cheese, and Cornish yarg cheese is traditionally wrapped in these leaves. Several species are cultivated for their stem fibre, which is of high quality and used to make cloth, fishing

EUDICOTS

nets and ropes. For instance fibre of U. dioica was used to make khaki cloth for military uniforms, notably in the Second World War. Ramie (Boehmeria nivea) is widely cultivated in East Asia and is the toughest, longest and silkiest of all vegetable fibres. Similarly fibre is collected from Girardinia diversifolia subsp. triloba, Sarcochlamys pulcherrima, Urtica dioica and many other species. Orange wild rhea, Debregeasia longifolia, also has good fibre, and its edible fruit is known as janatsi or yanagi. Fruits of uvilla, Pourouma cecropiifolia, taste like grapes. Tubers of Pouzolzia tuberosa are eaten in tropical Asia. The fruit of corkwood, Myrianthus arboreus, is eaten in Africa, and its wood is lighter than cork. Several species of Pilea and Soleirolia soleirolii are widely cultivated as ornamentals.

stinging hair breaks off upon contact, leaving a sharp point that readily pierces skin and allows the irritating liquid inside the hair to enter flesh. The hollow hairs act like a miniature hypodermic needle. Despite the painful result, nettles (Urtica dioica) were pulled out using bare hands into the 20th century, in the belief that the resulting inflammation was a useful treatment against arthritis and rheumatism. As quickly growing pioneer trees in the Neotropics, Cecropia attracts fire ants with glycogen-rich food bodies (Mullerian bodies) at the base of their petioles. The ants enter the plant there and make the hollow stems their home, which of course they defend against intruders. Even though these species do not have stinging hairs, this is another one to handle with care.

Do not touch this nettle: Stinging hairs in Urticaceae have a distinct bulbous or cylindric base and a stiff, translucent apex. The tip of the

Etymology: Urtica is the Latin name for nettle, first used by Plinius. It probably originated from Latin urere, to sting or burn.

FAGALES Families 188 to 195 make up Fagales. An order with this name has been included in many morphology-based classifications, often associated with other sets of wind-pollinated families in the so-called higher Hamamelidae. In that circumscription, Hamamelidae were, however, grossly polyphyletic. In modern molecular-based classifications, Fagaceae and their relatives are associated with other families in which nitrogen-fixation via root nodules occurs. The order is then associated with Curcurbitales, a mostly animal-pollinated order. There are fossils that exhibit morphological traits that are mixtures of those typical of these two orders.

188. NOTHOFAGACEAE Roble family

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Nothofagus antarctica, female branch, Royal Botanic Gardens, Kew, UK [188]

Nothofagus obliqua, male flowers, Royal Botanic Gardens, Kew, UK [188]

FAGALES

EUDICOTS

This is a family of evergreen and deciduous, bisexual trees and shrubs. The alternate leaves are usually in two ranks (distichous) or spirally arranged, and stipules are attached basally or peltate. Blades are simple, entire, serrate to doubly serrate, and the lamina is covered with globular-glandular hairs. Inflorescences are one- to three-flowered dichasia, up to seven (rarely up to 15) in female inflorescences. Male flowers are sessile or shortly stalked and have a fused, regularly splitting perianth composed of four to six fused tepals. The four to 90 stamens per flower have long, flexible filaments and basifixed anthers with a protruding connective. Female flowers are sessile or shortly stalked with a persistent involucre; they often have staminodes. The inferior ovary is tipped with short styles and usually curved stigmas. Fruits are two- or three-angled nuts surrounded by a one- to four-valved cupule with lamellar appendages. Distribution: This family can be found in southern South America, New Guinea, New Caledonia, eastern and southern Australia (New South Wales, Victoria, Tasmania) and New Zealand. Lithocarpus variolosus, Royal Botanic Gardens, Kew, UK [189]

Phylogeny and evolution: Nothofagus was previously included in Fagaceae, but it was found to be sister to all other Fagales in molecular studies. Therefore they have been placed in their own family. This family had a wider distribution during the Cretaceous and Tertiary in the Southern Hemisphere, and fossils are known from Australia, New Zealand, Antarctica, South America and the Falkland Islands. It was suggested that the current distribution is the result of continental drift, but modern clades evolved after the breakup of Gondwana. It is therefore more likely that the family reached this distribution via long-distance dispersal; for instance they reached New Zealand by long-distance dispersal c. 30 million years ago. Genera and species: The sole genus is Nothofagus with 43 species. The genus has been subdivided into subgenera (sometimes accepted as genera): Fuscospora (6), Lophozonia (7), Nothofagus (5) and Trisyngyne (25), but these are not widely accepted as genera. Nothofagus in the broad sense is monophyletic, and the advantages of splitting it are unclear, making this subdivision not strictly necessary.

Quercus crassifolia, Royal Botanic Gardens, Kew, UK [189]

Uses: Timber of southern beech or roble, Nothofagus, is hard and strong with a close grain and is valued especially for cabinetry and turning. It is also locally used for general construction and railway sleepers. Etymology: Nothofagus is composed of the Greek νόθος (nothos), false, and Latin fagus, a beech tree.

189. FAGACEAE Beech family

These trees (rarely shrubs) are bisexual and deciduous or evergreen. Simple leaves are usually alternate, rarely in whorls of three (Trigonobalanus), and have entire, serrate, sinuate or lobed blades with pinnate venation; they are often covered with branched or

Fagus sylvatica, Royal Botanic Gardens, Kew, UK [189]

Fagus sylvatica, Bix Bottom, UK [189]

Quercus ilex, Mt Saint-Victoire, France [189]

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FAGALES stellate hairs. Stipules are free and deciduous. Inflorescences are usually reduced spikes, heads or cymules, sometimes with solitary flowers in the leaf axils and male flowers sometimes in pendent catkins. The unisexual flowers are actinomorphic and have a usually six-, sometimes four- to nine-lobed perianth. Male flowers have usually 12, sometimes four to 20, stamens with free, filiform filaments and linear to kidney-shaped, dorsifixed or basifixed anthers opening by lengthwise slits. A pistillode is sometimes present among the stamens. Female flowers are surrounded by a cupule, the perianth usually six-lobed, but it is sometimes poorly developed. Staminodes can be absent, but when they are present there are six to 12 surrounding the ovary. The superior ovary is two- to six-loculed and has as many short styles on the top as there are locules, the stigmas capitate or linear and covering the inside of the styles. The fruit is a one- to threecelled nut that can be round, three-angled or winged, but always attached to a sessile or stalked cupule that can be saucer- or cupshaped or encloses the fruit, which can be dehiscent and variously ornate or not. Distribution: Fagaceae are distributed across the temperate zones of the Northern Hemisphere, extending into the tropics in Central America and northwestern South America, Cuba, tropical Asia and New Guinea. Phylogeny and evolution: Fossil beechwood (Fagoxylon) is known from Upper Cretaceous deposits across the Northern Hemisphere. The family was more diverse in the Tertiary. The crown group has been estimated to be c. 34–37 million years old. They are sometimes divided into two subfamilies: Fagoideae including Fagus only, and Quercoideae encompassing the rest. Genera and species: Fagaceae have eight genera with 927 species: Castanea (8), Castanopsis (136), Chrysolepis (2), Fagus (10), Lithocarpus (336), Notholithocarpus (1), Quercus (431) and Trigonobalanus (3). Uses: Sweet chestnuts, Castanea sativa, have been cultivated since Roman times and

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were staple food for Roman soldiers, which caused the species to be introduced from the Mediterranean to all of Europe. It is still cultivated for its edible seeds, although mostly they are grown in Asia and then are often the Chinese chestnut (C. mollissima), which has been cultivated there, also for thousands of years. The American chestnut (C. dentata), Japanese chestnut (C. crenata) and chinquapin (C. pumila) and species of Castanopsis, Chrysolepis, Fagus and Lithocarpus densiflorus also have edible seeds and are eaten locally. Ground acorns (seeds of Quercus spp.) have been eaten as a famine food in Europe and even as a substitute for instant coffee. In Spain, ham from pigs fed on acorns is considered a delicacy and commands a higher price than other ham. The thick bark of the cork oak (Quercus suber) is harvested every eight to ten years, and this is the cork of commerce, used in flooring, insulation, bottle corks, floats etc. Beechwood is highly valued for cabinetry and furniture. Oak is frequently harvested for timber, and English oak (Quercus robur) was especially important in the past for the construction of houses, ships, furniture and wine barrels, giving a particular flavour to wine, port and sherry. Semi-fossilised oak wood (due to burial in peat) is known as bog oak, and due to its extreme hardness, was the wood of choice for heavy clubs (known as a shillelagh) in Ireland. Bark of Castanopsis and Quercus is used to make dyes, and oak galls were collected to make ink. Acorns were used for tanning, but tannins are now more frequently produced in the laboratory. Many species are highly valued ornamentals and frequently planted as shade trees especially in the temperate zones. Being worshipped in Europe since ancient times, oaks have become the national symbols of Germany, Ireland and the United States. Mistletoe (Viscum album, Santalaceae) was considered more sacred by druids when it grew on oak trees than on other trees. In British folklore, the amount of rain was thought to be predicted by the sequence of leafing out in oak and ash (Fraxinus excelsior, Oleaceae). If the oak leafed out first, relatively little rain would fall, but if the ash leaves came first, it was thought to presage a wet summer.

Etymology: Fagus is the classical Latin name for a beech tree, from its Indo-European root bhehgos. It probably is a cognate with classical Greek φαγειν ( fagein), to eat, in reference to its edible nuts. The surname of one of the authors of this book (Fay) is believed to be derived from the Norman French common name for Fagus sylvatica.

190. MYRICACEAE Bayberry family

Evergreen shrubs and small trees make up this family. Plants are unisexual or bisexual and are often covered with resinous, peltate glands, making the plants aromatic. Roots frequently have nitrogen-fixing nodules. They have simple alternate leaves usually lacking stipules and nearly sessile blades. Blades are entire to irregularly serrate or lobed, pinnatifid and subtended by stipules in Comptonia. Inf lorescences are unisexual spikes, and f lowers usually lack a perianth (Canacomyrica has a sixlobed perianth) and are wind-pollinated. Male f lowers are solitary in the axils of an inf lorescence bract, and each is usually subtended by two to four bracteoles. The two to eight (rarely up to 20) stamens occur in groups at the base of the bract, progressively fewer towards the tip of the inf lorescence. Filaments are short, free or slightly fused at base, and the erect anthers open by lengthwise slits. Female f lowers are up to four per inf lorescence bract, and each f lower is subtended by two to four bracteoles. The more or less inferior ovary (superior in Comptonia) is composed of two fused carpels forming a single locule. The two styles are free or fused at the base. Fruits are drupe-like nutlets, often with wart-like, wax-covered papillae.

FAGALES

EUDICOTS

Distribution: This family is widespread across both hemispheres, occurring mostly in temperate, subtropical or tropical-montane regions. Phylogeny and evolution: Myricaceae are sister to Juglandaceae and form a well-suppor ted subclade in Fagales, an association well supported by their morphology and fossil history. Myrica is sometimes subdivided, and genera such as Gale, Faya and Morella are then accepted. They nevertheless are similar morphologically. Comptonia is closely related to Myrica and sometimes included there. The genus was widespread across the Northern Hemisphere during the Tertiary, but is now restricted to eastern North America. New Caledonian Canacomyrica monticola is sister to the rest of the family and is known from fossils in New Zealand. Myricaceae are associated with Frankia, nitrogen-fixing filamentous bacteria, and mycorrhiza, common in other Fagales, appear to be absent.

Myrica gale, Helsinki Botanical Garden, Finland [190]

Genera and species: This family includes three genera and 57 species: Canacomyrica (1), Comptonia (1) and Myrica (55). Uses: Chinese bayberry or yumberry (Myrica rubra) has sweet, edible fruit, and several large-fruited cultivars are grown commercially in China. The fruit is eaten fresh, dried, jammed, canned and juiced and fermented into alcoholic beverages. A dye is prepared from the bark. The astringent fruit of M. faya is also sometimes eaten. Bayberry wax, harvested from the fruits of several Myrica species, was used traditionally to make candles and soap, especially by the early settlers in America. The scented foliage of bog-myrtle or sweet gale (Myrica gale) is sometimes used as insect repellent and to flavour beer and other alcoholic drinks in Scandinavia. Etymology: Myrica is derived from classical Greek μυρίκη (myrike), a tamarisk bush, in turn derived from from μύρων (myron), a scent, referring to the aromatic leaves.

191. JUGLANDACEAE Walnut family

These deciduous to evergreen usually bisexual trees and shrubs have a tight bark and large terminal buds. The alternate (rarely opposite) leaves lack stipules (or are caducous in Rhoiptelea) and are usually oddly (sometimes evenly) pinnate, rarely trifoliate or simple, the leaflets bearing glandular, peltate scales. The often aromatic leaflets are alternate, and leaflet blade margins are serrate, rarely entire, with pinnate venation. The pendent (sometimes erect) terminal or lateral inflorescences are usually panicles composed of unisexual spikes or male and female spikes solitary. Flowers are

Myrica nana, Kunming Botanical Garden, China [190]

Platycarya strobilacea, Royal Botanic Gardens, Kew, UK [191]

Juglans regia, Romania [191]

Comptonia peregrina, Brooklyn Botanical Garden, New York, USA [190]

Rhoiptelea chiliantha, fruits, Yunnan, China (ZZ) [191]

Juglans cathayensis, Royal Botanic Gardens, Kew, UK [191]

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FAGALES unisexual (sometimes bisexual in Rhoiptelea) and actinomorphic. Male flowers are subtended by a trilobed bract and often two bracteoles. They have up to four fused tepals or none, and three to 40 (rarely to 100) stamens per flower; filaments are short and fused. Anthers are nearly sessile and open by lengthwise slits. Female flowers are subtended by entire or trilobed bracts and usually have two or three bracteoles. Tepals can be up to four or none, when present fused basally to each other and the ovary. The inferior (rarely superior) ovary is composed of two fused carpels and a single style on top with two stigmas that are usually fused, sometimes four-lobed or plumed. The infructescence is an elongate pendulous spike or a short erect cone-like structure, with drupelike nuts that have a dehiscent or indehiscent husk, a two- or three-winged samara or a discwinged nutlet. Distribution: This family is widely distributed across the Northern Hemisphere, occurring in the Americas from Canada through Central America to the Andes (South America), the Caribbean and Atlantic Brazil, and in the Old World from the Balkans to the Caucasus, Central Asia and temperate and tropical East Asia, throughout Malesia and New Guinea. Phylogeny and evolution: Rhoiptelea is sister to the rest of Juglandaceae and because of its superior ovary and stipules has sometimes been placed in its own family Rhoipteleaceae, but the two share many other characters, so APG III combined them. These two diverged an estimated 85.8 million years ago. The oldest fossils assignable to Juglandaceae are c. 78–90 million years old. The family has a rich fossil record and was formerly more widespread and diverse in the Northern Hemisphere. Genera and species: Juglandaceae have nine genera and 63 species: Alfaroa (7), Carya (18), Cyclocarya (1), Engelhardtia (5), Juglans (20), Oreomunnea (2), Platycarya (3), Pterocarya (6) and Rhoiptelea (1). Uses: Walnuts are harvested from Juglans regia, a tree that has been grown for its

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nuts since ancient times in Central Asia, the Near East and southern Europe. It is now the walnut of commerce and used frequently in baked goods and confectionery. Oil from the seeds is tasty and also valued for its ability to lower blood pressure. The husks produce a dye that can be used to stain wood and dye wool and was used by women to dye their hair red-yellow in Ancient Greece. The bark and fruits of Platycarya strobilacea yield a black textile dye used in East Asia. Pecan trees, Carya illinoinensis, produce a delicious dessert nut, used in the same way as hazelnuts or walnuts in confectionery, baked goods etc. Pecan oil is mostly unsaturated, making it good for cooking and cosmetics. It is the state tree of Texas. Several other species have edible nuts that are enjoyed locally, including shellbark hickory (C. laciniosa), shagbark hickory (C. ovata), mockernut hickory (C. tomentosa), Japanese walnut (Juglans ailanthifolia), butternut (J. cinerea) and black walnut (J. nigra). Hickory (Carya) wood is sometimes used for smoking food. The wood of many species is valuable and used for furniture, stair banisters, cabinetry and formerly aeroplane propellors. Several species are planted as ornamentals.

the United States, Andrew Jackson, had the nickname “Old Hickory”, in reference to his toughness on the battlefield. Etymology: Juglans is the classical Latin name for the walnut, and is composed of Jovis, the god Jupiter, and glans, an acorn (also used in English for the distal end of a penis).

192. CASUARINACEAE She-oak family

Trivia: Hickory wood has the reputation of being very durable. The seventh President of

This is a family of evergreen bisexual and unisexual trees and shrubs with green, articulated, horsetail-like branchlets. Internodes are ridged (phyllichnia), in between which stomates occur. Leaves are reduced to a whorl of four to 20 tooth-like structures, each at the apex of an internode ridge. Inflorescences are alternate with whorls of tooth-like bracts, within which there are two lateral scale-like bractlets and a single flower. Male inflorescences are whorled spikes, and

Allocasuarina torulosa, male flowers, Australian National Botanic Garden, Canberra [192]

Casuarina equisetifolia, Santiago de Chile [192]

FAGALES

EUDICOTS

male flowers have one or two hooded scalelike tepals and a single stamen with a basifixed anther. Female inflorescences are globular or ovoid heads. Female flowers lack a perianth, and the ovary is composed of two fused carpels. The style is branched into two and has two elongated stigmas. Infructescences are more or less woody cones, the two bracteoles hardened and enlarged as valves, with the fruit itself a winged nut (samara). Distribution: The family occurs naturally in Madagascar, Malesia, Australia, Melanesia, New Caledonia and other Pacific islands. It is frequently cultivated and naturalises easily throughout the tropics and subtropics, in some cases becoming invasive. Phylogeny and evolution: Ceuthostoma and Gymnostoma had a much wider distribution in the Southern Hemisphere in the Eocene and Miocene, and they are known from several macrofossils. Some diversification in Australia occurred with aridification of the central zones, c. 13 million years ago, isolating western and eastern Australian clades.

Uses: Casuarina equisetifolia is much planted on tropical beaches as a wind break and to stabilise sand. The wood of Casuarinaceae is similar to that of oak (Quercus, Fagaceae), and they are therefore called ‘she-oaks’. Etymology: Casuarina is derived from kasuari, the Malay word for the cassowary bird, alluding to the similarity between the feathers of the bird and the pendent branchlets of the plant.

193. TICODENDRACEAE Tico-tree family

Genera and species: This family has four genera and 91 species: Allocasuarina (61), Casuarina (14), Ceuthostoma (2) and Gymnostoma (14).

This is a family of unisexual trees with alternate, evergreen petiolate leaves and stipules that are subulate and encircle the stem. Leaf blades are simple and have doubly serrate margins and pinnate venation. Male inflorescences are spike-like thyrses with f lowers in cymules that are arranged in whorls along the inflorescence axis, each

Allocasuarina nana, female in fruit, Australian National Botanic Garden, Canberra [192]

Gymnostoma chamaecyparis, New Caledonia [192]

subtended by a bract. Male flowers lack a perianth and have eight to ten free stamens. Anthers are almost sessile and basifixed and open by lengthwise slits. Female flowers are borne solitarily in leaf axils, and each is subtended by a pair of bracts. A perianth is almost absent, reduced to a little rim. The inferior, tetralocular ovary is composed of two fused carpels, each topped with an elongate stigmatic style. The fruit is a drupe. Distribution: Ticodendraceae are restricted to Mesoamerica. Phylogeny and evolution: Morphologically this family is closest to Nothofagaceae, but similarities with other fagalean families also exist. Molecular evidence places Ticodendron as sister to Betulaceae, in which it could be included, although differing in several characters. Eocene fossils assignable to this family are known from Oregon and the London Clay, and the family may be c. 50 million years old. Genera and species: This family consists of a single species, Ticodendron incognitum. Etymology: Ticodendron is composed of tico, a nickname for a person from Costa Rica, and Greek δένδρων (dendron), a tree. Ticodendron incognitum, fruit, Volcán Barva, Heredia, Costa Rica (CD) [193]

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FAGALES

194. BETULACEAE Birch family

This is a family of deciduous, bisexual trees and shrubs. Betulaceae have loose bark that remains unchanged or exfoliates in thin layers and is often marked with lenticels. Their alternate, simple, petiolate leaves are pinnately veined and usually have toothed margins. Stipules are free and fall off early. Unisexual inflorescences are formed terminally or in the axils and are bracteate. Male flowers are clustered into more or less elongate, pendent catkins, consisting of oneor three-flowered cymules or are reduced to clusters of flowers. Female flowers are formed in pendulous or erect, short catkins or in woody cones formed from the fused bracts. The small flowers are unisexual, windpollinated and lack a perianth, but sometimes one to six free or fused scale-like tepals are present. The usually four, sometimes one to 12 stamens have free or basally fused filaments. The dorsifixed anthers open by Carpinus caroliniana, fruiting and with autumn colour, Michigan. USA [194]

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lengthwise slits. The superior ovary consists of two (or three) fused carpels, each bearing a linear, filiform, free style. Infructescences are often woody cone-like or leaf-like structures with persistent or deciduous scales. Fruits are laterally compressed nuts or two-winged samaras, each with a single seed.

Coryloideae – Carpinus (39), Corylus (16), Ostrya (9) and Ostryopsis (3).

Phylogeny and evolution: Even though placement in Fagales has been supported by most characters, a morphological similarity to Hamamelidaceae (with Corylopsis as putative relative) has also been suggested, which is not backed by molecular evidence. Betulaceae are classified into two subfamilies based on the numbers of flowers in each cymule of the male inflorescence. The oldest fossils known from this family are c. 83–94 million years old, and the stem lineage is estimated to have an age of at least 130 million years. Coryloid fossils are known from the Palaeocene, and Alnus and Betula were well differentiated by the middle Eocene.

Uses: Hazelnuts are harvested from the wild (mainly Corylus avellana and C. colurna) and plantations (mainly cultivars of common hazel, C. avellana, and filbert, C. maxima), and are an important crop in some parts of the world. The fine-grained wood of Alnus and Betula is valued for furniture making, plywood, electric guitars, skateboards and loudspeakers and is sometimes used in the paper industry. Wood of hornbeam, Carpinus betulus, is hard and even though difficult to work was used when resistance to heavy wear was required, such as for tool handles, water and cart wheels etc. Birch, Betula pendula, is tapped for its sweet spring sap in Fennoscandia, Siberia and northern China as a refreshing drink, syrup or hair lotion. In England birch sap wine is produced in small quantities. Branches of this species, which give off a pleasant fragrance, are collected to make sauna whisks (vihta) and decorate boats at midsummer in the Nordic countries. A number of species are cultivated as ornamentals and for hedging and street trees. Betula species are often dominant in boreal forests.

Genera and species: Betulaceae include six genera with 167 species in two subfamilies: Betuloideae – Alnus (38) and Betula (62);

Etymology: Betula is the classical Latin name for a birch tree, borrowed from the Gaulish name for it (e.g. Welsh bedwen).

Betula utilis var. prattii, Royal Botanic Gardens, Kew, UK [194]

Corylus maxima, plantation near Trabzon, ºTurkey [194]

Distribution: This family is distributed across temperate and boreal forests of the Northern Hemisphere, but they extend into tropical mountain regions of Central and South America and Southeast Asia.

CUCURBITALES

EUDICOTS

CUCURBITALES Families 195 to 202 form the Cucurbitales, an order that evolved some 90–78 million years ago. The order is an unexpected assemblage of families, with few morphological characters in common. Coriariaceae with their free carpels were thought to be ‘primitive’, and placed variously among Ranunculales. Anisophylleaceae were once associated with Rhizophoraceae (Malpighiales) or Cunoniaceae (Oxalidales), but molecular evidence firmly places them in Cucurbitales. Most relationships in the order remain poorly understood, although Corynocarpaceae and Coriariaceae are clearly found to be sister taxa in molecular analyses and share wood-anatomical characteristics. Placement of parasitic Apodanthaceae, often included in Rafflesiaceae (Malpighiales), has long been unclear, but they are now supported as part of this order.

195. APODANTHACEAE Stemsucker family

This is a family of endoparasitic plants without chlorophyll. Only the flowering shoots are visible outside the host. They parasitise the roots and stems of woody plants (Fabaceae, Salicaceae, Burseraceae and Meliaceae) and have diffuse and delicate vegetative parts; stems and leaves are absent. Plants are unisexual, and actinomorphic flowers emerge solitarily or in groups or rows directly from the host tissue. Flowers are subtended by two whorls of scale-like, free or fused bracts. There are four or five free or basally fused tepals in both male and female flowers, with cushions of hairs inside. Nectaries are formed in a ring at the base of the staminal column. Male flowers have numerous fused stamens, forming a central column, with the anthers at the top; these open by transverse slits. Female flowers also have a stamen column, but they lack anthers. The inferior or semiinferior ovary lacks a style and is composed of four carpels fused into a single locule. Fruits are irregularly dehiscing berries containing small seeds.

Distribution: This widespread family has a patchy distribution, but due to their enigmatic nature and small stature they are probably frequently overlooked. They are known in the Americas from California to Argentina, tropical East Africa, southwestern Asia and southwestern Australia. Phylogeny and evolution: Diverse endoparasitic plants were previously all included in Rafflesiaceae, and due to their elusive nature, lack of vegetative parts and transfer of genes from the host, these taxa have been a major challenge in resolving phylogenetic relationships. However, molecular evidence now firmly places this family in Cucurbitales.

Pilostyles coccoidea, Western Australia (KD) [195]

Apodanthes only parasitises Casearia and Xylosma (Salicaceae). Pilostyles parasitises mostly Fabaceae and rarely Burseraceae or Meliaceae. Berlinianche from East Africa is only found on members of tribe Amherstieae (Fabaceae), but it does not differ much from Pilostyles and is now included in it. Genera and species: There are two genera with c. 23 species: Apodanthes (9) and Pilostyles (c. 14). Etymology: Apodanthes is composed of the Greek prefix α- (a-), without, and the words ποδός ( podos), foot, and άνθος (anthos), flower, e.g. a flower without a plant. Pilostyles aethiopica, Ogooue River, Gabon (CD) [195]

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Anisophyllea disticha, flower detail (WA) [196]

Combretocarpus rotundatus, Bako National Park, Sarawak, Malaysia (CD) [196]

Combretocarpus rotundatus in fruit, Bako National Park, Sarawak, Malaysia (CD) [196]

Anisophyllea disticha, in fruit (WA) [196]

196. ANISOPHYLLEACEAE

Female flowers often have staminodes and a (semi-)inferior ovary composed of usually four (sometimes three or five) fused carpels topped with free stylodia. The fruit is a fewseeded drupe or berry, dry and winged in Combretocarpus.

Uses: Inoi nuts are the oily seeds of Poga oleosa, which are locally eaten in tropical Africa as a wild food. The fruit of monkeyapple (Anisophyllea laurina) is eaten in West Africa. Many species become large trees, and their wood is sometimes exploited for timber, with Anisophyllea griffithii reported to reach a commercially exploitable size.

Leechwood family

Distribution: This family is found in South America, Southeast Asia and tropical Africa, where it occurs in primary, wet tropical and swamp forests. These bisexual trees and shrubs have alternate leaves that are sometimes dimorphic, one set scale-like (on the upper surface of the shoot) and the other normally leafly. Blades are sessile or shortly petiolate and lack stipules. Venation is pinnate, but often there are three to five prominent veins emerging from the base (a single midvein in Combretocarpus). Inflorescences are open, bracteate, axillary racemes or panicles with f lowers that are unisexual by abortion (bisexual in Combretocarpus). Usually four (rarely three or five) sepals are fused, and the four (sometimes three or five) free petals are usually lobed or dissected (entire in Polygonanthus). Male flowers are often smaller than female flowers and have twice the number of stamens as petals (usually eight). Anthers are dorsifixed, and stamens have a nectar disc at the base. A pistillodium is often present under the nectary. 284

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Phylogeny and evolution: Previously Anisophylleaceae were included in Rhizophoraceae, although it has been recognised on morphological grounds that the two families are widely separated (R hizophoraceae now placed with Erythroxylaceae in Malpighiales). A symposium on the matter was held in 1988. The putatively clear morphological similarity with Cunoniaceae (Oxalidales), once suggested, is due to convergence. Anisophylleaceae are now placed as sister to the remaining Cucurbitales based on molecular studies. Fossil pollen of Combretocarpus is known from late Miocene deposits in Borneo. Genera and species: This is a family of four genera and 34 species: Anisophyllea (c. 30), Combretocarpus (1), Poga (1) and Polygonanthus (2).

Etymology: Anisophyllea is composed of Greek άνισος (anisos), unequal, and φύλλων ( fyllon), a leaf.

197. CORYNOCARPACEAE Cribwood family

These evergreen, bisexual (rarely unisexual in Corynocarpus laevigatus) trees and large shrubs often have their branches in false whorls. Their simple leaves are alternate and often crowded at the end of the shoots. They are glossy and petiolate and appear to be without stipules, but a stipule subtending

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Corynocarpus laevigatus, in fruit, Dunedin Botanical Garden, New Zealand (JC) [197]

Corynocarpus laevigatus, flowers, Dunedin Botanical Garden, New Zealand (JC) [197]

Corynocarpus laevigatus, Dunedin Botanical Garden, New Zealand (JC) [197]

axillary buds is present, falling as leaves develop. At the nodes there are trichomoids, structures that are part hair, part scale. Inflorescences are terminal or (rarely) axillary, umbellate thyrses. The bisexual (rarely unisexual) flowers are actinomorphic and have a shallow cup-like receptacle. The five sepals and petals are free, and of the ten stamens, one whorl is staminodial and petal-like, irregularly dissected with a large globular nectary at the base. Five fertile stamens are attached to the petals at their bases and have dorsifixed anthers that open lengthwise. The superior ovary is composed of one or two fused carpels topped with one or two styles with a capitate stigma. Fruits are drupes, distributed by bats and pigeons.

Genera and species: The single genus in this family, Corynocarpus, consists of five species.

nitrogen-fixing bacteria (Frankia). Their simple leaves are opposite or (three-)whorled and subtended by minute stipules at the base of the petioles. Blades are entire and have palmate venation with three to five primary veins. Inf lorescences are racemes that terminate lateral or terminal branches or bear solitary flowers. The bisexual or functionally unisexual flowers are actinomorphic. There are five persistent sepals and smaller petals that are keeled and become fleshy and larger in fruit. The ten stamens are in two free whorls or the outer whorl with filaments fused to the petals. Anthers are basifixed or somewhat dorsifixed and open lengthwise. The ovary is superior and composed of five to ten free carpels, and each carpel has a long slender style that is papillose and stigmatic along its entire surface. Fruits are free, small nutlets enclosed in the fleshy persistent petals (pseudodrupe).

Uses: Fruits of Corynocarpus are poisonous when fresh, but after soaking and steaming the toxins can be removed and the fleshy fruit can be eaten. This is only done by indigenous people of the Solomon Islands, Vanuatu and New Zealand. Wood of Corynocarpus has been used for canoes and carving. Etymology: Corynocarpus is composed of the Greek words κορυνε (koryne), a club, and καρπός (karpos), a fruit.

198. CORIARIACEAE Tanner-bush family

Distribution: Corynocarpaceae are found in Australasia: New Guinea, the Solomon Islands, Vanuatu, New Caledonia, eastern Australia and New Zealand. Phylogeny and evolution: Fossil pollen and fruits of Corynocarpus are known from the Miocene of New Zealand. Molecular studies place Corynocarpaceae as sister to Coriariaceae in Cucurbitales, the molecular age for this clade being c. 50 million years. The two families share several morphological and anatomical characters, such as the trichomoids and minute stipules.

This is a family of upright and scrambling shrubs and woody herbs with quadrangularly ribbed branchlets and structures that are part hair, part scale (trichomoids) at the nodes. The roots have a symbiosis with

Distribution: Coriariaceae have a patchy distribution in the mountains of Central and South America (Mexico to Chile), the western Mediterranean region, the Himalayas, northern Burma, southern China, Taiwan, Japan, Luzon, New Guinea, New Britain, South Pacific islands and New Zealand. Phylogeny and evolution: Fossil flowers, leaves and seeds of Coriaria are known Plants of the World

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Coriaria sarmentosa, Royal Botanic Gardens, Kew, UK [198]

Coriaria sarmentosa, Royal Botanic Gardens, Kew, UK [198]

Coriaria ruscifolia in fruit, Ecuador [198]

from Oligocene and Pliocene sediments in Europe, the oldest being 33 million years old. As with many members of Cucurbitales, affinities of Coriaria have been problematic, but DNA sequence analyses placed it sister to Corynocarpus (Corynocarpaceae), with which it shares some characters.

199. CUCURBITACEAE

in leaflets of compound leaves. Inflorescences are axillary racemes, thyrses, panicles, spikes, umbels or fascicles with unisexual (rarely bisexual), actinomorphic or occasionally zygomorphic flowers. There are five basally fused sepals and petals (sometimes three to ten) and the petals can be partly fused and entire, bilobed or fringed, sometimes with a basal scale or fused for their entire length, forming a funnelshaped corona. A nectary is often present, and flowers of some species have oil glands. Male flowers have three to five stamens that are often fused along their filaments into a central column. Anthers are usually basifixed, straight or variously bent or contorted, rarely fused into a ring, the thecae opening lengthwise. Female flowers have an inferior or semi-inferior ovary composed of three (rarely as few as one or up to five) fused carpels, the styles free or fused and stigma entire, lobed or split. Fruits are many- to one-seeded, soft-skinned berries or (explosive) capsules, rarely a samara or achene. Seeds are flattened, winged or not.

Cucumber family

Genera and species: The sole genus in this family is Coriaria with 14 species. Uses: Leaves, stems and fruit of Coriaria contains gallic and ellagic acids that form tannins, and the tanner bush or redoul, Coriaria myrtifolia, has long been used for tanning of leather, especially during Mediaeval times. The black fruit of some species is used for dying, and the fleshy petals of Himalayan C. terminalis are sometimes eaten or made into a drink. All other parts including the seeds are poisonous and have to be consumed with caution. Etymology: Coriaria is derived from Latin corium, leather, in reference to some species being used for tanning.

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Cucurbitaceae are a family of unisexual and bisexual, annual and perennial herbaceous climbers and woody perennial vines, rarely trees (Dendrosicyos), often with tuberous roots or rhizomes or the plants with leafless, succulent, photosynthetic stems. Leaves are spirally arranged, petiolate and without stipules, often with an opposite and simple or once- or twicebranched tendril usually at a right angle (c. 90 degrees) to the petiole. Leaf blades are simple or palmately to pedately compound, and margins are entire to pedately or palmately lobed or dissected, often with small glandular teeth. Venation is palmate in entire leaves, pinnate

Distribution: This essentially tropical and subtropical family has only a few species in

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temperate regions, their vegetative parts being sensitive to frost. Phylogeny and evolution: Based on molecular phylogenetic studies, it has been suggested that the family originated somewhere in Asia during the Late Cretaceous (c. 80 million years ago). Fossils of Cucurbitaceae are sparse, but the oldest known are seeds from the London Clay (Upper Palaeocene or Lower Eocene, c. 65 million years old). The American clades evolved from only a few trans-Atlantic dispersal events. The only tree in the family, Dendrosicyos socotranus, evolved some 34 million years ago, before its current native island, Socotra, existed (it is only 10 million years old). Therefore, it is thought that this represents a relic of a lineage that became extinct on the continent. Some genera are in need of recircumscription, and revisions of the larger genera are greatly needed. There are numerous small to monospecific genera based on trivial morphological differences from their relatives, and many of these are embedded in other genera in phylogenetic studies. For instance, Sicyos now includes Costarica, Microsechium, Parasicyos, Pterosicyos, Sechiopsis, Sechium, Sicyocaulis and Sicyosperma; Melothria includes Cucumeropsis, Melancium and Posadaea; Cucumis has been expanded with Cucumella, Dicoelospermum, Mukia, Myrmecosicyos and Oreosyce; and Echinopepon is expanded to include Apatzingania, Brandegea and Vaseyanthus. Psiguria may have to be recircumscribed to include Gurania and Helmontia, and Ibervillea may also have to be expanded. An analysis with broader sampling

Marah oreganus, near San Francisco, California, USA [199]

is needed to verify these nomenclatural reassignments. The traditional division into subfamilies does not hold up in molecular studies. Four clades have been found that are recognised at the tribal level. Genera and species: Cucurbitaceae include 97 genera and c. 990 species: Abobra (1), Acanthosicyos (2), Actinostemma (3), Alsomitra (1), Ampelosycios (3), Anisosperma (1), Apodanthera (15), Austrobryonia (4), Baijiania (5), Bambekea (1), Bayabusua (1), Benincasa (2), Borneosicyos (1), Bryonia (c. 10), Calycophysum (5), Cayaponia (c. 55), Cephalopentandra (1), Ceratosanthes (4), Cionosicyos (5), Citrullus (4), Coccinia (c. 30), Cogniauxia (2), Corallocarpus (13), Ctenolepis (2), Cucumis (c. 55), Cucurbita (c. 15 wild ones + 5 domesticated), Cucurbitella (1), Cyclanthera (c. 40), Cyclantheropsis (3), Dactyliandra (2), Dendrosicyos (1), Dieterlea (3), Diplocyclos (4), Doyerea (1), Ecballium (1), Echinocystis (1), Echinopepon (c. 20), Eureiandra (8), Fevillea (8), Frantzia (5), Gerrardanthus (5), Gomphogyne (2), Gurania (c. 40), Gynostemma (c. 10), Halosicyos (1), Hanburia (7), Helmontia (2), Hemsleya (c. 30), Herpetospermum (3), Hodgsonia (2), Ibervillea (8), Indofevillea (1), Indomelothria (2), Kedrostis (23), Khmeriosicyos (1), Lagenaria (6), Lemurosicyos (1), Linnaeosicyos (1), Luffa (7), Marah (7), Melothria (c. 12), Melothrianthus (1), Momordica (c. 60), Muellerargia (2), Neoalsomitra (c. 12), Nothoalsomitra (1), Papuasicyos (8), Penelopeia (2), Peponium (c. 20), Peponopsis (1), Polyclathra (6),

Alsomitra macrocarpa, seeds, Mt Datuk, Malaysia [199]

Psiguria (c. 12), Pteropepon (5), Raphidiocystis (5), Ruthalicia (2), Schizocarpum (11), Schizopepon (8), Scopellaria (2), Seyrigia (6), Sicana (4), Sicydium (7), Sicyos (c. 75), Siolmatra (2), Siraitia (4), Solena (3), Tecunumania (1), Telfairia (3), Thladiantha (c. 30), Trichosanthes (c. 100), Tricyclandra (2), Trochomeria (8), Trochomeriopsis (1), Tumamoca (2), Wilbrandia (5), Xerosicyos (5), Zanonia (1) and Zehneria (c. 60). Uses: A great number of species are of economic importance, usually as vegetables, fruits and medicines (cucurbitacin B is antiinf lammatory); their seeds produce oils frequently used for cooking. Tough-skinned species are used to make utensils and the spongy remains of the fruits of some species are used as vegetable sponges. The main uses are listed below. Vegetables: the commercially most widely grown is the cucumber (Cucumis sativus), a species known to have been cultivated some 3,000–8,000 years ago in the Middle East, as it was listed as one of the foods of ancient Ur of the Chaldeans. There currently are many cultivars, some bred to be eaten fresh (slicing cucumbers) for salads etc., some seedless, sweet varieties (burpless cucumbers), and other cultivars are bred for pickling as gherkins. Cucumber extract is also used in cosmetics as a face cleanser. Armenian cucumbers (Cucumis melo var. flexuosus) are similar and marketed locally in the Middle East. Butternut squash (Cucurbita moschata) was domesticated c. 9,200 years ago in the Andes of northern Peru. It is now commonly

Citrullus colocynthis, near Oodnadatta, South Australia [199]

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Sicyos edulis, Jardin des Plantes, Paris, France [199]

EUDICOTS

Ecballium elaterium, Bergius Botanical Garden, Stockholm, Sweden [199]

cultivated, but especially in climates with warmer summers. Squash (Cucurbita pepo) seeds were found in caves in Tamaulipas, Mexico, and these have been carbon-dated to c. 7,000 years old. This cultigen, unknown in the wild, may have been independently derived in North and Central America from closely related species: C. texana in the eastern USA giving C. pepo subsp. ovifera, and C. fraterna from Mexico giving C. pepo subsp. pepo. There are numerous cultivars, some more pumpkin-like, other more cucumber-like. Courgette or zucchini (C. pepo subsp. pepo var. cylindrica) and crookneck squash (C. pepo subsp. ovifera var. torticollia) are eaten immature and cultivated on a large commercial scale. Other cultivars produce the non-keeping summer pumpkins (C. pepo subsp. pepo var. pepo), pattypan or scallop (C. pepo subsp. pepo var. clypeata), cocozzelle (C. pepo subsp. pepo var. longa), marrow and summer squash (C. pepo subsp. pepo var. fastigiata) and ‘vegetable spaghetti’ (Spaghetti Squash Group). Most winter squash (sometimes referred to as “pumpkins”, especially in the UK) are varieties of C. maxima, recognised by its corky stem. This was first domesticated four millenia ago in South America. These can look much like cultivars of C. pepo, but can be distinguished by their thick, sponge-like stem. The stem of C. pepo is thin and woody at maturity. Both come in many cultivars, some small and some incredibly large, the heaviest known from California weighed 691 kg! These are the pumpkins carved at Halloween and made into pumpkin pie for Thanksgiving in the United 288

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Cucurbita maxima, private allotment, Kingston upon Thames, Surrey, UK [199]

States (and increasingly elsewhere around the world). Canned pumpkin purée is usually made from winter squash or silverseed gourd (C. argyrosperma) or butternut squash (C. moschata). Native to Mexico, the chayote or christophine (Sicyos edule, formerly Sechium edule) has fruits that each have a single large seed. When growing, the entire fruit is planted, and it is now cultivated throughout the tropics. The fruit is a staple in many tropical cuisines, and the starchy tubers and the young shoots are eaten as a vegetable, especially in Asia. There are a number of other minor vegetable crops, the (young) fruits often being cooked or pickled; they are a major ingredient of Indian and Pakistani curries. Examples are: narras (Acanthosicyos horridus), gemsbok cucumber (A. naudinianus), wax gourd or winter melon (Benincasa hispida), tinda (B. fistulosus, formerly Praecitrullus fistulosus), ivy gourd (Coccinia grandis), papasan (C. cordifolia), tindori (Cucumis anguria), stuffing gourd or achocha (Cyclanthera pedata), sinqua (Luffa acutangula), bitter cucumber (Momordica charantia), balsam apple (M. balsamina) and snake gourd (Trichosanthes cucumeroides). Young fruits of bottle gourd (Lagenaria ciceraria) are also eaten, peeled and dried in Japanese cuisine. Tubers of Coccinia grandis, C. abyssinica and Trichosanthes kirilowii, among others, are a source of starch. Additionally the leaves and young shoots of many species are boiled and eaten as a vegetable, especially in Africa and Asia. Fruits: melon (Cucumis melo) is a species of unknown origin, but wild forms (subsp. agrestis) are known from Africa. Their seeds

were once exported for their oil. They first appeared in Europe during Roman times, and melons are among the first plants to have been domesticated. There are seven cultivar groups: e.g. true cantaloupes (cantalupensis group); muskmelons, rockmelons and false cantaloupes (reticulatus group); honeydew and Chinese melons (inodorus group), snake melons (flexuosus group), pickling melons and sweet melons (conomon group), orange melons (chito group) and dudaim melons (dudaim group), the first three being the most frequent in commerce. Of African origin, the watermelon (Citrullus lanatus) is now cultivated worldwide for its refreshing, fleshy fruits, that can be eaten fresh or made into juices or drinks, boiled into syrups or used in cosmetics. They were possibly selected c. 4,000 years ago out of bitter apple (C. colocynthis), a species that has been cultivated in the Middle East since ancient times and was used as a purgative. Watermelon plants are extremely drought tolerant due to their deep root system. Kiwano or horned melon (Cucumis metuliferus) is a traditional food plant in Africa, being one of the few sources of fresh water in the Kalahari during the dry season, the other being Gemsbok cucumber (Acanthosicyos naudinianus). Because of the odd shape and green flesh of kiwano, it has become a fashionable dessert fruit and is now cultivated elsewhere. Casabanana (Sicana odorifera) has fragrant sweet fruits that can be made into preserves and is used as a pleasant scent in the house or closet. Zehneria anomala also has a sweet fruit and is sometimes collected from the wild in the

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Middle East. Not naturally sweet, cidra or Malabar gourd (Cucurbita ficifolia) is often candied and served as a stringy dessert called ‘cabello de angel’ (angel hair). In the Balearic Islands, it is made into a stringy marmalade. Seeds of Cucumis melo and Cucurbita species are commonly roasted and eaten, especially in the Middle East. Pumpkin seed oil is produced from roasted pumpkin seeds and used for cooking and salad dressing, especially popular in Austria and Eastern Europe. Egusi seeds (Melothria mannii and wild forms of Citrullus lanatus) are an important source of oil and protein in dryland Africa. Seeds of Actinostemma lobatum are used for cooking oil. Apodanthera aspera has oil-rich seeds and has been used for food since pre-Columbian times in Mexico. Fevillea cordifolia and F. pedatifolia are widely cultivated for their oil-rich seeds, which are used to make candles and as a purgative. Fluted pumpkin (Telfairia occidentalis) and oyster nut (T. pedata) are often cultivated in tropical Africa for their oily seeds, useful for soap and candle making. The hard seeds of Hodgsonia heteroclita are eaten in Sikkim; after roasting, they taste like lard. Seeds of Cayaponia kathematophora and Echinocystis lobata are used as beads for necklaces, etc. Marah macrocarpus seeds produce a red dye. Flowers of many species are edible, and especially courgette and squash flowers are popular in some cuisines, in salads or stuffed with ricotta and cooked. In parts of Mexico, the cooked flowers are eaten in omelettes. Tetrameles nudiflora on the Ta Prohm Temple, Angkor Wat, Cambodia [200]

Utensils: loofah or sponge gourd (Luffa cylindrica) has a spongy vascular system when mature and after bleaching becomes a vegetable scrubbing sponge for use in the kitchen or bath. Bottle gourd or calabash (Lagenaria ciceraria) has long been used for vessels, and Central American fragments have been carbon dated to 10,000 years, although it probably originated in the Old World (known by Pliny) and was distributed across the Pacific from the Americas. Mature fruits are used for flasks, cups, musical instruments and penis sheaths in Papua New Guinea. Ornamental gourds (Cucurbita pepo subsp. ovifera) come in many cultivars and are popular as seasonal decorations and in flower arrangements.

200. TETRAMELACEAE False hemp-tree family

Etymology: Cucurbita is the Latin name for a gourd. It is possibly a cognate with cucumis, a cucumber.

Large deciduous and evergreen, unisexual trees with buttressed trunks and soft wood make up this family. Their simple leaves are alternate and long petiolate and lack stipules. The simple, heart-shaped leaf blades are palmately veined with three to five main veins and entire or toothed margins. Inflorescences are bracteate, long pendent spikes or spike-like thyrses, clustered at the apices of the stems, the males usually branched, the females usually simple and sometimes axillary. The four to eight sepals are fused into equal or unequal lobes or are mostly free, fused only below the ovary. Petals are absent (Tetrameles) or six to eight, erect, narrowly triangular and green in male flowers of Octomeles. Male flowers have four to eight stamens, the filaments elongate, somewhat flattened and the anthers basifixed, opening lengthwise. A sterile pistillode is often present. Female flowers lack staminodes, the inferior ovary with a single locule, four to eight short, stout styles and large, capitate or obliquely oblong-decurrent stigmas. Fruits are capsules that open at the top between the

Tetrameles nudiflora, male inflorescence (DV) [200]

Octomeles sumatrana, Lungmanis, Sabah, Malaysia (CD) [200]

Gliders and squirters: Some Cucurbitaceae have strange seed dispersal mechanisms. Seeds of Alsomitra macrocarpa have among the largest wings of any samara, to 12 cm across. These aerodymanic seeds glide from the canopy to the ground in its native Malesian rainforests. Fruits of Ecballium elaterium (squirting cucumber) detach explosively from the pedicel when ripe, and the seeds are forcefully released in a slimy liquid. Native to Europe, North Africa and parts of temperate Asia, this species is sometimes grown in gardens, where this dispersal mechanism causes much amusement.

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CUCURBITALES persistent styles (Tetrameles) or split in two layers and open lengthwise (Octomeles). Distribution: This family is distributed from Sri Lanka, eastern India and Nepal through tropical Southeast Asia and Malesia to the Solomon Islands and northeastern Australia. Phylogeny and evolution: In the past, Octomeles and Tetrameles were placed with Datisca in Datiscaceae. However, that group would be paraphyletic with regard to Begoniaceae, and thus two families are now recognised: Datiscaceae, comprising of Datisca only, and Tetramelaceae, including the other two genera. Tetramelaceae with Datiscaceae and Begoniaceae are sister to Cucurbitaceae. The two genera are easily distinguished by the numbers of flower parts, reflected by their names: Tetrameles is usually four-parted, whereas Octomeles is usually eight- (sometimes six-) parted. Fossil wood of Tetrameles from Early Eocene deposits of India has been described.

for coastal underwater constructions. For the same reason, it is sometimes used to make canoes in New Guinea and Melanesia. Etymology: Tetrameles is derived from Greek τετρα, tetra, four, and μέλος (melos), member or part, in reference to the four sepals.

201. DATISCACEAE Durango-root family

Female flowers have three to eight fused sepals with short lobes, without staminodes (but rarely two to four stamens in bisexual flowers). The inferior ovary is composed of three to four fused carpels forming a single locule, each carpel topped with a free style that is deeply forked, with papillate stigmatic branches. Fruits are capsules that open at the top between the persistent styles. Distribution: The family shows a marked disjunction, with one species in western North America, in California, Nevada and Baja California (northwestern Mexico) and the other in Crete, Turkey, the Caucasus and the Levant, and from Central Asia to Nepal. They grow in moist places, often by streams in forests.

Uses: Although their wood is not particularly hard, Octomeles is used for cabinetry, furniture and general construction, and Tetrameles is resistant to marine borers and is therefore used

These are large unisexual (rarely bisexual) perennial herbs that have a symbiosis with nitrogen-fixing Frankia bacteria. Their oddly pinnate to deeply pinnatifid leaves are alternate and lack stipules. The upper ones are slowly reduced from trifoliate and lobed to simple unlobed bracts, the leaflets pinnately veined and margins serrate. Inflorescences are bracteate, long pendent spike-like thyrses, clustered at the apices of the stems or in axillary fascicles. Male flowers have three to four (to five) fused sepals and lack petals. The six to 15 (to 25) stamens have subsessile basifixed anthers that open lengthwise.

Phylogeny and evolution: Datisca is widely disjunct between mediterranean climates in western North America (D. glomerata) and the Mediterranean-Central Asia (D. cannabina). Such disjunctions between America and the Mediterranean were achieved during the Palaeocene. The species are similar and lack morphological distinctions, although chemical and DNA studies have shown that populations from the eastern Mediterranean (D. cannabina) are genetically closer to Californian D. glomerata than to the Central Asian-Himalayan populations; as a result, the latter may have to be accepted as a third species, D. nepalensis. Datiscaceae are sister to Begoniaceae.

Datisca cannabina, male flowers, private garden, Kingston upon Thames, Surrey, UK [201]

Datisca cannabina, male and female plant, private garden, Kingston upon Thames, Surrey, UK [201]

Datisca cannabina, female flowers, private garden, Kingston upon Thames, Surrey, UK [201]

Genera and species: There are two genera in Tetramelaceae, each with a single species: Octameles sumatrana and Tetrameles nudiflora.

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Genera and species: This family has a single genus Datisca with two or perhaps three species. Uses: Datiscin is a white crystalline glucoside extracted from bastard hemp (D. cannabina), which is an ancient oriental dye. Datisca cannabina is occasionally grown as a garden ornamental. Durango root (D. glomerata) was used by native Americans as a fish poison. Etymology: The name Δατηςκα (Datiska) first appeared in Dioscorides’ De Materia Medica, where it is cited as a name for Catananche (Asteraceae). The name is probably derived from Ancient Greek δατεσσαι (datessai), to heal, and εξισώ (exiso), to equate, in reference to its medicinal properties. Linnaeus borrowed the name and applied it to this unrelated plant.

202. BEGONIACEAE Begonia family

Begoniaceae are bisexual, perennial (rarely annual), succulent herbs, sometimes somewhat woody and forming subshrubs but rarely large tree-like herbs to four metres. They grow terrestrially and epiphytically, also often on rocks or cliff faces. Stems are erect or creeping and rooting at the nodes, sometimes forming large stem tubers or stemless, rarely climbing with adventitious roots or stoloniferous. Their simple or palmately compound leaves are alternate (rarely opposite or whorled) or in a basal rosette. They are usually petiolate, and stipules are ovate to triangular and sometimes large and papery. Blades are usually oblique, asymmetrical basally, sometimes peltate, the margins entire or variously lobed, irregularly serrate or divided. Venation is palmate or pinnate, and surfaces can be colourfully variegated or covered in hairs or scales. A bulbil is sometimes present in the leaf axils.

is usually bi- to tetraloculate (rarely up to eight locules). Styles are commonly as many as locules, usually two or three, free or fused at the base and once or twice forked, with a spirally twisted, U-shaped, capitate or reniform stigma. The fruit is an unequally three-winged or three- to four-horned, dry capsule or berry.

Inflorescences are bracteate, axillary mono- or dichasial cymes, or flowers solitary or paired in the leaf axils, sometimes inflorescences terminal raceme-like cymes or panicles. The unisexual flowers are actinomorphic, zygomorphic or irregular, usually sexually dimorphic; the perianth is not differentiated in Begonia, but a calyx and corolla are present in Hillebrandia. Male flowers have two or four petal-like tepals, usually the outer larger than the inner. The three to over 100 stamens have distinct or variously clustered filaments, basifixed and opening lengthwise or by an apical slit or pore and sometimes with an extended connective. Female flowers have two to five, rarely up to 11 free (rarely fused) petal-like tepals. The inferior (Begonia) or semi-inferior (Hillebrandia) ovary is usually composed of two or three fused carpels and

Phylogeny and evolution: Begoniaceae have been variously associated with other

Begonia luxurians, Royal Horticultural Society Garden, Wisley, UK [202]

Begonia dregei, private garden, Kingston upon Thames, Surrey, UK [202]

Begonia cucullata, Réunion [202]

Begonia coccinea, Glasgow Botanic Gardens, Scotland, UK [202]

Distribution: Begoniaceae have a pantropical distribution from North America (Mexico) and the Caribbean to southern South America (northern Argentina and Uruguay), throughout tropical Africa, Socotra and tropical Asia, extending into the temperate regions of China and Taiwan and throughout the Pacific to Hawaii.

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CUCURBITALES cucurbitalean or malpighialean families (because of their often parietal placentation), and even though the composition of associated families changed over the years, Datiscaceae (including Tetramelaceae) have always been considered close, which has been corraborated by molecular studies. These families share embryological and seed characteristics and inferior ovaries with parietal placentation, placing them firmly in Cucurbitales. Begonia in the broad sense is monophyletic, although some authors have accepted a number of segregate genera in the past, most are now recognised at the sectional level in Begonia. No subdivisions are accepted here. Only the morphologically, genetically and geographically isolated Hillebrandia is accepted as separate, although this could also be included because it is sister to Begonia. The split between Begonia and Hillebrandia has been estimated to have occured between

EUDICOTS

58 and 45 million years ago, which of course questions the origins of Hillebrandia because the current set of Hawaiian Islands is not old enough for the species to have evolved there, although it may have hopped around other Pacific islands that are now submerged. Begonia diversified some 30–25 million years ago, during a period of global cooling. Genera and species: There are two genera in Begoniaceae, the widespread Begonia with 1,803 species (and many more undescribed ones) and Hillebrandia sandwicensis only found in Hawaii. Uses: Leaves and petioles of some species (especially Begonia muricata) are sometimes eaten. Begonias are important ornamental plants, often used for summer bedding or as houseplants in the temperate zones, all originating through complex hybridisation.

Semperflorens begonias, used for bedding (B. semperflorens-cultorum group), are a result of crosses between various forms of B. cucullata and B. schmidtiana. Tuberous begonias (B. ×tuberhybrida) are complex hybrids that involve B. boliviensis, B. pearcei and B. veitchii. Rex begonias with their ornamental leaves (B. rex-cultorum group) are crosses between B. rex and other Asian or Mexican species. Winter-f lowering begonias (B. hiemalis group and B. ×cheimantha) involve crosses of tuberous begonias with B. dregei and D. socotrana and selections of mutants thereof. Etymology: Begonia is named for French marine intendant and plant collector Michel Bégon (1638–1710), Governor of French Canada. He visited Charles Plumier in the Caribbean, who named this genus in his honour, which later was established by Linnaeus.

CELASTRALES Families 203 and 204 comprise the order Celastrales. This order is closely related to Oxalidales, and indeed some species of Oxalidaceae were once placed in Lepidobotryaceae. Huaceae (here in Oxalydales) have been found to belong to this clade in some analyses, although this is not supported by others. The crown group of Celastrales evolved c. 50–80 million years ago. Placement of the COM clade (Celastrales, Oxalidales and Malpighiales) here is based entirely on plastid DNA analyses (including whole plastid genome analyses, in which it is highly supported), but in results from mitochondrial and nuclear gene analyses, this clade is sister to the malvids instead of the fabids, which could be the result of an ancient reticulation event.

203. LEPIDOBOTRYACEAE Snail-cedar family

This is a family of unisexual trees. Their simple leaves are actually compound but reduced to a single leaf let, which is

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petiolulate and articulate to a petiole with a pulvinus. Stipules are present but soon fall off, the blades entire and pinnately veined. Inf lorescences are bracteate racemes or spikes, sometimes cone-like. The unisexual flowers are actinomorphic and have five free sepals and petals placed on a short saucershaped hypanthium. Male flowers have ten stamens in two whorls, the filaments fused into a tube with a nectar disc at the base. Anthers are basifixed (Ruptiliocarpon) or dorsifixed (Lepidobotrys) and open lengthwise. The superior ovary is composed of two or three fused carpels, each carpel topped with a free style. The fruit is a

Ruptiliocarpon caracolito, Rancho Quemado, Osa Peninsula, Puntarenas, Costa Rica (CD) [203]

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septicidal capsule with the outside separating from the inside and exposing black seeds with an orange aril. Distribution: The family occurs in tropical America (Costa Rica, Colombia, Suriname, Peru) and the Congo region of Central Africa. Phylogeny and evolution: The genera of Lepidobotryaceae were originally included in Linaceae but were later moved to Oxalidaceae, in which they were considered somewhat anomalous. They were subsequently assigned to a number of families (e.g. Erythroxylaceae, Meliaceae), from which they differ markedly. Stackhousia monogyna, Mt Benia, Western Australia [204]

Molecular studies found Lepidobotryaceae to be sister to Celastraceae. The two genera of Lepidobotryaceae are morphologically similar, and generic distinction between them may not be warranted.

204. CELASTRACEAE Spindle family

Genera and species: This family has two similar genera, each with a single species: Lepidobotrys staudtii in Africa and Ruptiliocarpon caracolito in the Neotropics. Etymology: Lepidobotrys is composed of the Greek words λέπις (lepis), a flake or scale, and βότρυς (botrys), trusses, bunches or racemes.

These are unisexual and bisexual trees, shrubs, vines, and annual and perennial herbs, rarely ericoid subshrubs and epiphytes. Stems

Parnassia palustris, Pyrenees, France [204]

Lepuropetalon spathulatum, Freestone County, Texas, USA (CD) [204]

Brexia madagascariensis, Helsinki Botanical Garden, Finland [204]

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CELASTRALES are unarmed or with thorns, sometimes the branches terminating in sharp points. Leaves are simple, alternate, opposite or whorled, sometimes in basal rosettes, or fascicles on short shoots, usually petiolate with small stipules. Blades are usually leaf-like, rarely scale- or needle-like, or reniform to orbicular and fleshy, the margins entire, toothed, crenate, serrate, notched, spinose or glandular-toothed, the venation pinnate, sometimes with domatia in axils of larger veins. Inflorescences are axillary or terminal thyrses, cymes or fascicles, rarely epiphyllous (Polycardia), cauliflorous, umbellate, paniculate, racemose or spicate, in Parnassioideae the flowers solitary on scapes or lateral shoots. The actinomorphic (rarely weakly zygomorphic) flowers are bisexual or unisexual. The four to five sepals and petals (rarely three to seven or absent) are free or with the median petals fused. A nectary disc is present in the flower cup, usually cup-shaped with the margins upturned, sometimes absent. Five glandular staminodes occur between the motile stamens in Parnassioideae. Stamens sit between petals and are (two or) three to five, rarely more numerous, all equal or sometimes three long and two short, staminodial or absent in female flowers. Anthers are basifixed to dorsifixed and open lengthwise, along the width or obliquely so, the connective sometimes apiculate. The superior or semiinferior ovary is often partially sunken in the nectary disk and has (one to) two to five (or up to ten) fused or partially fused carpels. In male flowers, the ovary is absent or a sterile pistillode. The style is simple and terminates the ovary, topped with a simple or lobed stigma. The fruit is a loculicidal or septicidal capsule or a schizocarp that is composed of two

to five indehiscent parts, sometimes a drupe or a berry or a samara with a surrounding wing, three to five lateral wings or one apical wing, rarely a nutlet. Seeds are one to several per fruit and sometimes are winged or arillate, minute in Parnassioideae.

Phylogeny and evolution: The estimated age of Celastraceae is 71–85 million years. Celastraceae are now considered to include a number of previously recognised families (e.g. Brexiaceae, Hippocrateaceae, Parnassiaceae, Stackhousiaceae), but these form a clade with the previous concept of Celastraceae and are thus treated as a single family. Phylogenetics of the family was originally problematic because it was polyphyletic, with Bhesa and Goupia being distinct (now placed in Malpighiales, in Centroplacaceae and Goupiaceae, respectively). Perrottetia was morphologically and genetically unlike other members (and is now placed in Dipentodontaceae of Huerteales) and Forsellesia is now a synonym of Glossopetalon in Crossosomataceae. Plagiopteron was part of the now defunct Flacourtiaceae, Brexia was once included in Hydrangeaceae and Parnassia was in Saxifragaceae or its own family Parnassiaceae. With the reorganisation of membership of Celastraceae, they have become morphologically more uniform, but the generic and subfamilial classifications need much work. Nicobariodendron is only

Genera and species: Celastraceae are a family of c. 96 genera and c. 1,200 species, possibly divided into two subfamilies: Parnassioideae – Lepuropetalon (1) and Parnassia (c. 70); Celastroideae – Acanthothamnus (1), Allocassine (1), Anthodon (2), Apatophyllum (5), Apodostigma (1), Arnicratea (3), Bequaertia (1), Brassiantha (1), Brexia (1), Brexiella (2), Campylostemon (8), Canotia (2), Cassine (3), Catha (1), Celastrus (2), Cheiloclinium (11), Crossopetalum (c. 26), Cuervea (5), Denhamia (15), Dicarpellum (4), Elachyptera (7), Elaeodendron (c. 40), Empleuridium (1), Euonymopsis (4), Euonymus (c. 130), Fraunhofera (1), Gloveria (1), Glyptopetalum (c. 20), Goniodiscus (1), Gyminda (4), Gymnosporia (c. 85), Hartogiopsis (1), Haydenoxylon (3), Hedraianthera (1), Helictonema (1), Hexaspora (1), Hippocratea (3), Hylenaea (3), Hypsophila (2), Kokoona (8), Lauridia (2), Loeseneriella (16), Lophopetalum (c. 20), Lydenburgia (2),

Maytenus octogona in fruit, Galápagos Islands

Euonymus verrucosus, Helsinki Botanical Garden, Finland [204]

Euonymus maximowiczianus, East Bergholt Place Arboretum, UK [204]

[204]

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Distribution: This nearly cosmopolitan family occurs from the Arctic to Antarctic America, throughout Europe, North Africa, Sub-Saharan Africa, Madagascar, Sri Lanka, northern, western, central and eastern Asia, Southeast Asia, Malesia, Melanesia and Australia.

known from the type specimen and is placed here tentatively. Lepuropetalon (here in subfamily Parnassioideae) has sometimes been placed in its own family (Lepuropetalaceae) or in Saxifragaceae (to which it is only distantly related). It is one of the smallest terrestrial flowering plants, being only a few millimetres tall when in flower, and some authors consider it the smallest. The minute monocot, Wolffia (Araceae), is certainly smaller, but Lepuropetalon is probably the smallest eudicot. Even if Gymnosporia is segregated, Maytenus remains a polyphyletic and polymorphic genus. Denhamia and Haydenoxylon are therefore accepted and Moya and Tricerma are synonymised.

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Macgregoria (1), Maurocenia (1), Maytenus (c. 200), Menepetalum (4), Microtropis (c. 66), Monimopetalum (1), Mortonia (5), Mystroxylon (1), Orthosphenia (1), Paxistima (2), Peripterygia (1), Peritassa (13), Plagiopteron (1), Platypterocarpus (1, possibly extinct), Plenckia (4), Pleurostylia (8), Polycardia (4), Pottingeria (1), Prionostemma (5), Pristimera (24), Psammomoya (4), Pseudosalacia (1), Ptelidium (4), Pterocelastrus (4), Quetzalia (11), Reissantia (6), Robsonodendron (2), Rzedowskia (1), Salacia (c. 200), Salacighia (2), Salaciopsis (6), Salvadoropsis (3), Sarawakodendron (1), Schaefferia (c. 23), Semialarium (2), Simicratea (1), Simirestis (8), Siphonodon (7), Stackhousia (c. 16), Tetrasiphon (1), Thyrsosalacia (4), Tontelea (31), Torralbasia (1), Tripterococcus (2), Tripterygium (1), Tristemonanthus (2),

Wimmeria (12), Xylonymus (1) and Zinowiewia (17). Unplaced – Nicobariodendron (1). Uses: The leaves of qat (Catha edulis) are chewed or brewed into a tea and consumed as a stimulant. It is considered an illicit drug in several countries, where it is now outlawed, but it is still in frequent use in tropical Africa and Asia. It is a legal cash crop in Ethiopia. Several species have edible seeds or fruits, e.g. Arnicratea grahamii, mfukufuku (Brexia madagascariensis), Hylenaea comosa, Laurida tetragona, Maurocenia frangula, Mystroxylon aethiopicum, Paxistima myrsinites and Salacia species. The wood of Euonymus is hard and traditionally was turned into sharp pointed spindles for spinning wheels. Most parts of Euonymus are poisonous and may lead to death — although not by

pricking your finger on a spindle, as told in the folk tale Sleeping Beauty popularised by the Brothers Grimm — but consuming the fruit is not advisable. Wood of several tropical species is used for timber, and the bark of Kokoona and Lophopetalum contains an oily layer that can be used to start a fire. Several species are grown as garden ornamentals, especially species of Cassine, Celastrus, Elaeodendron, Euonymus, Maytenus, Parnassia, Paxistima, and Tripterygium. Etymology: Κελαστρος (kelastros) is a Greek name for an evergreen tree, with fruits remaining on the tree throughout the year according to Theophrastus. It was erroneously used by Linnaeus for these deciduous vines which do not occur in the Mediterranean, although the fruit does remain on the vine in winter.

OXALIDALES Families 205 to 211 belong to Oxalidales, an unexpected association of families that has few obvious synapomorphies. The age of the order is estimated to be c. 90 million years. Huaceae are the first branching clade, followed by a clade of Connaraceae and Oxalidaceae being sister to the rest. Placement of the COM clade (Celastrales, Oxalidales and Malpighiales) here is based entirely on plastid DNA analyses, but in results from mitochondrial and nuclear gene analyses, the COM clade are sister to the malvids instead of the fabids, which could be the result of an ancient hybridisation event.

205. HUACEAE Cameroon-garlic family

Huaceae are a family of trees and shrubs with a strong scent of garlic. The alternate, simple leaves often have glands at the base of the blade or along the entire margin. Stipules are present but soon fall. Inflorescences are cymes or the flowers solitary in the axils. The bisexual,

actinomorphic flowers have five free or partly fused sepals and five free petals. Sepals often bear glands, and petals are usually hairy. The ten (or eight) stamens are of equal length, all fertile and free and arranged in a single whorl. Anthers are basifixed, sometimes with an apical appendage (Afrostyrax), and open by apical or lengthwise slits. The superior ovary is composed of five fused carpels, forming one (Hua) or five (Afrostyrax) locules. Fruits are capsules that split or not, holding one or two large seeds.

Afrostyrax lepidophyllus, Cameroon (CD) [205]

Distribution: This family is restricted to tropical West and Central Africa. Phylogeny and evolution: These two genera were previously placed in Malvaceae (Hua) Plants of the World

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and Styracaceae (Afrostyrax), each in an isolated position within these families. Since their relationship was recognised in 1913, the family has been suggested to have a great number of relatives, although few similarities were obvious. The first molecular studies suggested a relationship with Celastraceae, but with low support, and stronger support is now found for them as sister to the other families of Oxalidales, which is where they are now placed.

206. CONNARACEAE

Genera and species: This small family includes two genera and three species: Afrostyrax kamerunensis, A. lepidophyllus and Hua gabonii.

Etymology: Hua is named for French botanist Henri Hua (1861–1919), curator at the herbarium of the Muséum national d’Histoire naturelle in Paris. He participated in extensive expeditions to Africa.

These are evergreen and deciduous treelets, shrubs and vines. The alternate leaves are petiolate and lack stipules. Leaf blades are oddly pinnate, trifoliate or simple, and petioles have a transversely ridged pulvinus. Leaflets are subopposite or alternate and usually leathery with an entire (rarely lobed) margin. Inf lorescences are terminal or axillary, bracteate panicles or racemes. The small actinomorphic flowers are bisexual or rarely unisexual. There are (four or) five sepals, free or united only basally, persistent and clasping the base of fruits. The five (or four) free petals are rarely slightly fused at the middle. Stamens are five or ten in two whorls

Agelaea borneensis in fruit, Singapore (KH) [206]

Jollydora sp., Etome Village, southwestern Cameroon (CD) [206]

Uses: Young shoots, leaves and seeds of all species can be used as a condiment called mufira. It has an aroma of garlic with fresh terpenic notes.

Zebrawood family

that are alternately longer and shorter, those opposite the petals often shorter and staminodial. Filaments are free or shortly fused at their base, the anthers dorsifixed and opening lengthwise. The disk is thin when present and surrounded by the base of the stamens. The superior or semi-inferior ovary is composed of usually five (rarely one to eight) free, one-celled, hairy carpels, each tipped with a thin style and a capitate uni- or bi-lobed stigma. Styles can be of different lengths in flowers on the same plant (heterostylous). Fruits are usually solitary, sessile or stalked follicles that open along the suture, rarely circumscissile at base or indehiscent. Seeds are usually arillate, the aril colourful and fleshy. Distribution: This pantropical family occurs mainly in Africa and tropical Asia, with only a few species in the New World. Phylogeny and evolution: Due to the similarity of their cuticular wax and compound leaves with a pulvinus similar to those of Fabaceae, the family was sometimes placed near them, although they differ in many characters. Molecular studies placed Connaraceae close to Oxalidaceae, with which they have many similarities. Genera and species: Connaraceae have c. 12 genera and c. 180 species: Agelaea (8), Burttia (1), Cnestidium (2), Cnestis (12), Connarus (77), Ellipanthus (7), Hemandradenia (2), Jollydora (4), Manotes (5), Pseudoconnarus (5), Rourea (c. 57) and Vismianthus (1). Uses: Vines of Agelaea have fibre used for making rope, and some Connarus and Ellipanthus species produce good hard wood (zebrawood). Plants are often poisonous but are sometimes applied medicinally. Etymology: Connarus is derived from ancient Greek κονναρος (konnaros), the name of a prickly evergreen shrub (possibly Paliurus spina-christi, Rhamnaceae), a name used by Linnaeus to describe this mainly tropical genus.

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shorter filaments. Filaments are fused at the base into a ring, and anthers are dorsifixed and open via lengthwise slits. The superior ovary is composed of five fused carpels forming five locules, topped with five free styles, each with a capitate or two-cleft stigma. Styles can be of different lengths in flowers on the same plant (heterostyly). Fruits are loculicidal capsules or fleshy berries with five pronounced ribs. Seeds often have a basal aril involved in their explosive ejection from the capsule; the aril is fleshy in Dapania.

207. OXALIDACEAE Wood-sorrel family

Perennial, rarely annual, herbs and shrubs, trees and vines make up this family. They are sometimes succulent with fleshy stems and petioles or have underground bulbs, tubers or scaly rhizomes. Leaves are alternate or whorled, clustered or in a basal rosette, the blades pinnately or palmately compound, sometimes quadrifoliate, trifoliate or unifoliate, often folding at night or in response to touch (Biophytum, Averrhoa), the petioles sometimes with small stipules at the base or stipules absent. Inflorescences are paniculate thyrses, often composed of cymes organised in umbels, spikes, racemes or heads or flowers solitary. The bisexual f lowers are actinomorphic, often with different types on a plant (heteromorphic, sometimes cleistogamous). The five sepals are free or basally fused, and the five petals are free or sometimes basally slightly fused, often clawed at base (petals are absent in cleistogamous flowers). The ten stamens are in two whorls, the outer whorl usually with Oxalis spiralis subsp. vulcanicola, San Francisco Botanical Garden, USA [207]

Distribution: This is a cosmopolitan family, but they are most diverse in tropical and subtropical regions, especially East Asia, southern Africa and South America. Oxalis is most diverse in southern Africa but extends into the temperate and subarctic regions in North and South America, Eurasia and Australia and New Zealand. The family is absent from the great deserts. Phylogeny and evolution: In morphologybased classsifications, the family was placed with Geraniaceae, but molecular evidence places Oxalidaceae close to Connaraceae, which was an association already suggested by some morphological similarities. An age of 34–56 million years is estimated for the family, which is not in line with the earlier suggestion that the current distribution of the family is due to tectonics. The woody Averrhoa, Dapania and Sarcotheca are closely related, supporting an Asian origin for carambola or starfruit

Averrhoa carambola, Santa Catarina, Brazil [207]

(A. carambola), which is unknown in the wild. The evolution of tunicate bulbs (unique among eudicots) allowed Oxalis to diversify in the winter-rain climates in the Cape of South Africa and southern South America, especially Chile. Hypseocharis was previously placed here, but DNA results showed it to be a member of Geraniaceae. Genera and species: This is a family of five genera and c. 570 species: Averrhoa (2), Biophytum (c. 50), Dapania (3), Oxalis (c. 500) and Sarcotheca (11). Uses: Oca (Oxalis tuberosa) is commercially grown in South America and locally an important alternative to potatoes. The starfruit or carambola (Averrhoa carambola) is a commonly grown fruit crop in the tropics, mainly for juice, and it is exported as an exotic fruit, often used for garnish of desserts, salads etc. The more acid bilimbi (A. bilimbi) is also grown for juice and pickles, and the sap is used as a bleaching agent. The acid fruits of Sarcotheca species are also locally eaten in curry. Leaves of many Oxalis species are edible in moderation. For example, leaves of O. acetosa (wood sorrel) can be chewed for their refreshing taste on the few hot days in a typical English summer. Etymology: Oxalis (οξαλις) is the ancient Greek name for garden sorrel (Rumex acetosa, Polygonaceae), which is similarly rich in oxalic acid but unrelated to Oxalis. Biophytum sensitivum, Helsinki Botanical Garden, Finland [207]

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Geissois pruinosa, New Caledonia [208]

Bauera rubioides, Australian National Botanic Gardens, Canberra [208]

Cunonia deplanchei, New Caledonia [208]

Weinmannia tinctoria, Réunion [208]

208. CUNONIACEAE

the node (interpetiolar). Inflorescences are terminal or axillary panicles, thyrses or cymes, sometimes in heads, always bracteate and bracts often with stipules. Flowers are rarely solitary in the leaf axils (Eucryphia). The actinomorphic flowers are bisexual, rarely unisexual and then on separate plants. The four or five (rarely three or up to ten) sepals are free or basally fused, and petals are as many as the sepals or absent in several genera, sometimes more numerous than sepals (Bauera). Stamens are usually twice as many as the sepals or more numerous in one to several whorls. The free filaments are long and slender, and anthers are usually dorsifixed and open by lengthwise slits or sometimes apically. A nectary disc is usually present, composed of segments or confluent into a ring. The ovary is superior to inferior and composed of two to five (up to 14 in Eucryphia) carpels that are fused or distinct, and each carpel is topped with a free style. The fruit can be a capsule or follicle or an indehiscent drupe or samara, the wings made up of the sepals or carpel walls, sometimes the carpels swollen and bladderlike (Platylophus).

Distribution: This is a mostly Southern Hemisphere family found throughout tropical America from Mexico and the Caribbean to southern Chile and Patagonia, the Cape of South Africa, Madagascar, the Malesian Archipelago north to the Philippines, Melanesia, New Caledonia, Australia, New Zealand and Polynesia. Weinmannia is widespread, occurring on New Caledonia, Indian Ocean islands and mountains of southern Mexico, the Antilles, Central and South America.

Butterspoon-tree family

Cunoniaceae are a family of trees and shrubs that are usually terrestrial but can be hemiepiphytic and strangling trees (some Weinmannia); they are mostly evergreen but deciduous in some species of Eucryphia. Leaves are opposite or whorled (three or four per node), rarely alternate and spirally arranged (Davidsonia). Blades are oddly pinnate, trifoliate or palmately compound, rarely unifoliate, the margins are toothed and often glandular, rarely entire, and venation is pinnate, with small tufts of hairs or domatia sometimes present along the midvein. Stipules are usually evident between the petioles on

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Pancheria hirsuta, New Caledonia [208]

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Phylogeny and evolution: The genera of Cunoniaceae were previously placed in Saxifragaceae, but since the family was established, they have had various circumscriptions, with Bauera, Eucryphia and Davidsonia either included or recognised in their own families. Phylogenetic analysis of DNA sequences has led to a recircumscription of this family and placed Cunoniaceae in Oxalidales, close to Elaeocarpaceae, Cephalotaceae and Brunelliaceae, the last also previously included in Cunoniaceae. Aphanopetalum (now Aphanopetalaceae, Saxifragales), Gumillea (still unplaced)

OXALIDALES

EUDICOTS

and Paracryphia (now Paracryphiaceae, Paracryphiales) were all once included in Cunoniaceae. The family has a fossil record in Australian Cenozoic deposits, with the oldest fossil known being a Late Palaeocene fossil leaf of Eucryphia, which suggests that the diversification of the family happened sometime during the Late Cretaceous. A flower in Burmese amber (c. 100 million years old) is similar to Ceratopetalum. Genera and species: Cunoniaceae include 27 genera and c. 330 species: Ackama (4), Acrophyllum (1), Acsmithia (16), Aistopetalum (2), Anodopetalum (1), Bauera (4), Caldcluvia (1), Callicoma (1), Ceratopetalum (9), Codia (12), Cunonia (25), Davidsonia (3), Eucryphia (7), Geissois (c. 18), Gillbeea (3), Hooglandia (1), Lamanonia (5), Opocunonia (1), Pancheria (c. 30), Platylophus (1), Pseudoweinmannia (2), Pullea (3), Schizomeria (10), Spiraeanthemum (6), Spiraeopsis (6), Vesselowskya (2) and Weinmannia (c. 155).

that can be used to make preserves and wine. Schizomeria fruits can be similarly consumed. Weinmannia tinctoria is used locally for tanning. Many species produce high-quality wood and are used for a variety of purposes, from general construction to cabinetry, furniture, boats, canoes, musical instruments and wagons. Weinmannia, Eucryphia and Platylophus forests are used for commercial honey production in Madagascar, New Zealand, Chile, Tasmania and South Africa. Eucryphia is sometimes grown as an ornamental, and cultivars are especially popular in Britain and New Zealand. In Australia, the New South Wales Christmas bush, Ceratopetalum gummiferum, is a popular ornamental.

209. ELAEOCARPACEAE Fairy-petticoats family

Uses: Davidson’s plum (Davidsonia pruriens) is sometimes cultivated for its edible fruit

Etymology: Cunonia is named for Dutch poet and garden appreciator Johan Christian Cuno (1708–1780), who wrote a poem describing his exotic Amsterdam garden in 1750. He also wrote a poem that accompanied the description of Wachendorfia (Haemodoraceae), honouring Dutch botanists E. J. van Wachendorff and J. Burman.

This is a family of (semi-)evergreen trees and shrubs, which can be large rainforest canopy trees with buttressed trunks or small ericoid shrubs of heathland. Their leaves are alternate and spirally arranged or in a plane, sometimes opposite or whorled. They are petiolate and sometimes have stipules. Blades are simple (rarely pinnately dissected or pinnate in juveniles) and have entire to serrate margins and pinnate venation, sometimes (Vallea) with three main veins at the base. Inf lorescences are axillary or terminal racemes, corymbs, panicles or fascicles, or the flowers are solitary. The bisexual flowers

Elaeocarpus grandiflorus, Singapore (WA) [209]

Vallea stipularis, Royal Botanic Gardens, Kew, UK [209]

Sloanea javanica, Singapore [209]

Crinodendron hookeriana, Kilmacurragh Botanic Gardens, Ireland [209]

Tremandra stelligera, Kings Park and Botanic Garden, Perth, Australia [209]

Sloanea caribaea, Guadeloupe [209]

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OXALIDALES are actinomorphic and subtended by bracts or not. The four or five sepals are basally fused or free, and the four or five petals (absent in some Sloanea) are free or fused and variously toothed or lobed, rarely entire. The four to 300 stamens have free filaments that are borne on or around a circular or lobed disk, and anthers are basifixed and open by one or two apical pores or lengthwise slits with the connective sometimes extended (awned) or tipped with hairs. The superior ovary is composed of two to nine locules topped with a simple single style that is branched at the tip or (rarely) the styles free. Fruits are drupes, berries or loculicidal capsules; they can be bristly or spiny outside in some Sloanea. Distribution: This family occurs mostly in tropical and subtropical regions of the Americas and Asia. They can be found from Mesoamerica and the Caribbean to southern Chile, Madagascar, tropical Asia, Japan, Malesia, New Guinea, Melanesia, Australia, New Zealand and Pacific islands including Hawaii. Remarkably, they are absent from Africa.

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in Oxalidales. The former Tremandraceae are embedded in Elaeocarpaceae as sister to Elaeocarpus+Sericolea+Aceratium. Even though the current distribution of the family is centred on the Southern Hemisphere, there are Late Cretaceous and Tertiary fossils known from Europe, Greenland and North America. The age of the family has been estimated to be 38–59 million years, although >100 million years has also been suggested.

Etymology: Elaeocarpus is composed of Greek ελαιο (elaio), oil, and καρπός (karpos), a fruit, in reference to the similarity of some fruits to an olive.

210. CEPHALOTACEAE Albany-pitcherplant family

Genera and species: This is a family of 12 genera and c. 615 species: Aceratium (20), Aristotelia (5), Crinodendron (5), Dubouzetia (11), Elaeocarpus (c. 350), Peripentadenia (2), Platytheca (2), Sericolea (c. 16), Sloanea (c. 150), Tetratheca (c. 50), Tremandra (2) and Vallea (2). These are perennial, evergreen carnivorous herbs. The stem is an underground knotty rootstock, with leaves in a rosette at its tip. Leaves lack stipules and are petiolate and of two forms, some spathulate and flat, pinnateparallel-veined, others pitcher-shaped and insect trapping. A third leaf type is found as scales on the rhizome, and all leaves have sessile glands. Pitchers are hooded, the body winged or ribbed and the mouth ribbed. Inflorescences are elongate thyrses composed of scorpioid cymes. The bisexual flowers are actinomorphic and sweetly scented. The perianth is of a single whorl (sepals) composed of six fused parts (petals absent). There are 12 stamens in two whorls that emerge from a papillose disk. Anthers are dorsifixed and

Phylogeny and evolution: Elaeocarpaceae were often included in Malvales in the past, and genera such as Muntingia (Muntingiaceae, Malvales) and Petenaea (Petenaeaceae, Huerteales) were previously included. Molecular evidence places Elaeocarpaceae close to Brunelliaceae and Cephalotaceae

Uses: Fruits of Aceratium oppositifolium and macqui (Aristotelia chilensis) are edible as are those of some Elaeocarpus species, e.g. blueberry ash (E. obovatus) and whitewood (E. kirktonii). Fruits of E. robustus and E. serratus are usually pickled or used in curry in South and Southeast Asia. Seeds of Elaeocarpus are also edible, but only largefruited species are worthwhile (for instance karanda nuts, E. bancroftii). The wood of some large Elaeocarpus and Sloanea species is used for timber. Several species of Aristotelia, Crinodendron, Tetratheca and Vallea are grown as ornamentals in temperate regions, whereas others are sometimes grown as ornamentals or shade trees in the tropics.

Cephalotus follicularis, Golden Gate Park, San Francisco, California, USA (CD) [210]

Cephalotus follicularis, near Albany, Western Australia [210]

Christenhusz, Fay & Chase

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versatile, with a connective appendage and thecae opening by lateral slits. The superior ovary is composed of six free carpels, each forming a single locule topped with a straight style. The fruit is a one- or two-seeded hairy follicle. Distribution: This family is confined to sandy and peaty swamps in southwestern Western Australia, particulary near the town of Albany. Phylogeny and evolution: Although the pitchers of Cephalotus are superficially similar to those of Nepenthes, the two genera differ in so many other important characters that they were not considered closely related, even before the advent of molecular systematics. Cephalotus has mostly been associated with Saxifragaceae or Crassulaceae, but molecular phylogenetic studies indicated that Cephalotaceae belongs to Oxalidales, within which they are related to Brunelliaceae (which have the same floral diagram) and then to Eleaocarpaceae and Cunoniaceae. Cephalotus is thus only distantly related to any other carnivorous taxa, and it is unusual in belonging to a clade in which glands are not always present. Mucilage-producing epidermal cells and hairs do occur in other families of Oxalidales (in some genera of Elaeocarpaceae and Cunoniaceae) and evidently can serve as antecedents for the development of the digestive and absorptive glands in Cephalotaceae. Genera and species: The sole species in this family is Cephalotus follicularis. Carnivory: Like other pitcher plants of the Americas (some but not all Sarraceniaceae) and Asia (Nepenthaceae), Cephalotus produces its own digestive enzymes and is thus an “active” carnivore (most species of Heliamphora, Sarraceniaceae, rely upon symbiotic bacteria to digest their captured prey). It attracts insects by ultraviolet absorption patterns in the traps and nectar glands inside the mouth of the pitcher. It grows in wet, sunny sites where other carnivorous taxa (e.g. Drosera species, Droseraceae, Caryophyllales) also occur.

Use: It is popular with carnivorous plant enthusiasts and is frequently grown in specialist collections. Etymology: The name is derived from Greek κεφαλωτος (kefalotos), headed, in reference to the floral disc (confused with filaments by the original author) that resembles a head.

211. BRUNELLIACEAE Palo-bobo family

Stamens are twice as numerous as sepals and in two whorls, the anthers dorsifixed and opening by lengthwise slits with a protruding connective. Staminodes are present in female flowers. Superior ovaries are composed of as many carpels as there are sepals, which are free and basally sunken in the disk. Each carpel is hairy and topped by a style that is hooked or curled at the tip with linear stigmas. Male flowers have rudimentary pistillodes. The fruit is composed of separate hairy follicles, each opening individually along the suture, the styles curving outward and the seeds staying attached to the follicle by a placental stalk. Distribution: This family is restricted to the Neotropics, from southern Mexico and the Greater Antilles throughout the Andes from Colombia to northern Bolivia and in the mountains of Venezuela (Paría Peninsula, Roraima Massif).

These evergreen trees have branches with angular internodes and swollen nodes. The simple to (evenly) pinnate leaves are opposite or in whorls of three. Blades have toothed to crenate margins and pinnate venation. Stipules are lateral, and stipels are often present at the insertion of the pinnae on compound leaves. Inflorescences are axillary thyrsoid panicles, the flowers with small prophylls. The actinomorphic flowers are bisexual or unisexual by abortion and then the plants unisexual. There is a single perianth whorl composed of four to six (rarely to eight) basally fused sepals; there are no petals. A cup-shaped, lobed nectary disc is adnate to the calyx and surrounds the stamen filaments at the base.

Etymology: Brunellia is named for 18th century Italian botanist G. Brunelli.

Brunellia comocladifolia, Guyana (PM) [211]

Brunellia comocladifolia (DS) [211]

Phylogeny and evolution: Traditionally the family was included in Cunoniaceae, with which it shares the stipulate opposite compound leaves with stipels. The free ovaries are reminiscent of Rosaceae, but this is a parallelism. Molecular studies have shown that their closest relative is Australian Cephalotaceae, which have an identical floral diagram. Genera and species: The single genus Brunellia includes c. 60 species.

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MALPIGHIALES

EUDICOTS

MALPIGHIALES Families 212 to 247 compose the large and diverse order Malpighiales, which started radiating during the Cretaceous c. 101–114 million years ago, probably at the same time when tropical rainforests developed, although it is also suggested that these did not develop before the start of the Tertiary. It is a large order comprising nearly 8% of eudicot diversity and is an important component of tropical rainforests. Placement of the COM clade (Celastrales, Oxalidales and Malpighiales) is here based entirely on plastid DNA analyses, but in results from mitochondrial and nuclear gene analyses, the COM clade are sister to the malvids instead of the fabids, which could be the result of ancient reticulation. At the time when the linear APG (Haston et al. 2009) was compiled, the relationships of the families in this order were not yet well known. We now have better estimates on relationships and the families are reorganised following APG IV (2016).

Panda oleosa, Ogooue River, Gabon (CD) [212]

Microdesmis puberula in fruit, Mokoko Forest Reserve, Cameroon (CD) [212]

212. PANDACEAE

or two whorls with free filaments that are sometimes unequal in length, the inner ones (all in female flowers) reduced to staminodes. Anthers are dorsifixed and open lengthwise, the thecae fused to an elevated connective. The superior ovary is composed of two to five fused carpels, each of one locule and topped with a two- to five-lobed style. The fruit is a drupe with a bony, hard endocarp, sometimes opening by valves upon germination, sometimes a capsule.

Kana-nut family

This is a family of unisexual trees and shrubs, the stems with prominent axillary buds and small stipules. Leaves are petiolate, alternate and arranged in a plane (distichous), the blades simple, with entire or serrate margins and pinnate venation. Inflorescences are terminal or cauliflorous raceme-like thyrses or axillary and fasciculate; in some the flowers are solitary. The actinomorphic flowers are unisexual with five free or fused sepals and five free petals, sometimes with a small disk. Male flowers have five to 15 stamens in one

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Distribution: This family occurs in tropical West and Central Africa and in tropical Southeast Asia throughout the Malesian Archipelago to the Solomon Islands. Phylogeny and evolution: Formerly these genera were part of Acalyphoideae of Euphorbiaceae, but molecular evidence placed these as sister to Irvingiaceae. Thus Pandaceae have been reinstated. It has been estimated that the family separated from Irvingiaceae sometime around 97–119 million years ago.

Microdesmis caseariifolia (WA) [212]

Genera and species: This family has three genera with 17 species: Galearia (5), Microdesmis (11) and Panda (1). Uses: Seeds of Panda oleosa contain c. 50% oil, which is extracted in West Africa for domestic use. They can also be eaten after cooking. The wood is much valued for carpentry and canoe-making. It is not cultivated due to its slow growth and erratic germination, but it remains a valuable native forest product. Etymology: Panda is named in honour of Congolese agronomist Paul Panda Fernana M’fumu (1888–1930), who was the first Congolese man to receive higher education (in Belgium and France) and actively denounced colonialism in Africa.

MALPIGHIALES

EUDICOTS

213. IRVINGIACEAE Ogbono-nut family

These evergreen trees (sometimes up to 90 m tall in Allantospermum borneense) have extremely hard wood and slime cells in leaves and stems. The alternate leaves are simple and petiolate and have large unequal, interpetiolar stipules that encircle the bud. Blades have an entire margin and pinnate venation. Inf lorescences are axillary or terminal panicles with small flowers. Its bisexual flowers are actinomorphic. The five small sepals are free, and the five free petals exceed them. The ten stamens are longer than the petals and inserted around a nectary disk. Anthers are almost basifixed, and thecae open by slits. The superior ovary sits upon the nectary disk and is composed of four or five carpels, each forming a locule. A single style with a short pointed stigma tops the ovary. The fruit is a drupe or a broadly winged samara (former Desbordesia).

with one species (Irvinga malayana) in tropical Asia and another genus (Allantospermum) in Peninsular Malaysia and Borneo. Phylogeny and evolution: Due to the presence of mucilage cells, Irvingiaceae were previously associated erroneously with Simaroubaceae or Ixonanthaceae. They were later included in a broad Linaceae, but some authors recognised this family, as corroborated by molecular analyses. Their exact placement in Malpighiales has been contentious, different phylogenetic analyses giving different placements, although the latest most comprehensive treatment places them as sister to Pandaceae. Fossil wood from the Pleistocene of Ethiopia (Irvingiaceoxylon) shows that in the past this area was covered in rainforest. Allantospermum was previously placed in Ixonanthaceae, but it fits better here morphologically and genetically. Desbordesia is embedded in Irvingia. Genera and species: This family has three genera and 12 species: Allantospermum (2), Irvingia (8) and Klainedoxa (2).

valued for their high levels of protein and fat. They are grown commercially in Cameroon. The nuts are also ground to make dika bread or Gabon chocolate. Etymology: Irvingia was named for Scottish navy surgeon and plant collector Edward George Irving (1816–1855).

214. CTENOLOPHONACEAE Litoh family

These are trees with buttressed trunks and tufts of stellate hairs on shoots, stipules and perianths. Leaves are opposite, simple and petiolate with interpetiolar stipules that soon fall off. Blades have an entire margin and pinnate venation. Inflorescences are axillary or terminal, bracteate, thyrsoid panicles, and flowers are bisexual and regular. The five basally fused sepals are swollen and persistent in fruit. The five petals are contorted in bud and usually shortly clawed at the base. A welldeveloped cup-shaped disk surrounds the ten stamens. Filaments are basally fused into a short corona, and dorsifixed anthers open

Distribution: The family is found in moist forests of West and Central Africa,

Uses: Irvingia gabonensis bears edible drupes that resemble mangos. The nuts of I. gabonensis have been shown to be effective in weight-loss treatments, often marketed as “African mango”, a common name that causes confusion with the unrelated mango fruit (Mangifera indica, Anacardiaceae) and explains the images on the packaging. Seeds of Irvingia are called ogbono nuts and are

Irvingia gabonensis, fruit, Aburi Botanic Garden, Ghana (CD) [213]

Irvingia gabonensis seedlings sprouting from elephant dung, Gabon (AP) [213]

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MALPIGHIALES

EUDICOTS

was placed in various families in the past, including Celastraceae, Linaceae, Icacinaceae and Oleaceae but is now treated as a family close to Erythroxylaceae and Rhizophoraceae, with which it shares some characters. Genera and species: This family has a single genus, Ctenolophon, with two geographically and palynologically isolated species, C. englerianus and C. parvifolius. Etymology: Ctenolophon is derived from the Greek κτένα (ktena), a comb, and λόφων (lofon), hills.

215. RHIZOPHORACEAE Mangrove family

Ctenolophon parvifolius. Illustration from Oliver (1873) Descriptions of three new genera from the Malayan Herbarium of the late Dr. A. C. Maingay. Translactions of the Linnean Society of London 28: 515–518, plate XLIII [214]

lengthwise. The superior ovary is composed of two fused carpels, each forming a locule. The fruit is a one-celled woody capsule that splits lengthwise in two halves exposing a solitary seed that is retained in the capsule.

Rhizophora samoensis, Nouméa, New Caledonia

These are evergreen trees and shrubs that often have aerial roots and swollen nodes. They have interpetiolar stipules that sheath the terminal bud and fall off when the leaf develops. The simple petiolate leaves are opposite or in whorls, often clustered at the shoot tips. Leaf blades are leathery, and margins are entire or apically or completely serrulate, with pinnate venation. Inflorescences are slender axillary cymes or fascicles. The bisexual flowers are actinomorphic often with a hypanthium. Paradrypetes has unisexual, petal-less flowers. The three to 16 sepals are free or slightly fused at the base and inserted on the rim of the hypanthium (if present). Petals are as many as sepals and are free with the margin entire, lacerate, or two-cleft, often with a terminal arista and filiform appendages on the two lobes, each petal usually enveloping one to five stamens. Stamens are twice as many as sepals or numerous. Filaments are borne around the base of an entire or lobed nectary disk and are free or fused, short in Paradrypetes. Anthers are dorsifixed, near the base of the connective, and they open lengthwise or by a valve. The superior to

[215]

Pellacalyx axillaris, Singapore (WA) [215]

Rhizophora samoensis, New Caledonia [215]

Bruguiera gymnorhiza, Royal Botanic Gardens, Kew, UK [215]

Distribution: This is a palaeotropical family, found in West Africa (Nigeria to Angola), the Malayan Peninsula, Sumatra, Borneo, Banka, the southern Philippines (Mindanao, Leyte, Samar) and New Guinea, possibly relictual in nature. Phylogeny and evolution: Ctenolophon englerianus-like plants have an extensive fossil record dating back to the Upper Cretaceous (Maastrichtian) of Central Africa. Palynological finds have shown that this pollen type was widespread across the tropics in the Palaeocene and Eocene but disappeared from many regions, including the Neotropics and the Indian subcontinent, during the Tertiary. The earliest Ctenolophon parvifolius pollen types are found in Palaeocene deposits in Africa, from where it spread to India and Southeast Asia during the Tertiary. Ctenolophon

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inferior ovary is composed of two to five (20) carpels forming up to eight locules, topped with a single style (three in Paradrypetes) and an entire or lobed, capitate stigma. The fruit is a pulpy or leathery indehiscent capsule, berry or drupe with one to many seeds. Germination while seeds are still on the tree (vivipary) is common, allowing the seedling to form a thick sturdy root (radicle; up to 80 cm long) for establishment in muddy soil in shallow seas or river beds. Carallia and Pellacalyx germinate on the ground. Distribution: This pantropical family, typically in mangroves in warm shallow seas, extends north to Bermuda, the Red Sea and southern Japan, and south to southern Brazil and northern New Zealand. Phylogeny and evolution: Traditionally the genera of Anisophylleaceae were thought to belong to this family, but molecular studies have shown that Anisophylleaceae belong to Cucurbitales, whereas Rhizophoraceae are most closely related to Erythroxylaceae in Malpighiales. Crown Rhizophoraceae have been dated to c. 50 million years, the family diverging from Erythroxylaceae c. 60–63 million years ago. Fossils are known from

the mangrove genera Ceriops and Bruguiera in the Early Eocene, London Clay, and from Kandelia from the Middle Eocene of Alaska. Genera and species: Rhizophoraceae include 16 genera and c. 147 species: Anopyxis (1), Blepharistemma (1), Bruguiera (6), Carallia (11), Cassipourea (c. 62), Ceriops (2), Comiphyton (1), Crossostylis (c. 12), Dactylopetalum (15), Gynotroches (1), Kandelia (2), Macarisia (7), Paradrypetes (2), Pellacalyx (7), Rhizophora (8) and Sterigmapetalum (9). Uses: The wood of many mangrove species is used for making charcoal or as ornamental wood in terraria, resulting in over-exploitation. Mangroves (especially Bruguiera and Rhizophora) are important for coastal storm protection and extensive programmes are now carried out replanting mangroves to prevent coastal erosion and to stabilise soil. Timber of some species (e.g. Carallia) is commercially valuable. Etymology: Rhizophora is composed of the Greek ρίζα (rhiza), a root, and φόρων ( foron), duty or carrying, in reference either to the stilt-roots or the seeds germinating on the tree.

Erythroxylum coca, Hortus botanicus, Leiden, the Netherlands [216]

216. ERYTHROXYLACEAE Coca family

This family of evergreen and deciduous shrubs and trees sometimes has milky or coloured sap. Leaves are simple and alternate or rarely opposite with two stipules on either side of the petiole. Leaf blades have an entire margin with pinnate venation and often have two parallel lines or folds on the leaf blade (because young leaves are involute in bud, leaving these lines). Inflorescences are axillary fascicles or (stalked umbellate) cymes. The usually bisexual flowers are actinomorphic. The five (rarely four) sepals are basally fused and persistent in fruit. The five (or four) petals are free and often shortly clawed, usually with a bilobed appendage at their base on the inner side. The usually ten, rarely five, 12 or 20 stamens, usually in two whorls with filament

Erythroxylum sechellarum in fruit, Seychelles [216]

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MALPIGHIALES bases fused into a tube or cup. Anthers are basifixed and open with lengthwise slits. The superior ovary is composed of three to four fused carpels, each forming a locule, with all or only one of the locules fertile. Styles are two to three, distinct, partly or completely fused, with oblique stigmas; they are frequently of different lengths. Fruits are one-seeded drupes or two- to three-seeded capsules. Distribution: The family has a pantropical distribution with centres of diversity in South America and Madagascar. Phylogeny and evolution: Formerly the family was considered closely related to Linaceae, with which it shares leaf anatomical and seed characteristics. Molecular analyses placed them close to Rhizophoraceae, with which they share involute leaf blades and similar chemistry (protein crystals in sieve tube plastids, similar alkaloids). These families have been merged by some authors, but are now maintained separate but considered closely related; together they are sister to Ctenolophonaceae. Some Eocene fossils of Erythroxylaceae are known from Argentina. Genera and species: This family includes four genera and c. 242 species: Aneulophus (2), Erythroxylum (c. 230), Nectaropetalum (8) and Pinacopodium (2). Uses: Leaves of coca (Erythroxylum coca and E. novogranatensis) are chewed by the Andean people often mixed with alkaline lime or plant ash as a catalyst, a habit dating back thousands of years. Coca leaves were already an important trade commodity during the time of the Incas. Coca leaves work as a stimulant, allowing the chewer to endure hunger and exhaustion and to cope with low oxygen levels at high elevations. Apart from cocaine, the leaves contain many vitamins and minerals and are locally not considered a drug. A tea from coca leaves is also commercially available, but in this form cocaine content is low. Large quantities of leaves are needed just to produce a few grams of crystallised cocaine. Cocaine as a narcotic drug has serious

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EUDICOTS

social, political and economic impacts on communities and nations around the world, which is why it is banned in most countries, although eradication of plantations in South America is not possible due to the cultural importance of this sacred plant to the native Andean people. During the 19th century, a number of products based on coca extracts were commercially marketed, especially in candies, biscuits, cigarettes, wine and soft drinks. These became popular as stimulants, medicines and, of course, recreational drugs. Because a number of people became addicted to alcoholic drinks like Vin Tonique Mariani and Pemberton’s Wine Coca and because of the popularity of Coca Cola and other soft drinks in the USA, cocaine has been removed from these drinks since 1906. Coca Cola and Red Bull Cola still contain coca leaf extract, but without the cocaine, whereas Agwa de Bolivia, Coca Sek and Coca Colla are drinks with natural coca extracts available in Bolivia. Catuaba (E. vacciniifolium) is used as an aphrodisiac in Brazilian drinks. Wood of some species of Erythroxylum and Nectaropetalum is hard and durable.

217. OCHNACEAE Mickey Mouse-plant family

Etymology: Erythroxylum is derived from the Greek ερυθρός (erythros), red, and ξύλων (xylon), wood.

These are evergreen trees, shrubs and sometimes herbs that are bisexual or occasionally unisexual (Quiinoideae). Their leaves are alternate in Ochnoideae, opposite in Medusagynoideae and whorled to opposite in Quiinoideae, simple or rarely pinnately compound (pinnately compound in juveniles, simple in adults in many species of Quiinoideae) and petiolate. Stipules are free or sometimes fused at the base between the petioles, usually entire but sometimes laciniate, sometimes leaf-like, bristle-like or absent. Leaf blades have serrate, (glandular) toothed or ciliate, rarely entire margins with pinnate venation. Inflorescences are terminal or axillary bracteate cymes or racemes, rarely uniflorous or in bunch-like thyrses (Froesia). The usually bisexual

Quiina blackii, Rio Yanayacu, Pacaya-Samiria, Nauta, Loreto, Peru (CD) [217]

Medusagyne oppositifolia, Mahé, Seychelles [217]

MALPIGHIALES

EUDICOTS

Sauvagesia erecta, Guadeloupe [217]

Campylospermum flavum, Royal Botanic Gardens, Edinburgh, UK [217]

Ochna kirkii in fruit, Hong Kong Zoological and Botanical Gardens, China [217]

Lophira alata, fruit, Korup National Park, Cameroon (CD) [217]

Ochna mauritiana, Réunion [217]

(often unisexual in Quiinoideae) flowers are mostly actinomorphic, rarely zygomorphic, with geniculate pedicels. The usually five (sometime as few as two or up to 15) sepals are free or rarely fused at the base, sometimes unequal. Petals are also usually five (rarely three to eight) and mostly free and often clawed and contorted in bud. In Quiinoideae, petals and sepals are sometimes adnate to form a floral cup. Stamens are five, ten, 15 or numerous, sometimes fascicled, with free and persistent filaments, or the anthers sessile. Anthers are basifixed and open by lengthwise slits or apical pores. Staminodes are sometimes present and sometimes petallike and free, sometimes fused into a tube. Nectaries are not present. The superior ovary is composed of three or five (sometimes two to 15, or 16–25 in Medusagyne) carpels that are fused at least at their base or along the central axis. The ovary is sometimes stalked (gynophore) and can be entire or deeply lobed, generally topped by a long style and an entire or divided stigma or each carpel

topped with a free style and capitate stigma (Medusagynoideae and Quiinoideae). Fruits are fleshy or not but usually a septicidal capsule, sometimes a nut with accrescent persistent sepals or a drupe. In some genera the carpels separate into blackish drupelets on a coloured accrescent receptacle (e.g. Ochna) or each carpel opens from the central column along the margin (Medusagyne and some Ochnoideae); it is a berry-like capsule in Quiinoideae. Seeds are winged or not.

(Sauvagesioideae of traditional Ochnaceae) and Luxemburgiaceae (Luxemburgioideae of traditional Ochnaceae), which together with Testulea form a grade leading to core Ochnoideae. The family in the broad sense is characterised by having mucilage cells, stratified phloem and several floral characters, such as lack of floral nectaries and petals contorted in the bud. The circumscription of Sauvagesia, which is not monophyletic in the latest phylogenetic studies and the subdivision of and relationships in Ochninae require further investigation.

Distribution: The family has a pantropical distribution, with Sauvagesia erecta being a pantropical weed. Medusagyne oppositifolia (Medusagynoideae) is endemic to the island of Mahé in the Seychelles. Quiinoideae are only found in the Neotropics (from Belize to Brazil). Phylogeny and evolution: Formerly treated as separate families in Theales, subfamilies Medusag y noideae and Quiinoideae together are sister to Ochnoideae, which include the former families Sauvagesiaceae

Genera and species: This is a family of 32 genera with c. 550 species in three subfamilies: Medusagynoideae – Medusagyne (1); Quiinoideae – Froesia (6), Lacunaria (c. 10), Quiina (c. 34) and Touroulia (2); Ochnoideae – Adenarake (2), Blastemanthus (3), Brackenridgea (7), Campylospermum (c. 65), Cespedesia (6), Elvasia (15), Euthemis (2), Fleudydora (1), Godoya (2), Idertia (4), Indosinia (1), Krukoviella (1), Lophira (2), Plants of the World

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Luxemburgia (18), Ochna (c. 85), Ouratea (c. 200), Perissocarpa (3), Philacra (4), Poecilandra (3), Rhabdophyllum (8), Rhytidanthera (5), Sauvagesia (c. 40), Schuurmansia (3), Schuurmansiella (1), Testulea (1), Tyleria (13) and Wallacea (2). Uses: The seeds of Lophira are used to make cooking oil. Lophira alata makes excellent timber, which is hard and durable. South African Ochna serrulata (bird’s eye bush) is widely cultivated in tropical gardens for its yellow flowers and peculiar “bird’s eye” fruit, which is reminiscent of the head of Walt Disney’s cartoon character Mickey Mouse.

Ploiarium alternifolium, Singapore Botanical Garden [218]

Ploiarium alternifolium in fruit, Singapore Botanical Garden [218]

Bonnetia paniculata, Rio Wawaime, Cordillera del Condor, Ecuador (CD) [218]

Bonnetia stricta, near Mucugê, Bahía, Brazil [218]

bundles. Anthers are basifixed, opening laterally. A superior ovary with three to five locules sits in the middle topped by free stylodia or these fused into a simple style with papillate stigmas. The fruit is a capsule that opens around the persistent central column.

be in Calophyllaceae, making Bonnetiaceae more homogeneous.

Conservation: Bois meduse or jellyfish tree (Medusagyne oppositifolia) was thought to be extinct until it was rediscovered in the 1970s on the island of Mahé. Since then, it has been a flagship species for plant conservation in the Seychelles and elsewhere. Etymology: Ochna is Latinised Greek of ochne, a wild pear tree.

218. BONNETIACEAE Cascarilla family

These trees and shrubs have spirally arranged leaves at the tips of branches. Leaves are simple on a short petiole or sessile and lack stipules. Blade margins are finely toothed with setae, and venation is pinnate-parallel with closely ascending veins running parallel to the main vein. Inflorescences are bracteate cymes or simple flowers, the pedicels with two prophylls or several bracts. The bisexual flowers are actinomorphic and have five unequal free sepals and five free petals. Stamens are numerous and have free slender filaments or basally fused filaments in five

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Distribution: Bonnetiaceae have a disjunct distribution in tropical America and Asia. They occur in Cuba, continental South America and southern Indochina and Malesia. Phylogeny and evolution: Previously thought to be intermediate between the unrelated Theaceae and Clusiaceae or included in the polyphyletic Guttiferae, Bonnetiaceae were found in DNA analyses to be sister to Clusiaceae. Several genera (Caraipa, Kielmeyera and Neotatea) were previously included here, but these have been shown to

Genera and species: Bonnetiaceae are a family of three genera and 35 species: Archytaea (2), Bonnetia (c. 30) and Ploiarium (3). Uses: The durable wood is sometimes harvested locally in Asia. Etymology: Bonnetia is named for Swiss philosopher and naturalist Charles Bonnet (1720–1793), who was a pioneer in physiological psychology. Bonnet was the first to use the term evolution in a biological context, and he developed the catastrophe theory, in which the earth undergoes natural catastrophies periodically, which aids the surviving taxa in their evolutionary development.

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EUDICOTS

219. CLUSIACEAE Mangosteen family

formed singly. The bisexual or unisexual f lowers are actinomorphic and usually subtended by prophylls. The two, four or five (to 20) sepals are usually free, sometimes fused, and the usually four or five (sometimes three or up to eight) petals are free, sometimes absent. The numerous (but sometimes as few as four) stamens are free or grouped in bunches or fused in bundles. Anthers open by longitudinal slits. The superior ovary has one to five (rarely 20) locules, with styles free or fused into one or the stigmas sessile. Fruits are berries or septicidal capsules exposing the winged or arillate seeds.

These evergreen shrubs and trees can grow terrestrially or epiphytically, the stems frequently producing prop-roots or kneeroots, and epiphytic species may only make secondary contact with the ground through aerial roots. All parts of the plants are glandular and often have resin canals exuding white or yellow latex when damaged. The opposite (rarely verticillate) leaves lack stipules, but paired glands may sometimes be present. Blades are simple, usually petiolate with pinnate venation, often with closely set parallel secondary venation united by an intramarginal vein. Inf lorescences are terminal or axillary modified cymes, sometimes the f lowers

Phylogeny and evolution: Clusiaceae were previously variously treated (often including Calophyllaceae, Hypericaceae and Bonnetiaceae) as Guttiferae, an alternative name, which was associated with Theaceae

Clusia mangle, Guadeloupe [219]

Symphonia globulifera, Braulio Carillo National Park, Heredia, Costa Rica (CD) [219]

Clusia rosea, Curaçao (AP) [219]

Garcinia brasiliensis, New York Botanical Garden, USA [219]

Distribution: Clusiaceae are a pantropical family occurring from Mexico and the Antilles to southern Brazil, throughout SubSaharan Africa, Madagascar, southern India and Sri Lanka, Southeast Asia, southern China throughout the Malesian Archipelago to New Guinea, Melanesia and northeastern Australia.

s.l., but this was a heterogeneous assembly. Aquatic Podostemaceae were found to be sister to Hypericaceae and hence the former Guttiferae were dismembered (although the alternative of an expanded Clusiaceae would also have been an option, but was not preferred by most taxonomists). Placed in Malpighiales, the ‘guttifer’ clade is sister to Ochnaceae. Known from a Late Cretaceous fossil from New Jersey, Clusiaceae are probably at least 90 million years old. Eocene and Miocene fossil wood and pollen assignable to Clusiaceae are frequently found, sometimes outside the current range of the family, suggesting a wider range in the past. Genera and species: Clusiaceae include 14 genera and c. 750 species: Allanblackia (10), Chrysochlamys (57), Clusia (c. 350), Dystovomita (4), Garcinia (c. 260), Lorostemon (5), Montrouziera (5), Moronobea (7), Pentadesma (5), Platonia (1), Symphonia (23), Thysanostemon (2), Tovomita (c. 26) and Tovomitopsis (3). Uses: One of the finest tropical fruits, the mangosteen (Garcinia mangostana) is widely cultivated. The inner part of the fruit (pericarp) Clusia major in fruit, Netherlands Antilles (AP) [219]

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MALPIGHIALES

Mammea americana, Guayaquil Botanical Garden, Ecuador (CD) [220]

EUDICOTS

Caraipa tereticaulis, Allpahuayo-Mishana Reserve, Loreto, Peru (CD) [220]

is eaten and has a creamy acid-sweet flavour. Other Garcinia species also produce edible fruit but are often more acidic; these are used to flavour curries. Fruits of bacuri or manil (Platonia insignis and Moronobea coccinea) and bakupari (Garcinia brasiliensis) are sometimes cultivated for their fruits in South America. The oily seeds of Allanblackia are used as a butter alternative in West Africa, and seeds of kokam (Garcinia indica) yield butter used in Asia. The latex of some species provides a dye called gamboge, the origin of the country name Cambodia. A fruit extract of Garcinia gummi-gutta is marketed under its synonym, G. cambogia, as a weightloss supplement. Etymology: The genus Clusia is named in honour of Flemish physician and botanist Carolus Clusius (Charles de l’Écluse, 1529– 1609). He was the most famous of 16th century horticulturalists and prefect of the imperial medicinal garden in Vienna. After making botanical expeditions to the Iberian Peninsula, the Austrian Alps and the Pannonian Plains, he helped to establish the Hortus botanicus of Leiden University in the Netherlands, the first botanical garden north of the Alps. Among many other plants, he is responsible for the introduction of horse chestnuts (Aesculus hippocastanum, Sapindaceae) into Western Europe and for popularisation of tulips (Tulipa, Liliaceae) in the Netherlands. 310

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Calophyllum inophyllum, Seychelles [220]

220. CALOPHYLLACEAE Takamaka family

septicidal capsules, and seeds lack arils but can be embedded in a fleshy pericarp. Distribution: This pantropically distributed family is found from the Caribbean and Mexico to Argentina, tropical West and Central Africa, Madagascar and other Indian Ocean and Pacific islands, southern India, Southeast Asia and the Malesian Archipelago east to New Guinea, New Caledonia, Melanesia, Polynesia and northern Australia.

This family of evergreen trees usually has all parts with glands and/or resin canals exuding white or yellow latex. The opposite or alternate leaves lack stipules, but there may sometimes be paired glands. Blades are simple, usually petiolate with pinnate venation and closely set (or not) parallel secondary venation; tertiary veins are well developed. Inflorescences are terminal or axillary modified cymes or racemes, or flowers are solitary. The bisexual or unisexual flowers are actinomorphic (rarely zygomorphic in Marila asymmetralis). The two, four or five (up to 20) sepals are usually free, sometimes fused, and the usually four to six (or up to eight) petals are free, sometimes absent. The numerous (or just four) stamens are free and normally not grouped in bundles. Anthers open by longitudinal slits and are often fixed to a glandular connective. The superior ovary has one to five (rarely 20) locules with a simple fused style. Fruits are

Phylogeny and evolution: Due to the frequently spirally arranged leaves, an association with similarly spirally leaved Theaceae was made. Calophyllaceae were later included as subfamily Kielmeyeroideae in Clusiaceae, but this renders that family polyphyletic. Endodesmia is sister to the rest of the family. Genera and species: Calophyllaceae include 14 genera and c. 475 species: Calophyllum (190), Caraipa (22), Clusiella (8), Endodesmia (1), Haploclathra (5), Kayea (70), Kielmeyera (c. 50), Lebrunia (1), Mahurea (2), Mammea (c. 70), Marila (c. 20), Mesua (30), Neotatea (4) and Poeciloneuron (2). Uses: Mammey apples (Mammea americana) are a tropical delicacy. Flowers of Mammea siamensis are used in the perfume industry. Seeds of Calophyllum yield minor oils that can

MALPIGHIALES

EUDICOTS

be used for cooking and soap making. Wood of ironwood (Mesua ferrea) is extremely hard. Several species are of ornamental value, especially takamaka (Calophyllum inophyllum), which is frequently planted along tropical beaches or as a street tree and occasionally used for timber. Etymology: Calophyllum is composed of the Greek words κάλλος (kallos), beauty, and φύλλων ( fyllon), a leaf.

221. PODOSTEMACEAE Riverweed family

Aquatic perennial and annual herbs that often resemble seaweeds, algae, mosses or lichens, make up this unusual family. They are attached with flattened, thalloid or filiform roots to rocks or other hard surfaces in fresh, rapidly flowing streams, rapids or waterfalls. Leaves arise from buds directly from the roots and are distichous, scattered or imbricate, the base of the blade often sheathing, sometimes with stipule-like appendages, and blade margins entire, lobed or dissected and completely submerged. Inflorescences emerge when the water level drops; these are solitary or clustered racemes or cymes, sometimes flowers surrounded by a cupule or enclosing Macarenia clavigera, Colombia (MF) [221]

bract (spathella). The bisexual flowers are actinomorphic or zygomorphic, opening above the water or cleistogamous under water. A complete or incomplete whorl of two to five (or up to 20) tepals is present, often only on one side of the flower, free or more or less fused. There are one to 40 stamens, in one or two complete or incomplete whorls, or stamens are fused into a Y-shaped structure carrying two stamens. Filaments can be fused or free, and anthers open by longitudinal slits. The superior ovary is uni-, bi- or tri-locular and topped by two or three styles. Fruits are smooth or ribbed septicidal capsules, splitting in two or three equal valves and exposing numerous small seeds. Distribution: This family is widespread in tropical regions, with a few species in warm-temperate areas. Plants are found in clean fresh-water streams, rapids, cataracts and waterfalls, where they are attached to submerged rocks or wood. Water pollution and eutrophication are a major threat to these species. Phylogeny and evolution: Because of their divergent morphology and as a result of their aquatic habitat, relationships of this family were long considered speculative. An association with Hydrostachyaceae (Cornales), a similarly odd family of submerged aquatics had been suggested, but these are not closely related, their seaweed-like morphology being a case of convergent evolution. Molecular results place Podostemaceae in the old Clusiaceae s.l. (Guttifereae) in Malpighiales, as sister to Hypericaceae. The oldest fossil records of Podostemaceae are found in Upper Eocene

Marathrum squamosum, Colombia (MF) [221]

formations in Russia. Three subfamilies are generally recognised. Some genera may need to be recircumscribed, and consolidation may be appropriate in the future. Genera and species: This family consists of c. 46 genera and some 300 species in three subfamilies: Tristichoideae – Cussetia (2), Dalzellia (5), Indodalzellia (1), Indotristicha (2), Terniopsis (8) and Tristicha (2); Weddellinoideae – Weddellina (1); Podostemoideae – Angolaea (1), Apinagia (c. 50), Autana (1), Aulea (1), Castelnavia (6), Ceratolacis (2), Cipoia (2), Cladopus (9), Diamantina (1), Dicraeanthus (2), Djinga (1), Endocaulos (1), Farmeria (1), Griffithella (1), Hanseniella (2), Hydrobryum (23), Hydrodiscus (1), Inversodicraea (20), Jenmaniella (7), Ledermanniella (26), Leiothylax (3), Letestuella (1), Lophogyne (1), Macarenia (1), Macropodiella (6), Maferria (1), Marathrum (10), Monostylis (1), Mourera (8), Noveloa (2), Oserya (5), Paleodicraeia (1), Paracladopus (2), Podostemum (11), Polypleurum (c. 18), Rhyncholacis (23), Saxicolella (5), Sphaerothylax (2), Stonesia (5), Thawatchaia (1), Thelethylax (2), Wettsteiniola (3), Willisia (2), Winklerella (1), Zehnderia (1) and Zeylanidium (6). Uses: Dicraeanthus africanus can be eaten as salad. Rhyncholacis species are dried and used as a seasoning similar to pepper. In large species like Mourera fluviatilis, salt accumulates, which can be extracted by burning. Etymology: Podostemum is the Latinised form of the Greek words ποδών (podon), a foot and, στεμα (stema), a crown or stamen. Marathrum rubrum, Colima, Mexico [221]

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MALPIGHIALES

Cratoxylum cochinchinense, Hong Kong Zoological and Botanical Gardens, China [222]

222. HYPERICACEAE St. John’s-wort family

EUDICOTS

Hypericum perforatum, Suomenlinna, Helsinki, Finland [222]

Vismia brasiliensis, Serra de Maranguape, Ceará [222]

These evergreen and deciduous shrubs, perennial and annual herbs contain resin or oil glands and/or canals in most parts, the exudates often being yellow or purple. The opposite leaves are simple, sessile or shortly petiolate and lack stipules. Blades are often dotted with pellucid and/or black glands and usually have entire margins and pinnate venation. Inflorescences are terminal (rarely axillary) cymes or panicles, rarely flowers single. The bisexual or unisexual flowers

Hypericum lanceolatum, Réunion [222]

are actinomorphic and often have bracteoles just below the calyx inserted on the pedicel. Sepals are four or five (rarely two) and free. The four or five (rarely three) petals are contorted in bud and often oblique. Stamens are numerous (rarely nine), free and usually aggregated into three, four or five bundles, or the filaments are variously fused. Anthers are dorsifixed and open by longitudinal slits, the connective often bearing glands, and staminodes are sometimes present.

A third system of classification — Robert Folger Thorne (1920–2015) The former curator emeritus at Rancho Santa Ana Botanic Garden in California conducted research in the evolution of flowering plants during his long career. He also worked on plant communities in Australasia and California, and he advocated the conservation of endangered natural environments, resulting for instance in the conservation of the flora and fauna of Santa Catalina Island and other unusual natural areas in California. His system of angiosperm classification, first published in 1983, and updated in 1992, 2000 and 2007, is placed in the ’intuitive’ tradition established by Bessey, but organises families into a ’phyletic shrub’ based on similar characters. Family delimitations varied among the several versions of the Thorne System, the later ones incorporating some results of molecular studies, but a full list of synonymy helps place families not accepted in the system. On the basis of molecular studies and later anatomical and chemical analyses, we now know that some of these families were misplaced, but the system was a great step forward in the understanding of plant evolution and biogeography and should be considered in parallel with the Cronquist and Takhtajan systems that found more followers. The Central American genus Thornea (Hypericaceae) was named in honour of him in 1976.

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Photograph of Robert Thorne in 1960 by F.W. Kent at the Iowa State University Herbarium in Ames.

MALPIGHIALES

EUDICOTS

The superior ovary is composed of three to five carpels forming one locule or as many locules as carpels, each having a free style or with them fused more or less into a single style. Stigmas are broad, smooth and sticky or pointed and papillate. Fruits are berries or septicidal capsules, the seeds small and sometimes winged.

223. CARYOCARACEAE Souari-tree family

Distribution: This family has a worldwide distribution but is less common in dry regions and wet tropical lowlands. Phylogeny and evolution: The fossil record of the family is minimal, and many putative fossil hypericaceous seeds may actually be assignable to other families. Use as an age calibration for fossil seeds like Hypericum antiquum may not be useful. This has, however, been done, resulting in an age of 53.8 million years, although divesification of Hypericum probably did not happen earlier than 10 million years ago. Genera of this family were usually included in Clusiaceae s.l. in the past, but the position of Podostemaceae as their sister resulted in acceptance of the family Hypericaceae. Generic delimitations need to be revisited, especially with regard to Triadenum versus Hypericum and Harungana (including Psorospermum).

These are trees and shrubs with opposite and alternate, trifoliate leaves. Petioles have two to four stipules or the stipules absent; leaflets have serrate margins and often have stipels at their base. Inflorescences are terminal racemes with articulate pedicels. The bisexual flowers are large and actinomorphic. The five (or six) sepals are free, and the five (or six) petals are usually free but sometimes basally fused, in Anthodiscus fused at the top to form a hood (calyptra). Stamens are numerous, usually more than 50, up to c. 750, with the long and slender filaments usually fused in a ring at the base, often with sterile staminodia Caryocar microcarpum, Rio Momon, Iquitos, Loreto, Peru (CD) [223]

on the inner side that are sometimes recurved, and often with spirally arranged vesicles along their length and basally with glandular nectaries in Caryocar. Anthers are basifixed and open by slits. After pollination, stamens often fall with the petals as an entity. The superior ovary is composed of four (to six) fused carpels in Caryocar and eight to 20 fused carpels in Anthodiscus, each topped with a single style and a punctiform stigma. The fruit is a drupe with one to four or eight to 20 seeds, eventually splitting into pyrenes or mericarps. Caryocar is pollinated by bats. Distribution: The family can be found around the Neotropics from Costa Rica to southern Brazil. Phylogeny and evolution: In most traditional systems, Caryocaraceae were placed in Theales, but molecular analyses firmly place them in Malpighiales. Genera and species: This is a family of two genera and 26 species: Anthodiscus (10) and Caryocar (16). Anthodiscus chocoensis, Rancho Quemado, Osa Peninsula, Puntarenas, Costa Rica (CD) [223]

Genera and species: Hypericaceae include nine genera and c. 590 species: Cratoxylum (6), Eliea (1), Harungana (perhaps 50), Hypericum (c. 470), Lianthus (1), Santomasia (1), Thornea (2), Triadenum (6) and Vismia (c. 50). Uses: There are a number of medicinal applications, like hypericin, extracted from St John’s wort (Hypericum perforatum), which is used to alleviate depression. Several species are cultivated as garden ornamentals but can potentially become weedy. Wood of Cratoxylum is soft and easily worked into drums and roof shingles in Indonesia.

Caryocar brasiliense, Brazil (JD) [223]

Etymology: Hypericum was hung above pictures to ward off evil spirits, hence its name is composed of the Greek υπέρ (hyper), above, and εικών (eikon), an image, picture or icon.

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MALPIGHIALES Uses: Several species of Caryocar produce edible nuts (souari or sawarri nuts), the best known being pekea nut, C. nuciferum, and pequi nut, C. brasiliense, the latter also pressed to yield an oil in Brazil. The trees also produce a strong timber, and C. costaricense has been overexploited for this, resulting in listing on Appendix 2 of CITES. Etymology: Caryocar is composed of the Greek κάρυο (karyo), a nut, and καρδιά (kardia), a heart.

224. LOPHOPYXIDACEAE Koteb family

EUDICOTS

These are bisexual, climbing shrubs and treelets with tendrils derived from axillary branchlets that look like watch springs often bearing a bud. Their regular branches have a lateral bud at their base, and leaves are arranged alternately and spirally around the stem. Leaf blades are simple, petiolate, the margins serrulate to crenulate with pinnate venation. Stipules are reduced and knob-like. Inflorescences are loose axillary panicles with the basal branches transformed into tendrils and apical branches bearing clusters of flowers. The unisexual flowers are actinomorphic and have five basally fused sepals and five free petals that are much smaller than the sepals and set upon a yellowish nectar disk. Male flowers have five (sometimes six) stamens with filiform filaments that emerge between two glands that are fused to the petals. Anthers are basifixed and open with lengthwise slits. A half-globular pistillode is present and hairy. Female flowers have the glands fused into a lobed disk, and the superior ovary is

Lophopyxis maingayi, illustration based on herbarium specimens [224]

composed of (four or) five fused carpels each forming a locule and each topped with a free style. The fruit is a single-seeded indehiscent capsule with five crests or wings. Distribution: The family is distributed throughout the Malesian Archipelago, east to the Solomon and Caroline Islands and west to Peninsular Malaysia. Phylogeny and evolution: Lophopyxis was formerly variously placed, most frequently in Euphorbiaceae, Icacinaceae or Celastraceae. Molecular analyses place it in Malpighiales as sister to Putranjivaceae, with which it shares several characters. Genera and species: This family consists of the single genus Lophopyxis with two species: L. maingayi and L. pentaptera. Etymology: Lophopyxis is derived from the Greek λόφων (lofon), a crest or hill, and πυξις (pyxis), a box, referring to the fruit.

Drypetes natalensis, Durban Botanic Garden, South Africa (CD) [225]

Drypetes euryodes in fruit, Monts de Cristal, Tchimbele, Gabon (CD) [225]

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MALPIGHIALES

EUDICOTS

Bhesa paniculata, Singapore (KH) [226]

225. PUTRANJIVACEAE Childlife-tree family

Bhesa robusta, Singapore (WA) [226]

three (or to six) fused carpels, each forming a locule, with as many styles as locules. Stigmas are peltate, reniform, disc-shaped, bilobed or petal-like. The fruit is a drupe. Distribution: This family is found throughout the tropical and subtropical regions.

This family of usually unisexual trees and shrubs produces glucosinolates (mustard oils). These plants are characterised by their simple or stellate hairs on their fruits. They have alternate, rarely opposite, leaves that often appear to grow in a plane (distichous). Blades are simple, petiolate, usually oblique at their base, the margins entire or (prickly) toothed, and venation is pinnate. Stipules are persistent or not. Inflorescences are axillary or in cauliflorous clusters or pedicellate cymes; sometimes the f lowers are solitary. The normally unisexual flowers are actinomorphic, rarely bisexual or polygamous. They usually have four or five (sometimes three or six) free sepals and no petals. Male flowers have two or three (Putranjiva) or four to 20 stamens (sometimes fewer or up to 50) with free filaments and subdorsifixed anthers that open by lengthwise slits. An intrastaminal disk and a small pistillode are sometimes present. Female flowers often have an annular disk. The superior ovary is composed of one to

Phylogeny and evolution: In morphologybased classifications, members of this family were placed in Euphorbiaceae subfamily Phyllanthoideae (tribe Drypeteae) with Lingelsheimia, which remains in Phyllanthaceae on the basis of its different morphology. Molecular studies have provided strong support for Putranjivaceae being sister to Lophopyxidaceae in Malpighiales. Sibangea, a genus of three species, is often accepted as separate from Drypetes, but it is embedded in that genus. Putranjivaceae are the only family that produce mustard oils outside of Brassicales, a chemical pathway that has evolved in parallel. Genera and species: This family has two genera with 216 species: Drypetes (212) and Putranjiva (4). Uses: Fruit of Indian boxwood, Drypetes sepiaria, is eaten in India and Sri Lanka, and several other species of Drypetes are eaten locally. The seeds of the childlife tree, Putranjiva roxburghii, are pressed to yield an oil that is used for burning and as an essential oil. The bark is used medicinally. This

species is widely cultivated as an ornamental throughout the tropics. Etymology: Putranjiva is derived from Sanskrit putra, son, and juvi, life or prosperity, which is also the basis for its common name.

226. CENTROPLACACEAE Biku-biku family

These evergreen, unisexual (Centroplacus) and bisexual (Bhesa) trees often have buttressed trunks. Their simple, alternate leaves are two-ranked or spirally arranged and have two stipules that often almost encircle the stem; the petiole is frequently pulvinate. Leaf blades have entire or toothed margins with pinnate venation. Inflorescences are axillary panicles, thyrses or racemes, and flowers are actinomorphic and usually bisexual, unisexual in Centroplacus. There are five free sepals and five free petals. Female flowers of Centroplacus lack petals. A nectary disc surrounds the stamens. The five stamens are free, the anthers basifixed with obliqueapical dehiscence. The superior ovary has

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MALPIGHIALES two or three locules, each topped with a free style that is spreading with expanded stigmas. The fruit is a septicidal or loculicidal capsule, splitting from the base, each locule with a single seed that has a fleshy aril (Bhesa) or caruncle (Centroplacus). Distribution: Centroplacus occurs in West Africa, whereas Bhesa occurs in tropical Asia, from southern India, Sri Lanka and Bengal through southern China and Malesia to New Guinea. Phylogeny and evolution: The two genera in this family are distinct morphologically. Bhesa was formerly placed in Celastraceae on the basis of gross morphology but always was oddly placed there in morphological terms. Centroplacus was variously placed in the now defunct Flacourtiaceae or Celastraceae but also considered part of Phyllanthoideae in Euphorbiaceae, where it was also a poor match. The two genera are closely related despite their morphological differences and form a clade sister to Malpighiaceae plus Elatinaceae. Genera and species: The two genera of Centroplacaceae include seven species: Bhesa (6) and Centroplacus glaucinus.

EUDICOTS

Etymology: Centroplacus is Greek composed of κέντρων (kentron), a sharp point, and πλακος (plakos), a flat plane, referring to the style.

227. ELATINACEAE Waterwort family

Elatinaceae are perennial and annual, aquatic and moist habitat herbs, rarely small shrubs (Bergia suffruticosa). They are glabrous (Elatine) or glandular/hairy (Bergia) and often dwarf and easily overlooked or confused with other similarly small plants of wet habitats. Leaves are opposite, rarely whorled (Elatine alsinastrum), with paired stipules on both sides of the petioles. Simple leaf blades have entire or serrate margins with pinnateparallel venation. Inflorescences are axillary cymes (dichasia) or clusters, or flowers are solitary in the leaf axils. The bisexual flowers

are actinomorphic, small and sometimes cleistogamous. The two to five sepals are free or basally fused, often with transparent margins. The two to five petals are free and persistent in fruit. There are as many or twice as many stamens as petals in one or two whorls. Anthers are dorsifixed and open via lengthwise slits. The superior ovary is usually three- to five-loculate and topped by three to five styles with capitate stigmas. The fruit is a thin-walled septicidal capsule with numerous tiny seeds. Distribution: The family is found in wet or seasonally wet habitats around the world, often aquatic in shallow water. They are absent or only locally distributed in the tropics and more common in the temperate zones, although Bergia is diverse in ephemeral pools of tropical Australia. Phylogeny and evolution: The family was formerly associated with Caryophyllaceae or Clusiaceae, but molecular analyses place the family close to Malpighiaceae (Malpighiales), with which they share only a few characters. Genera and species: Elatinaceae include two genera and c. 35 species: Bergia (c. 25) and Elatine (c. 10).

Malpighia glabra, Santa Catarina, Brazil [228]

Bergia perennis, Northern Territory, Australia

[227]

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Elatine alsinastrum, Finland (HV) [227]

Malpighia glabra, fruit, Santa Catarina, Brazil

[228]

MALPIGHIALES

EUDICOTS

Tristellateia australasiae, Seychelles [228]

Acridocarpus austrocaledonica, New Caledonia [228] Galphimia brasiliensis, Rio de Janeiro, Brazil [228]

Hiptage benghalensis, Réunion [228]

Uses: Elatine species are sometimes grown as unusual aquarium plants.

Distribution: This pantropical family is found north to southwestern USA and southern Florida, Yemen, the Himalayas and southern China, and south to Uruguay and Argentina, South Africa and northern New South Wales, with the greatest diversity in the Neotropics.

Etymology: Elatine is the feminine form of the Greek ελατινος (elatinos), from the silver fir, probably of Armenian origin (elevin). The word was used by Pliny for a species of snapdragon (possibly now Kickxia elatine, Plantaginaceae). It is unknown why the name was chosen for this genus.

228. MALPIGHIACEAE Acerola family

This family includes shrubs, trees, perennial herbs and woody and herbaceous vines that have a mix of simple and two-armed ‘compass-hairs’ (malpighioid hairs). Leaves are usually opposite, rarely alternate or in whorls of threes, and usually have free or fused stipules. Petioles often bear glands,

or the glands are on the lower leaf blade, which is simple with an entire (rarely lobed) margin, sometimes falsely toothed or ciliate around glands, and has pinnate venation. Inf lorescences are terminal or axillary racemes or panicles composed of corymbs or umbels of four or more f lowers but sometimes flowers are solitary. The bisexual or staminate (in some Stigmaphyllon) flowers are actinomorphic or more commonly zygomorphic and have two bracteoles at the point of articulation along the pedicel. The five sepals are free or partially fused, and Neotropical species almost always bear one or two glands basally per sepal on the outside of the calyx. The five petals are typically clawed and have a ciliate, toothed or fimbriate margin, and the upper petal (flag) is often different in size, colour and orientation from the others. The ten stamens (sometimes fewer or up to 15 in Lasiocarpus) are equal in length or one stamen much larger than the others. Filaments are usually fused at least at the base, and anthers open inwards by longitudinal slits or pores. Ovaries are superior (half inferior in Barnebya) and have three locules topped with three free styles, which may be fused into a single style. Fruits are schizocarps splitting into three or fewer winged samaras or indehiscent drupes.

Phylogeny and evolution: The crown group of Malpighiaceae has been estimated to be c. 32–36 million years old. The oldest fossils corroborate this estimate, being found in the Eocene Claiborne formation in Tennessee, which is c. 34 million years old. The closest relatives of Malpighiaceae are Elatinaceae, which may have separated some 94 million years ago. Both families have foliar glands and produce latex. The Acmanthera, Byrsonima and Galphimia clades are together sister to the rest of Malpighiaceae and are often placed in a separate subfamily: Byrsonimoideae. Lophanthera and Tetrapterys are not monophyletic in their traditional sense. There is evidence that the Neotropical distribution was achieved through dispersal from the Old World tropics via the North Atlantic (the Boreotropic hypothesis). Neotropical genera are pollinated by oil-collecting bees that grab the flag petal with their mandibles and scrape oil from the calyx glands, which is mixed with pollen and fed to their larvae. Genera

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MALPIGHIALES without calyx glands, in both the Neo- and Palaeotropics, produce nectar as a reward. In the Neotropics, members of the orchid subtribe Oncidiinae (Orchidaceae) mimic the flowers of the family and attract (through deceit) oil-collecting bees. Genera and species: Malpighiaceae are a family of 77 genera and c. 1,315 species in two subfamilies: Byrsonimoideae – Acmanthera (7), Blepharandra (6), Byrsonima (> 135), Coleostachys (1), Diacidia (11), Galphimia (26), Lophanthera (5), Pterandra (15), Spachea (6) and Verrucularia (2); Malpighioideae – Acridocarpus (29), Adelphia (4), Aenigmatanthera (2), Alicia (2), Amorimia (10), Aspicarpa (12), Aspidopterys (c. 17), Banisteriopsis (92), Barnebya (2), Brachylophon (3), Bronwenia (10), Bunchosia (c. 75), Burdachia (3), Calcicola (2), Callaeum (11), Camarea (7), Carolus (6), Caucanthus (3), Christianella (5), Cordobia (2), Cottsia (3), Diaspis (2), Dicella (7), Digoniopterys (1), Dinemagonum (1), Dinemandra (1), Diplopterys (4), Echinopterys (2), Ectopopteris (1), Excentradenia (4), Flabellaria (1), Flabellariopsis (1), Gallardoa (1), Gaudichaudia (10), Glandonia (3), Heladena (1), Henleophytum (1), Heteropterys (> 140), Hiptage (c. 25), Hiraea (c. 55), Janusia (17), Jubelina (6), Lasiocarpus (4), Lophopterys (7), Madagasikaria (1), Malpighia (c. 130), Malpighiodes (4), Mascagnia (c. 45), Mcvaughia (1), Mezia (10), Microsteira (25), Mionandra (1), Niedenzuella (16), Peixotoa (29), Peregrina (1), Philgamia (4), Psychopterys (8), Ptilochaeta (5), Rhynchophora (2), Sphedamnocarpus (12), Stigmaphyllon (c. 100), Tetrapterys (c. 90), Thryallis (5), Triaspis (12), Tricomaria (1), Triopterys (3) and Tristellateia (21). Uses: Barbados cherry or acerola (Malpighia emarginata) is a cultigen domesticated from wild, smaller fruited and also edible M. glabra. Both species are eaten fresh or made into juice, ice cream and alcoholic beverages. Fruits of chaparro or nanche (Byrsonima crassifolia) are harvested from wild and cultivated trees and sold in local markets to eat fresh or for ice-cream or wine. Other species with berry-like fruits are also eaten,

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such as ciruela (Bunchosia sp.). Ayahuasca is an infusion of the bark of Banisteriopsis caapi, often used for its psychotropic effects (tryptamines) and in combination with other hallucinogenics like Diplopterys cabrerana and Psychotria viridis (Rubiaceae). Callaeum antifebrile also reportly has hallicinogenic properties. Several species are popular ornamentals in the tropics, particularly locustberry (Byrsonima lucida), Callaeum macropterum (formerly Mascagnia macroptera), C. septentrionale, goldshower (Galphimia gracilis), Hiptage benghalensis, Lophanthera lactescens, Singapore holly (Malpighia coccigera), Spachea elegans, Stigmaphyllon ciliatum and Tristellateia australasiae.

filaments with anthers that open laterally, sometimes surrounding a rudimentary pistil. Female flowers form singly in axils of scaleleaves and are subtended by several spirally arranged, cup-shaped bractlets. The perianth is missing and the (superior) ovary is naked and composed of usually two (rarely three) fused carpels, each forming a locule. The ovary is topped by three basally fused styles that are once or twice forked at the tip, forming ribbonlike stigmas. The fruit is a fleshy berry with two or three hard seeds, and it is subtended by the persistent cup-like bracts.

Etymology: Malpighia is named in honour of Italian physician and biologist Marcello Malpighi (1628–1694), who was the first to employ microscopy in the study of anatomy and histology.

Phylogeny and evolution: Relationships of Balanopaceae have long been debated, and the genus was thought perhaps to be closest to Daphniphyllaceae, Euphorbiaceae, Hamamelidaceae or Pittosporaceae in the past. They somewhat resemble Myricaceae but lack the

Distribution: This family occurs in the southwestern Pacific, Vanuatu, New Caledonia, Fiji and northeastern Australia.

229. BALANOPACEAE Pimplebark family

Balanops pancheri, male flowers, New Caledonia (JM) [229]

This is a family of unisexual, evergreen trees and shrubs with white, circular lenticels on their bark. The alternate leaves are dimorphic, the small scale-like leaves are spread across the stem and followed by foliar leaves that often are restricted to the tips of branches or organised in false whorls. Foliar leaves are petiolate with two basal teethlike stipules, toothed margins and pinnate venation. Male inflorescences are catkins, the flowers positioned in the axils of a bract. Individual male flowers are subtended by one or several bractlets or a lobed rudimentary perianth. The three to six (sometimes as few as one or up to 12) stamens have short free

Balanops pancheri, female flowers, New Caledonia (JM) [229]

MALPIGHIALES

EUDICOTS

distinctive glands. Molecular analyses placed the family in Malpighiales, as sister to a clade including Dichapetalaceae and Trigoniaceae plus Chrysobalanaceae and Euphroniaceae, even though morphological characters would fit better with the euphorbioid rather than chrysobalanoid families in Malpighiales. Genera and species: The sole genus in this family, Balanops, has nine species. Etymology: Balanops is derived from the Greek word βάλανος (balanos), a glans or acorn, for its resemblance of the fruit to an oak seed. The alternative spelling Balanopsidaceae would linguistically be more appropriate and has been used in the past, but Balanopaceae is a conserved name with that spelling.

Trigonia nivea, Chapada Diamantina, Bahía, Brazil [230]

230. TRIGONIACEAE Triangle-vine family

These are trees, shrubs (sometimes scandent) and woody vines. Leaves are simple and alternate or opposite and petiolate with stipules often fused and interpetiolar (when leaves are opposite). Blades have an entire margin and pinnate venation. Inflorescences are panicles, thyrses or racemes, sometimes reduced to cymes, and pedicels have two bracteoles. The bisexual flowers are obliquely zygomorphic (papilionaceous), with the plane of symmetry through the third sepal. The five sepals are unequal in size and fused at the base or some sepals free. The five petals are free, contorted in bud, the two anterior petals forming a (saccate) keel, the standard petal also often saccate and the lateral petals (wings) spathulate. The four to eight stamens and zero to six staminodes are basally fused into a tube that is more developed on the keelside of the flower. The dorsifixed anthers open

Trigonia nivea, Chapada Diamantina, Bahía, Brazil [230]

by lengthwise slits. Up to four (or no) nectar glands are placed opposite the standard petal. The superior (or partly inferior) ovary has three (or four) locules with a simple terminal style and a capitate stigma. The fruit is a septicidal capsule or three-winged samara. Distribution: This disjunct family is most diverse in the Neotropics but also found in Madagascar (Humbertiodendron) and Malesia (Trigoniodendron). Phylogeny and evolution: The family was previously often associated with Polygalaceae, Linaceae or Vochysiaceae, but associations with Chrysobalanaceae and Dichapetalaceae have also been made in the past. Molecular analyses have shown that the last two are the

Trigonia villosa, near Matoury Reserve, French Guiana (CD) [230]

closest relatives of Trigoniaceae. Euphroniaceae were previously sometimes included in Trigoniaceae, but that family is closer to Chrysobalanaceae. Genera and species: Trigoniaceae are a tropical family of five genera and 28 species: Humbertiodendron (1), Isidodendron (1), Trigonia (24), Trigoniastrum (1) and Trigoniodendron (1). Uses: Trigoniastrum hypoleucum has hard wood that is sometimes used to make furniture. Etymology: Trigonia is composed of the Greek words τριγωνο (trigono), triangle, in reference to the three-parted fruits.

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MALPIGHIALES

Tapura guianensis, Camp Caiman, Kaw Mts, French Guiana (CD) [231]

Euphronia guianensis, Puerto Yuruaní, Bolívar, Venezuela (PM) [232]

Dichapetalum eickii, Taita Hills, Kenya [231]

Euphronia acuminatissima, Colombia (MF) [232]

231. DICHAPETALACEAE Ratbane family

This is a family of trees, shrubs and woody vines. Their simple leaves are alternately arranged, and blades have pinnate venation with an entire margin. Petioles have stipules that usually soon fall off. Inflorescences are corymbose cymes, heads or fascicles that are axillary or terminal, often attached to the petiole or the midrib of the leaf. The bisexual (rarely unisexual) flowers are actinomorphic or weakly zygomorphic and often have kneed (articulate) pedicels. The five sepals are equal 320

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or unequal in size and free or fused, sometimes forming a tube. The five petals are free and nearly equal in size or fused into a tube with the lobes equal or unequal and are usually clawed at the base and bilobed or bifid at the apex, often somewhat inflexed or cucullate. The usually five stamens are all fertile or up to three staminodial, often the filements fused to the corolla tube or receptacle. Anthers are dorsifixed and open lengthwise. A superior ovary with two to four locules is topped by a simple style with as many lobes as locules, rarely two to four styles free. The fruit is a dry or fleshy drupe. Distribution: This pantropical family extends into temperate South Africa. Phylogeny and evolution: Many different relationships were suggested in morphologybased classifications, but molecular analyses placed Dichapetalaceae close to Chrysobalanaceae, Euphroniaceae and Trigoniaceae in

Malpighiales. Structurally, Dichapetalaceae are most similar to Trigoniaceae. Genera and species: The family has three genera and c. 168 species: Dichapetalum (c. 135), Stephanopodium (13) and Tapura (20). Etymology: Dichapetalum is Latinised Greek, composed of the words δίχα (dicha), in twos, and πετάλων (petalon), a petal, referring to the bilobed petals.

232. EUPHRONIACEAE Euphronia family

MALPIGHIALES

EUDICOTS

Phylogeny and evolution: Originally published in Bonnetiaceae and later classified under Vochysiaceae or Rosaceae, the genus Euphronia was segregated into its own family because it is substantially different from these last two families. Morphological and molecular analyses place it close to Trigoniaceae in Malpighiales.

Distribution: Euphroniaceae are restricted to white-sand savannas and heath forests of northern South America (Venezuela, Guyana and adjacent Brazil).

This family of trees and shrubs have alternate leaves with stipules that are persistent or not

and sometimes fused to the petiole. Blades are simple with pinnate venation and an entire margin. Inflorescences are racemes, panicles or cymes, the branches bracteate and each flower with two bracteoles. The bisexual, rarely polygamous or unisexual flowers are actinomorphic to zygomorphic and have a short to elongate receptacle (hypanthium) that is sometimes gibbous at the base. A nectar disk forms a lining to the receptacle or is an annular or shortly tubular structure at the mouth of the receptacle. The five sepals are often unequal and can be reflexed. The five petals are often unequal in size and soon fall off, sometimes absent or clawed. Stamens are few (two or three) to numerous (100–300) and inserted on the hypanthium margin or on the disk surface, often the filaments fused to the disk at their bases; they can all be fertile or partially reduced to staminodes and lack an anther. Anthers are dorsifixed and open lengthwise. The superior ovary is inserted at the base, in the mouth or along the side of the hypanthium and has one or two locules. The style is filiform and fused to the hypanthium at the base of the ovary, the stigmas are trilobed. The fruit is a fleshy or dry drupe, which is often densely hairy within.

Maranthes corymbosa in fruit, Singapore (WA) [233]

Hirtella racemosa, Brazil (IP) [233]

Chrysobalanus icaco, fruit, Seychelles [233]

Parastemon spicatum, Bako National Park, Sarawak, Malaysia (CD) [233]

Couepia paraensis, Brazil (IP) [233]

Chrysobalanus icaco, Guadeloupe [233]

Trees and shrubs with spiderweb-like hairs make up this family. Their alternate leaves are simple and petiolate with tiny stipules. Blades have entire margins with pinnate venation. Inflorescences are terminal or nearly terminal racemes or thyrses with minute bracts. The bisexual flowers are unevenly zygomorphic with a cup-shaped receptacle that is lined with a nectary. The five sepals are fused basally and placed on the receptacle rim, the two outer sepals are narrower and shorter than the inner three. The three petals are free and of the same size, contorted in bud and clawed at their base. The two fertile stamen pairs are of two lengths and flank a large, hairy, pointed staminode; they are surrounded by two groups of two or three tooth-like staminodes. Filaments are fused, and the dorsifixed anthers open by lengthwise slits. The semiinferior ovary is triloculate and topped with a hairy style and capitate stigma. The fruit is a three-valved, septicidally dehiscent capsule with three seeds.

Genera and species: This family has a single genus, Euphronia, with three species. Etymology: Euphronia is composed of Greek ευ (eu), good, and φρόνως ( fronos), to think.

233. CHRYSOBALANACEAE Cocoplum family

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MALPIGHIALES

Vantanea obovata, Brazil (JD) [234]

EUDICOTS

Sacoglottis gabonensis, Fougère Brulée, Lope Humiria balsamifera, French Guiana [234] Reserve, Gabon (CD) [234]

Distribution: This pantropical family has its greatest diversity in the Neotropics, extending into the temperate zones in the southern USA and southern Brazil. Phylogeny and evolution: In most morphology-based classifications, this family was considered a subfamily of Rosaceae or a separate family in Rosales, but they differ markedly from members of Rosaceae and Rosales in their gynobasic style and wood anatomy, among other characters. Molecular analyses place Chrysobalanaceae in Malpighiales, close to Trigoniaceae, Dichapetalaceae and Euphroniaceae, which share anatomical floral characteristics. Genera and species: Chrysobalanaceae have 27 genera and 536 species: Acioa (6), Afrolicania (1), Angelesia (1), Atuna (8), Bafodeya (1), Chrysobalanus (3), Cordillera (1), Couepia (59), Dactyladenia (32), Exellodendron (5), Gaulettia (9), Geobalanus (3), Grangeria (2), Hirtella (109), Hunga (11), Hymenopus (29), Kostermanthus (3), Leptobalanus (31), Licania (100), Magnistipula (13), Maranthes (12), Microdesmia (2), Moquilea (49), Neocarya (1), Parastemon (3), Parinari (39) and Parinariopsis (1). Uses: Many species have edible fruits and are locally consumed. Icacos or cocoplums, the fruit of Chrysobalanus icaco, are often made into preserves or canned in syrup. Other edible fruits locally marketed are Couepia bracteosa and C. rufa. Parinari seeds have been found 322

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in archaeological sites in Malawi, and seeds of Atuna excelsa are the main ingredient of a dish called koku-koku in Ambon. Oils are pressed from some species, that of Acioa edulis and Couepia longipendula is used for cooking, and Licania arborea is flammable and used as lamp oil. Formerly L. rigida and Afrolicania elaeosperma oils were used in paint. Their wood and bark contains silica bodies, and the latter is thus hard and difficult to work. However, it is resistant to marine borers and can thus be used in coastal settings. Etymology: Chrysobalanus is composed of the Greek words χρυσός (chrysos), golden, and βάλανος (balanos), a glans or acorn, in reference to the fruits of some species.

234. HUMIRIACEAE Umiri family

Humiria balsamifera, fruit, Brazil (UM) [234]

or serrulate margin with pinnate venation. Inflorescences are axillary or terminal, often corymbiform panicles, branched in twos or threes with small bracts. The bisexual flowers are actinomorphic and pedicellate. The five sepals are more or less fused and the same size or the outer two smaller. The five petals are free and often cup-shaped (cochlear). Stamens are ten to 30 or numerous and usually in one or two whorls of different lengths, all fused into a tube at the base. Anthers are dorsifixed, and each theca opens with a lengthwise slit or by detachment. The connective is thick and fleshy, often with a tipped or tongue-shaped appendix. A cup-shaped or tubular disk is formed inside the stamen tube surrounding the ovary, which is often toothed, lobed, laciniate or composed of scales. The superior ovary is composed mostly of five fused carpels, each forming a locule. The single style tops the ovary and bears a five-lobed or five-radiate, capitate stigma. The fruit is a drupe with pulpy to fibrous flesh and a woody seed. Distribution: This family is mostly distributed in the Neotropics from Costa Rica to southern Brazil, with a single species in coastal West Africa (Sacoglottis gabonensis).

This is a family of evergreen trees and shrubs that often have balsam-scented wood. The alternate leaves are simple, often in a plane (distichous) and usually petiolate (rarely sessile) with (or without) small, lateral stipules. Blades have an entire, crenulate

Phylogeny and evolution: In early classifications, Humiriaceae were already placed in Malpighiales, in which they were often associated with Linaceae and Erythroxylaceae. Molecular analyses confirm this position, although their relationships within Malpighiales are less certain. Vantanea is sister to the

MALPIGHIALES

EUDICOTS

rest of the family, which dates back to the Eocene. Fossil seeds are found throughout South America in Neogene deposits. Pollen similar to that of extant Humiria has been found in early Miocene deposits from the Andes and western Amazonia.

These are bisexual, rarely unisexual, trees, shrubs and herbs, sometimes climbing (Ceratiosicyos) with clear sap. Leaves are alternate and spiral or two-ranked along the stem or aggregated into basal rosettes (Guthriea) with stipules and pulvinate petioles. Leaf blades are simple with an entire, crenate or serrate, lobed or dissected margin and pinnate venation. Crushed leaves may smell strongly

of cyanide. Inflorescences are few-flowered fascicles (cymes), spikes or racemes, or flowers are solitary in the leaf axils. The actinomorphic flowers are functionally unisexual, the male flowers lacking a pistillode. Flower parts may be in two distinct whorls or spirally arranged and not distinct. The two to five sepals are fused at the base with the lobes longer than the tube and the calyx opening in bud. The three to 15 petals are basally fused into a campanulate tube. There is no nectary present. Male flowers have three to five (to many) stamens and three to five staminodes that are unequal in length (the staminodes shorter), and the filaments are free but basally fused to the corolla tube. Anthers open inwardly by lengthwise slits, rarely pores, and are basifixed with a broadly expanded connective. Female flowers have a superior ovary that is composed of two to ten fused carpels forming a single locule topped with a single or branched styles and as many

Carpotroche longifolia with pollinators, Miraflores, Rio Tigre, Nauta, Loreto, Peru (CD) [235]

Hydnocarpus anthelminthica, Hong Kong Zoological and Botanical Gardens, China [235]

235. ACHARIACEAE Chaulmoogra family

Genera and species: This family includes eight genera and c. 56 species: Duckesia (1), Endopleura (1), Humiria (4), Humiriastrum (c. 16), Hylocarpa (1), Sacoglottis (8), Schistostemon (9) and Vantanea (16). Uses: Oil of the seeds of several species is used by indigenous people of the Amazon Basin. Etymology: Humiria is derived from houmiri, a Carib name for H. balsamifera from French Guiana. The name was originally published as Houmiri, but was later altered to Humiria by J. Saint-Hilaire.

Acharia tragodes, Eastern Cape, South Africa (CD) [235]

Grandidiera boivinii, Helsinki Botanical Garden, Finland [235]

Ryparosa hullettii, Singapore (MN) [235]

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MALPIGHIALES stigmas as carpels. The fruit is a dehiscent berry or capsule, with angular seeds that often have vascular bundles on the coat and are frequently surrounded by fleshy arils. Distribution: The family occurs pantropically but is especially diverse in Africa and Asia. Phylogeny and evolution: Originally the family only included three peculiar African genera (Acharia, Ceratiosicyos and Guthriea), but they were expanded to accommodate an additional 27 genera from the now superfluous, polyphyletic family Flacourtiaceae. Support for monophyly of Achariaceae in this circumscription is strong, although subclassification is still debated. The family has been dated to be c. 23 million years old. Cerolepis is an older name of Camptostylus, but combinations have not been made. Genera and species: This family includes 31 genera and c. 154 species: Acharia (1), Ahernia (1), Baileyoxylon (1), Buchnerodendron (2), Caloncoba (10), Camptostylus (3), Carpotroche (11), Ceratiosicyos (1), Chiangiodendron (1), Chlorocarpa (1), Dasylepis (7), Eleutherandra (1), Erythrospermum (4), Grandidiera (1), Guthriea (1), Gynocardia (1), Hydnocarpus (c. 40), Kiggelaria (1), Kuhlmanniodendron (2), Lindackeria (13), Mayna (6), Pangium (1), Peterodendron (1), Poggea (4), Prockiopsis (3), Rawsonia (2), Ryparosa

Rinorea anguifera, in fruit, Singaporet (WA) [236]

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(18), Scaphocalyx (2), Scottellia (3), Trichadenia (1) and Xylotheca (10). Uses: Keluak or kepayang, Pangium edule, is used to make a thick gravy called rawon. The seeds are poisonous when fresh and need to be boiled and fermented before consumption. There are selected cultivars with lower levels of cyanide compounds. Oil pressed from the seeds is sometimes used for cooking. Some Hydnocarpus species produce chaulmoogra oil used in Indian traditional medicine to treat leprosy, eczema and other skin conditions. Other members of this family are used for similar purposes elsewhere. Some tree-like species produce good timber (e.g. Kiggelaria, Scottellia). Etymology: Acharia is Latinised from Greek αχάρης, acharis, ungracefulness or clumsiness.

236. VIOLACEAE Violet family

Viola abyssinica, Taita Hills, Kenya [236]

This is a family of annual and perennial herbs, shrubs, vines and small trees. The simple leaves are petiolate (or nearly sessile) and usually alternate, sometimes opposite or in false whorls. Stipules are always present, scale- or leaf-like, in some species soon falling off. Leaf blades are entire or serrate (with salicoid, gland-tipped teeth), sometimes lobed or dissected, usually with pinnate, sometimes palmate, venation. Inflorescences are axillary or terminal spikes, thyrse-like panicles, bunches (racemes), fascicles or f lowers solitary, and usually subtended by a persistent bract. The bisexual or rarely unisexual flowers are actinomorphic or zygomorphic and often placed on kneed (bent) pedicels and subtended by two bracteoles. Flowers are sometimes cleistogamous (not opening and self-pollinating). The five sepals are free and persist in fruit, equal or unequal in size and shape. The five free petals are equal in actinomorphic flowers, but in zygomorphic species the lowermost petal is enlarged and often spurred, gibbous or saccate. The five (sometimes three) stamens are free or filaments fused into a tube, in zygomorphic flowers with two nectar glands enclosed by the spurred or saccate petal. Anthers are basifixed and open by lengthwise slits, the connectives often dilated into membranous appendages. The superior ovary is composed of usually three (sometimes two to five) fused carpels forming a single locule, topped with Melicytus crassifolius fruits, Chelsea Physic Garden, London, UK [236]

MALPIGHIALES

EUDICOTS

a single filiform or clavate style and a simple variously shaped stigma. The fruit is a capsule that opens along the carpel suture in three (sometimes two to five) valves, sometimes a berry or nutlet. Distribution: Violaceae occur nearly worldwide, but are particularly diverse in tropical regions. The herbaceous genus Viola is most diverse in temperate and montane regions.

vegatatively characterised by having serrate alternate stipulate leaves with raised venation. Hybanthus and Rinorea are not monophyletic in their traditional treatments and will have to be recircumscribed, either to include other genera or to be divided. Genera and species: Violaceae include 25 genera and c. 980 species: Acentra (1), Agatea (8), Allexis (4), Amphirrhox (1), Anchietea (5), Calyptrion (4), Decorsella (1), Fusispermum (3), Gloeospermum (12), Hekkingia (1), Hybanthopsis (1), Hybanthus (c. 120), Isodendrion (c. 7), Ixchelia (2), Leonia (6), Mayanaea (1, possibly extinct), Melicytus (10), Noisettia (1), Orthion (6), Paypayrola (8), Phyllanoa (1), Rinorea (c. 250), Rinoreocarpus (1), Schweiggeria (1) and Viola (c. 525).

nearly every colour of the rainbow. They are complex hybrids (usually referred to as V. ×wittrockiana) involving several species, including V. altaica, V. lutea and V. tricolor. Due to their tolerance of cold temperatures, they are frequently used in temperate climates for winterbedding. Etymology: Viola is a classical Latin name for a scented flower, probably originating from a pre-Indo-European language. The musical instruments viola and violin are named because they have a shape similar to a violet.

Phylogeny and evolution: Previously, in morphology-based classifications the family was placed among other families with salicoid teeth and parietal placentation, including Salicaceae and Passifloraceae. Molecular results corroborated its placement as sister to Goupiaceae (formerly in Celastraceae), with which they share stamen appendices. Even though they are obviously related to the other malpighialean families, morphologically Goupiaceae and Violaceae stand apart from them. Fusispermum and Leonia are sometimes segregated into their own subfamilies; they are sister to the rest of the family, but because each subfamily consists of only a single genus, we do not see the need for this classification. Some woody species, such as Melicytus, which have berries and actinomorphic f lowers, are often incorrectly identified as belonging to other families, but the family can be

Uses: Sweet violets, Viola odorata, are cultivated for their scented flowers, which are used in perfumes, jellies, syrups and sweets and as flavouring for chocolate. Leaves of some Viola species are sometimes eaten as a pot herb. Flowers of wild pansy, V. tricolor, and garden pansies (V. ×wittrockiana) are sometimes eaten in salads, although in excess they are reputed to have a laxative effect. Many species of Viola are cultivated as ornamentals, some mountain species treasured as unusual rockgarden plants, but most commonly pansies are grown as annual bedding plants, having been bred to include

These evergreen trees accumulate aluminium and have alternate, petiolate leaves that are placed in a plane (distichous) with narrow stipules on both sides of the petiole. The simple blades are palmately veined basally (usually with three or five main veins)

Melicytus chathamicus, Royal Botanic Gardens, Kew, UK [236]

Rinorea scheffleri, Helsinki Botanical Garden, Finland [236]

Hybanthus calycinus, near Perth, Western Australia [236]

237. GOUPIACEAE Kopi family

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MALPIGHIALES

EUDICOTS

Goupia glabra in fruit, Venezuela (VZ) [237]

with the midvein pinnate and margins entire. Inflorescences are axillary umbellike racemes. The bisexual f lowers are actinomorphic and have slender pedicels with triangular, hairy bracts. The five sepals are fused at the base, and the five free petals have a long, subulate apex. The nectary disk is large and cup-shaped, somewhat wavy along the margin. The five stamens are inserted on the edge of the nectary disk that surrounds them. The basifixed anthers are almost sessile, open with lengthwise slits and have an apical thickened, hairy connective. The superior ovary is partly sunk in the nectary disk but not fused with it. It is composed of five locules, each topped with a short free style. The fruit is a hard, globose drupe. Distribution: This is a family of rain forest trees found in Mesoamerica (Guatemala) and northern South America. Phylogeny and evolution: Formerly the genus was included in Celastraceae where it did not fit morphologically. DNA results placed Goupiaceae as sister to Violaceae with which they share the stamen connective appendages and haplostemony. Genera and species: The sole genus in this family is Goupia with three to five species. Etymology: Kopi or goupi, the Ndyuka name for Goupia glabra in French Guiana, has been French-Latinised to form Goupia.

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Goupia glabra, Guyana (LW) [237]

238. PASSIFLORACEAE Passionfruit family

These are perennial and annual herbs, shrubs, small trees and tendrillate climbers. They often have glandular hairs, and some species have a foetid smell (Malesherbia). Leaves are alternate, simple, pinnatifid or palmatifid to palmately lobed or compound, sessile or petiolate and (usually) with leaf-like or scale-like stipules at the base of most leaves and bracts and sometimes with stalked or sessile glands on the petioles, stipules or leaf blades. Blades are highly variable in shape, simple, crenate, toothed or lobed, laterally expanded or variously palmately compound with pinnate, palmate or pedate venation. Tendrils are often present in the leaf axils or terminate inflorescences, and branches are simple, bifid or trifid. Inflorescences are cymes, leafy panicles or racemes, the flowers rarely in fascicles, triads or solitary, sometimes epiphyllous or in heads. The bisexual flowers are usually actinomorphic, rarely zygomorphic, and pedicels often have prophylls. The perianth is usually fused at the

base into a saucer-shaped, tubular, funnelshaped or campanulate floral tube. The five (or three to eight) sepals are free or partially fused and inserted on the floral tube. The five (or three to eight) petals are clawed or not, free or partially united or (rarely) absent. In some genera, a corona is well developed and can be a ridge or composed of threads, fringes, tubercles, glands or scales, and like the petals is inserted on the rim of the floral tube in one to many rows. Nectaries are often present at the base of the floral tube. Male and female parts are usually placed together on a stalk (androgynophore), rarely the ovary sessile (in Turneroideae). The five (sometimes four to eight or numerous) stamens are usually fused to the stalk just below the ovary. Anthers are dorsifixed (or nearly basifixed) and open by longitudinal slits. The superior (rarely inferior) ovary is usually terminal on the androgynophore and composed of three or four fused carpels forming a single locule and three or four free styles at the top. Fruits are capsules or berries surrounded by the persistent perianth, containing seeds with a fleshy or pulpy aril. Distribution: Passifloraceae have a pantropical distribution, extending into the temperate zones in eastern North America, Argentina and southern Brazil, southern China and New Zealand. Malesherbioideae are restricted to Pacific South America, from Peru to Central Chile. Turneroideae are found throughout the American and African tropics.

MALPIGHIALES

EUDICOTS

Phylogeny and evolution: Formerly treated in three separate families and now accepted as subfamilies, Malesherbioideae, Passifloroideae and Turneroideae; these were always thought to be closely related, even before the molecular age. The discovery of Pibiria as sister to Passifloroideae and Turneroideae reinforces this relationship, but weakens the morphological distinction between these subfamilies. Hollrungia and Tetrapathea are included in Passiflora.

Genera and species: Passifloraceae include 28 genera and c. 982 species in three subfamilies: Malesherbioideae – Malesherbia (24); Passifloroideae – Adenia (93), Ancistrothyrsus (2), Androsiphonia (1), Barteria (4), Basananthe (37), Crossostemma (1), Deidamia (5), Dilkea (3), Efulensia (2), Mitostemma (3), Paropsia (12), Paropsiopsis (7), Passiflora (c. 560), Schlechterina (1), Smeathmannia (2) and Viridivia (1); Turneroideae – Adenoa (1), Erblichia (5), Hyalocalyx (1), Loewia (3),

Uses: The most extensively cultivated passion fruit is the maracuja, Passiflora edulis, which is eaten fresh or used for juice. The second most commonly grown passion fruit in the tropics is the sweet granadilla, P. ligularis, which has much sweeter fruit that is lighter in colour. The water lemon

Barteria fistulosa, Avebe River, Monts de Cristal, Gabon (CD) [238]

Malesherbia linearifolia by W. J. Hooker from Curtis’s Botanical Magazine vol. 61, plate 3362, 1834 [238]

Passiflora edulis, planted in a garden on Easter Island [238]

Mathurina (1), Pibiria (1), Piriqueta (44), Stapfiella (6), Streptopetalum (6), Tricliceras (16) and Turnera (c. 140).

Passiflora manicata, Ecuador [238]

Basananthe sandersonii, Kloof Gorge, KwaZulu Natal, South Africa (CD) [238]

Adenia globosa subsp. pseudoglobosa, Kifaru, Tanzania (CD) [238] Turnera ulmifolia, Royal Botanic Gardens, Kew, UK [238]

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MALPIGHIALES (P. laurifolia), sweet calabash (P. maliformis), giant granadilla (P. quadrangularis) and the wild maracuja (P. foetida) are locally grown but rarely found cultivated at a larger scale. Banana passionfruits (P. mollissima and P. tarminiana) are also locally grown but can be invasive. They should not be cultivated outside their native range. Maypop, P. incarnata, is a native of North America and also locally cultivated for its fruits and flowers. Brazilian Passif lora caerulea is the most widely cultivated passionflower in temperate gardens. It has edible fruits but is not worth cultivating for that purpose. It can also be invasive due to its underground runners. Many species of Passiflora, Piriqueta and Turnera are grown as ornamentals, and Adenia is sometimes found in specialist collections. Other genera may be of potential ornamental value. Herbivory and carnivory: Passiflora species are known for their glands that mimic insect eggs to fool Heliconius butterflies, which do not lay eggs if there are already “eggs” present on the leaves. In this manner they may prevent predation by caterpillars. Similarly, the peculiar, pantropical, weedy wild maracuja, Passiflora foetida, has long, glandular hairs on the bracts subtending its white-green flowers. These hairs have been hypothesised to produce digestive enzymes, but this has not been proven. It may just be another case of a sticky floral defence against herbivory. The preferred habitats of Passiflora do not fit those of other carnivorous taxa. Etymology: Passiflora is composed of the Latin words passio, passion, and flos, a flower. The flower parts were thought by Spanish missionaries to represent the passion of Jesus Christ. Hence, the ten petals and sepals represent the ten faithful apostles (excluding Peter and Judas), the radial filaments represent the crown of thorns, the central cup of the flower supposedly represents the Holy Grail, the five anthers represent the five wounds, and the three stigmas represent the three nails. Outside the Christian influence, these plants usually were referred to as clock-flower because the flower reminded them of the face of a clock.

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EUDICOTS

239. LACISTEMATACEAE Cemp-wood family

The superior ovary is composed of three fused carpels forming a single locule topped with a single style and three short stigmas. The fruit is a capsular berry with three valves. Distribution: Lacistemataceae are only found in tropical America.

These shrubs and small trees usually have hairy young branches. Their leaves are simple, alternate and arranged in a plane (distichous) with a short petiole and deciduous stipules. Leaf blades have serrate margins and pinnate venation. Inf lorescences are bracteate, axillary, fascicled spikes, racemes or panicles. The bisexual flowers are zygomorphic and subtended by two bracteoles. The one to six sepals are unequal in size, and petals are absent. The single stamen is on or inside a fleshy, sometimes cupular disk. The filament is usually bifid with each branch bearing a globose anther that opens by a lengthwise slit. Lozania nunkui, Cordillera del Condor, Ecuador (CD) [239]

Lacistema aggregatum, La Selva Biological Station, Heredia, Costa Rica (CD) [239]

Phylogeny and evolution: Although they were usually treated as a tribe of the now defunct Flacourtiaceae, the most recent classification favours the status of the family as sister to Salicaceae, in which most members of former Flacourtiaceae are now placed. The lack of a perianth is shared with Salicaceae s.l. Previous studies placed Lacistemataceae as sister to Ctenolophonaceae. Genera and species: This family includes two genera and 16 species: Lacistema (11) and Lozania (5). Etymology: Lacistema is composed of the Latin words lacerus, torn, and Greek stema, στεμα (stema), a penis, in reference to the deeply split stamens. Lacistema aggregatum in fruit, La Selva Biological Station, Heredia, Costa Rica (CD) [239]

MALPIGHIALES

EUDICOTS

Populus trichocarpa in fruit, Suomenlinna, Helsinki, Finland [240]

Carrierea calycina leaves with salicoid teeth, Royal Salix myrsinites, female, Ruissalo Botanic Gardens, Kew, UK [240] Botanical Garden, Turku, Finland

240. SALICACEAE Willow family

These are bisexual sometimes unisexual shrubs and trees, sometimes almost herbaceous (but woody at the base). Leaves are alternate, rarely subopposite (Salix purpurea) and usually arranged in two ranks (distichous) or spirally. The simple blades often have marginal salicoid teeth (where the vein ends in a dark, translucent, spherical gland) and usually pinnate, rarely palmate venation. Petioles often bear glands, and stipules are usually present (in Azara leaf-like and appearing opposite the leaf), sometimes absent. Inflorescences are racemes, spikes or catkins, sometimes fascicled or racemose cymes, with a bract subtending each flower, sometimes epiphyllous (Mocquerysia, Phyllobotryon). Flowers are usually unisexual, sometimes bisexual, and both occur on the same or different plants. A perianth can be present or absent, sometimes replaced by nectaries or a disk. Sepals when present are three to eight (sometimes to 15) and free or basally fused. Petals are usually absent, sometimes present and then free and as many

[240]

as the sepals. A nectary disk is often present and lobed. The one to numerous stamens have free or basally fused filaments, often fasciculate, and the anthers are dorsifixed and open via lengthwise slits towards the outside of the flower. The superior ovary is composed of one to ten fused carpels, each forming a locule and bearing a style that is free or fused into a single style and two to four, notchedcapitate stigmas. Fruits are berries, drupes or capsules with usually three valves, rarely samaras. Seeds sometimes have an aril or long silky hairs that aid the dispersal by wind (e.g. Populus and Salix). Distribution: This is a cosmopolitan family found from the Arctic to the tropics. They are absent from Antarctic regions, the Sahara and Arabian deserts and most of Australia and New Zealand. Salix arctica can be found at 83ºN latitude and is thus the most northerly distributed vascular plant together with Papaver radicastum (Papaveraceae), which occurs there as well. Phylogeny and evolution: In the past, a relationship with Salicaceae s.s. and part of Flacourtiaceae had been hypothesised based especially on the similar distinctive (salicoid) teeth and phenolic compounds such as salicin. Flacourtiaceae used to be a ‘dumping ground’ for taxa that were difficult to place, but after molecular phylogenetic studies, it was shown that the family was widely polyphyletic. Many genera were moved to other families and orders, including Aphloiaceae, Asteropeiaceae,

Salix repens, male, Buurserzand, the Netherlands [240]

Berberidopsidaceae, Celastraceae, Crypteroniaceae, Dioncophyllaceae, Euphorbiaceae, Gerrardinaceae, Malvaceae, Muntingiaceae, Myrtaceae, Passifloraceae, Putranjivaceae, Theaceae, Thymelaeaceae etc. Some genera remained in Malpighiales but were placed elsewhere, such as the tribe Oncobeae (but not most species of Oncoba), which are closer to Achariaceae, in which these genera that have cyanogenic glycosides and lack the salicoid teeth are now placed. Lacistemataceae were also previously included in Flacourtiaceae, but are now in their own family, sister to Salicaceae, where the vast majority of the former Flacourtiaceae now resides, including the type genus Flacourtia. A tribal classification has been proposed, but this still needs more investigation. It is preliminary because not all clades are well supported. Nevertheless, Casearia and relatives (formerly placed in Samydaceae or Passifloraceae), which lack salicoid teeth are sister to the rest of the family or recognised as the family Samydaceae. Trichostephanus is placed here tentatively. Crown Salicaceae have been dated to 40–63 million years. An Eocene fossil from North America links Salix with florally more conventional members of the family. Genera and species: Salicaceae in their current circumscription include 56 genera and c. 1,220 species: Abatia (9), Aphaerema (1), Azara (10), Banara (33), Bartholomaea (2), Bembicia (5), Bennettiodendron (3), Bivinia (1), Byrsanthus (1), Calantica (8), Carrierea (3), Casearia (180), Dissomeria (2), Dovyalis

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MALPIGHIALES

Abatia parviflora, Ecuador [240]

EUDICOTS

Homalium decurrrens, New Caledonia [240]

(15), Euceraea (2), Flacourtia (c. 15), Hasseltia (3), Hasseltiopsis (1), Hecatostemon (1), Hemiscolopia (1), Homalium (c. 180), Idesia (1), Itoa (2), Laetia (10), Lasiochlamys (13), Ludia (23), Lunania (14), Macrohasseltia (1), Macrothumia (1), Mocquerysia (2), Neopringlea (3), Neoptychocarpus (2), Neosprucea (5), Olmediella (1, possibly extinct in nature), Oncoba (4), Ophiobotrys (1), Osmelia (4), Phyllobotryon (5), Pineda (1), Pleuranthodendron (1), Poliothyrsis (1), Populus (35), Prockia (2), Pseudoscolopia (1), Pseudosmelia (1), Ryania (8), Salix (c. 450), Samyda (9), Scolopia (37), Scyphostegia (1), Tetrathylacium (2), Tisonia (14), Trichostephanus (2), Trimeria (2), Xylosma (85) and Zuelania (1). Uses: Kei apple (Dovyalis caffra) has edible fruits that are used in jellies and marmalades and frequently planted as hedging throughout the tropics. In Sri Lanka the edible fruits of D. hebecarpa are added to spirits to make a sherry substitute. Flacourtia species also have edible fruits, often used for jellies, including ramontchi (F. indica), batako plum (F. inermis), Indian plum (F. jangomas) and coffee plum (F. rukam), many of which are unknown in the wild (cultigens). The pulp of Oncoba spinosa fruit is also edible. The inner bark of Canadian aspen (Populus tremuloides) is edible and was eaten by native American tribes. The bark of white willow (Salix alba) has long been used medicinally as a painkiller in Europe and is a source of salicin, which 330

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Samyda dodecandra, Fairchild Tropical Botanical Garden, Florida, USA [240]

becomes salicylic acid in the body. This inspired the synthesis of aspirin, but it was first purified from Filipendula ulmaria (Rosaceae), and is now marketed worldwide. The leaves of Abatia rugosa and Pineda incana are the source of a black dye in Peru. Several species of willow (Salix) can be coppiced, and these pliable branches are frequently used to make baskets, fish traps, rugs and even hot-airballoon baskets etc. Many species produce valuable wood, especially Bivinia, Casearia, Homalium, Populus, Salix and Scolopia. Maracaibo boxwood (Casearia praecox) replaces box wood (Buxus sempervirens, Buxaceae) for commercial use in veneers and carving etc. Several genera and species are popular ornamentals, especially Azara, Dovyalis caffra, Idesia polycarpa, Oncoba spinosa, Poliothyrsis sinensis, Populus and Salix. Weeping willows (Salix babylonica complex, S. ×sepulcralis) are often planted along water, such as ponds and streams, as a quintessential element of the English countryside. Corkscrew willow (S. matsudana ‘Tortuosa’) is cultivated for its ornamental stems that are used in decoration and the cut-flower industry and are popular for Easter decorations in the Netherlands. There are numerous cultivars and hybrids of Populus and Salix. Populus euphratica occurs from North Africa to western China, but is only known from a single European site near Alicante, Spain. The Spanish population is clonal, consisting of only female plants. It was probably introduced by the Moors.

Oncoba spinosa, Botanical Garden, National Museums of Kenya, Nairobi [240]

Record: A clone of quaking aspen (Populus tremuloides) in Utah called ‘Pando’ is estimated to be c. 80,000 years old, the individual trunks not living that long, but it resprouts from an ancient underground rootstock. This may be the oldest living plant on Earth. Etymology: Salix is the classical Latin name for willow, from the Proto-Indo-European word salik.

241. PERACEAE Lightning-bush family

Peraceae are unisexual, sometimes bisexual trees, shrubs and perennial herbs without latex. Hairs are simple, compass-like (malpighioid), stellate or scale-like. Leaves are simple, alternate or rarely opposite, petiolate with or without stipules. Leaf blades are entire with pinnate venation. Inf lorescences are axillary glomerules in Pera surrounded by involucral bracts. Flowers are unisexual and actinomorphic. Sepals are absent or two to six, petals are

MALPIGHIALES

EUDICOTS

Clutia pulchella, male flowers, Hortus botanicus, Leiden, the Netherlands [241]

Clutia robusta, female flowers, Kenya [241]

Clutia robusta, male flowers, Kenya [241]

often absent, sometimes present and free. A disk is present (absent in Pera), usually glandular or annular. One to 20 stamens are free, partially fused or completely united into a column with anthers that are dorsifixed and open by lengthwise slits. Male flowers sometimes have a sterile pistillode, and in Pera reduced pistillate flowers may surround fertile male flowers. The usually tri- (rarely tetra-) locular ovary is superior and topped by free, usually bifid syles, or these fused into a thick peltate stigmatic structure (Pera). The fruit is a dehiscent, rarely indehiscent, capsule with a persistent columella and fragile membranaceous septa. Seeds often have a caruncle, consistent with dispersal by ants. They have a peculiar type of seed coat (with a tracheoidal exotegmen), distinguishing this family from Euphorbiaceae.

Distribution: Peraceae can be found in tropical America, Africa, Madagascar, Yemen, Sri Lanka, Southeast Asia and Borneo.

Etymology: Pera is derived from Ancient Greek pera (pera), a pouch or bag, in reference to the swollen fruits.

Phylogeny and evolution: Placement of Rafflesiaceae in Euphorbiaceae s.s. resulted in the acceptance of former Euphorbiaceae subfamily Peroideae as a separate family despite their numerous shared morphological characters.

242. RAFFLESIACEAE

Sapria himalayana, Thailand (MN) [242]

Corpse-flower family

Genera and species: Peraceae comprise five genera and c. 127 species: Chaetocarpus (13), Clutia (c. 75), Pera (c. 35), Pogonophora (1) and Trigonopleura (3). Uses: The young leaves of bukir (Chaetocarpus castanocarpus) are edible. Timber of some tree species is used as firewood.

Rafflesiaceae are difficult to locate when they are not flowering because most of the plant grows endophytically (inside the stems or

Rafflesia arnoldii, Tab. 15 accompanying the article describing the genus and species by Robert Brown, Trans. Linn. Soc. London 13 (1). 1821 [242]

Rafflesia keithii, Kinabalu National Park, Sabah, Malaysia (CD) [242]

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MALPIGHIALES roots of other plants). Only the inflorescence appears, rather randomly, from a root or stem. They are unisexual or bisexual parasites with filamentous tissues that resemble a fungal mycelium inside the host roots and stems. Stems and leaves are absent, but some scalelike inflorescence bracts subtend the flowers. Inf lorescences comprise a single f lower in a basal cup that bursts out of the host stem. The unisexual (sometimes bisexual in Rhizanthes) flowers are actinomorphic, and some species can have large flowers, some the largest known. Tepals are fused to each other and the ovary, forming a cup-shaped structure with five (Rafflesia), ten (Sapria) or 16 (Rhizanthes) lobes. Lobes either form a central opening (diaphragm) or have long appendages (Rhizanthes). The inside surfaces of the flower often have branched or clubshaped hairs (ramentae), which may function as nectaries or scent organs (osmophores). Five to 50 stamens are sessile and emerge on the lower side of the central disk-shaped column of the flower, the anthers with one or two pores. The disk-like column is expanded over the anthers and often forms many hornlike structures, where in female flowers, stigmas are situated. The inferior ovary is composed of four to eight fused carpels with a columnar style that is expanded at the tip into a disk topped by the stigmas. The fruit is a fleshy berry that sometimes opens irregularly exposing numerous seeds.

EUDICOTS

Genera and species: Rafflesiaceae include three genera and c. 25 species: Rafflesia (c. 20), Rhizanthes (2) and Sapria (2). Parasitism and pollination: Rafflesiaceae parasitise Vitaceae in Southeast Asia. Rafflesia lives inside Tetrastigma and other genera, whereas the other two genera are exclusively found on Tetrastigma. Their tissues are reduced and resemble fungi inside the vessels of the host. Previously suggested relationships with Vitaceae based on DNA sequence studies are most likely due to horizontal transfer of plastids between host and parasite. Many Raff lesiaceae have thermogenic f lowers similar in function to some Araceae, producing a pungent smell of rotting flesh and attracting carrion flies for pollination. The temperature in the flowers can be 7–9ºC above ambient. It has been suggested that the pollination syndrome is predominantly one of brood-site deception of the flies attracted by the smell, heat and appearance of the flowers. Etymology: Rafflesia was named for Sir Thomas Stamford Raff les (1781–1826), founder of the city of Singapore and leader of the expedition to Indonesia where it was first discovered. A colour illustration was made of the huge flower by its discoverer Joseph Arnold (1782–1818) and Lady Raffles, who accompanied the expedition.

Record: Rafflesia arnoldii from Sumatra holds the record for the largest flower, which can grow up to 1 metre in diameter and weigh up to 10 kg, but even the smallest species have relatively large flowers, e.g. flowers of R. manillana measure 20 cm in diameter. Flower size in Rafflesia has been hypothesised to have increased dramatically during the evolutionary history of the genus, perhaps related to parasitism and loss of leaves and roots, allowing them to invest more energy in flowers.

243. EUPHORBIACEAE Spurge family

This family comprises unisexual, sometimes bisexual, annual and perennial herbs, shrubs and trees, rarely vines, often succulent, sometimes cactus-like, frequently with milky (Euphorbioideae) or coloured latex, sometimes with clear sap. Hairs are simple, branched, compass-like (malpighioid),

Distribution: Rafflesiaceae are found in Southeast Asia, from Assam to western Malesia. Phylogeny and evolution: The family has been variously circumscribed in the past, often including other endoparasitic plants such as Apodanthaceae (Cucurbitales), Mitrastemonaceae (Ericales) and Cytinaceae (Malvales), all now placed in different orders due to our better understanding of relationships resulting from DNA analysis. Rafflesiaceae are placed in Malpighiales, sister to Euphorbiaceae (minus the former Peroideae, now an independent family), and are now restricted to three genera that are morphologically similar and genetically closely related.

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Hippomane mancinella, Galápagos Islands [243]

Excoecaria cochinchinensis, Royal Botanic Gardens, Kew, UK [243]

MALPIGHIALES

EUDICOTS

Euphorbia tithymaloides subsp. padifolia, Guadeloupe [243]

Euphorbia prostrata, Réunion [243]

Euphorbia segetalis, Barbentane, Provence, France [243]

Euphorbia serendipita, Botanical Garden, National Museums of Kenya, Nairobi [243]

Euphorbia pulcherrima, Mexico [243]

Euphorbia cotinifolia subsp. cotinoides, planted near Voi, Kenya [243]

stellate or scale-like. Leaves are simple, sometimes trifoliate or palmately compound, alternate, opposite or sometimes whorled, petiolate or sessile, with or without stipules, the stipules often bearing glands. Leaf blades are entire, toothed or palmately lobed and have pinnate or palmate venation, often bearing glands on blades or petioles. Inf lorescences are axillary or terminal panicles, racemes, spikes, glomerules or thyrses, often condensed and surrounded by involucral bracts that are frequently fused, condensed into a cup-shaped structure resembling a flower (cyathium, in Euphorbia and related genera), rarely with f lowers solitary. In Euphorbioideae, bracts often have two glands at their base. Flowers are unisexual and actinomorphic or (frequently) the perianth absent. Sepals are absent or

two to six (rarely up to 120), and petals are often absent, sometimes present, free and then usually four or five. A disk is present or not, usually glandular and segmented or annular. The one to numerous stamens have free or fused filaments and anthers that are basifixed to dorsifixed and open by lengthwise slits. Male flowers sometimes have a sterile pistillode. The usually trilocular (rarely bi- or tetralocular) ovary is superior and topped with free, usually forked or dissected, sometimes simple; styles fused into one (some Euphorbioideae), sometimes free (stylodia). The fruit is a dehiscent, rarely indehiscent, sometimes explosive capsule with a membrane-like septum and persistent columella, rarely a berry or drupe. Seeds often have a caruncle, enabling distribution by ants.

Distribution: This is a cosmopolitan family apart from cold regions in the Arctic and Antarctic. Phylogeny and evolution: Euphorbiaceae have been variously circumscribed in the past. Previously, Euphorbiaceae included a number of now segregate families, such as Peraceae, Phyllanthaceae, Picrodendraceae and Putranjivaceae. Due to placement of Linaceae and Ixonanthaceae, former subfamily Phyllanthoideae were accepted at the family level, excluding tribe Drypeteae, which are closer to Lophopyxidaceae and are therefore now treated as Putranjivaceae. Placement of Rafflesiaceae as sister to Euphorbiaceae resulted in the acceptance of Peraceae. Currently four subfamilies are recognised, with Cheilosoideae being sister to the rest. Plants of the World

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Dalechampia spathulata, Helsinki Botanical Garden, Finland [243]

Acalypha indica, Kenya [243]

Macaranga griffithiana (WA) [243]

Monotaxis grandiflora, Mt Benia, Western Australia [243]

The family is an estimated 90–102 million years old, with Acalyphoideae having diversified c. 70 million years ago and Euphorbioideae radiating c. 50 million years ago. A 23 millionyear-old fossil from New Zealand can be attributed to Mallotus or Macaranga. Genera and species: In their current circumscription, Euphorbiaceae include c. 210 genera and c. 6,252 species in four subfamilies: Cheilosoideae (two genera) – Cheilosa (1) and Neoscortechinia (6); Acalyphoideae (97 genera) – Acalypha (447), Acidoton (6), Adelia (10), Adenophaedra (3), Adriana (2), Afrotrewia (1), Agrostistachys (6), Alchornea

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(52), Alchorneopsis (2), Amperea (8), Amyrea (11), Angostylis (2), Aparisthmium (1), Argomuellera (12), Argythamnia (23), Astrococcus (1), Aubletiana (2), Avellanita (1), Bernardia (74), Bia (5), Blumeodendron (5), Bocquillonia (14), Botryophora (1), Caperonia (34), Caryodendron (4), Cephalocroton (7), Cephalomappa (6), Chiropetalum (23), Chondrostylis (2), Chrozophora (9), Cladogynos (1), Claoxylon (117), Cleidiocarpon (3), Cleidion (32), Cnesmone (11), Conceveiba (14), Crotonogynopsis (2), Cyttaranthus (1), Dalechampia (121), Discoclaoxylon (4), Discocleidion (1), Discoglypremna (1), Ditaxis (50), Doryxylon (1), Droceloncia (1),

Ricinus communis naturalised on Easter Island [243]

Mallotus japonicus, Mt Tianmu, Anhui, China [243]

Dysopsis (3), Enriquebeltrania (2), Epiprinus (6), Erismanthus (2), Erythrococca (41), Garciadelia (4), Haematostemon (2), Hancea (18), Homonoia (3), Koilodepas (12), Lasiococca (5), Lasiocroton (7), Leidesia (1), Leucocroton (26), Lobanilia (7), Macaranga (308), Mallotus (122), Mareya (4), Mareyopsis (2), Megistostigma (5), Melanolepis (2), Mercurialis (10), Micrococca (12), Monotaxis (8), Moultonianthus (1), Necepsia (3), Orfilea (4), Pachystylidium (1), Paranecepsia (1), Philyra (1), Platygyna (7), Plukenetia (20), Podadenia (1), Pseudagrostistachys (2), Ptychopyxis (10), Pycnocoma (18), Ricinus (1), Rockinghamia (2), Romanoa (1), Sampantaea (1),

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Seidelia (2), Spathiostemon (3), Speranskia (3), Sphaerostylis (2), Sphyranthera (2), Sumbaviopsis (1), Syndyophyllum (2), Tapoides (1), Thyrsanthera (1), Tragia (152), Tragiella (4) and Wetria (2); Crotonoideae (66 genera) – Acidocroton (13), Adenocline (3), Aleurites (2), Alphandia (3), Annesijoa (1), Astraea (8), Baliospermum (5), Baloghia (15), Benoistia (3), Bertya (28), Beyeria (24), Blachia (10), Borneodendron (1), Brasiliocroton (1), Cavacoa (3), Chlamydojatropha (1), Cladogelonium (1), Cnidoscolus (92), Cocconerion (2), Codiaeum (17), Croton (c. 1,185), Crotonogyne (16), Cunuria (6), Cyrtogonone (1), Deutzianthus (2), Dimorphocalyx (17), Ditta (2), Dodecastigma (4), Elateriospermum (1), Endospermum (11), Fontainia (9), Garcia (2), Givotia (4), Glycydendron (2), Grossera (8), Hevea (9), Hylandia (1), Jatropha (173), Joannesia (2), Karima (1), Klaineanthus (1), Leeuwenbergia (2), Manniophyton (1), Micrandra (6), Micrandropsis (1), Manihot (109), Mildbraedia (3), Myricanthe (1), Neoboutonia (3), Oligoceras (1), Omphalea (21), Ostodes (2), Pantadenia (3), Paracroton (4), Pausandra (8), Radcliffea (1), Ricinocarpos (28), Ricinodendron (2), Tetrorchidium (23), Sagotia (2), Sandwithia (2), Shonia (4), Strophioblachia (1), Suregada (31), Tannodia (9), Trigonostemon (84), Vaupesia (1) and Vernicia (4); Euphorbioideae (41 genera) – Actinostemon (19), Adenopeltis (1), Algernonia (12), Anomostachys (1), Anthostema (3), Balakata (2), Bonania (7), Calycopeplus (5), Colliguaja (5), Dalembertia (4), Dichostemma (2), Ditrysinia (1), Euphorbia (1,933), Excoecaria (41), Falconeria (1), Grimmeodendron (2), Gymnanthes (28), Hamilcoa (1), Hippomane (3), Homalanthus (23), Hura (2), Mabea (39), Maprounea (5), Microstachys (18), Nealchornea (2), Neoguillauminia (1), Neoshirakia (1), Ophthalmoblapton (4), Pachystroma (1), Pimelodendron (4), Plagiostyles (1), Pleradenophora (3), Sapium (23), Sclerocroton (6), Shirakiopsis (6), Sebastiania (72), Senefeldera (6), Senefelderopsis (5), Spegazziniophytum (1), Stillingia (29) and Triadica (3). Uses: The family has many uses, including food, oil, rubber, timber and ornamentals.

The most important food crop is the root of Manihot esculenta, which is variously called cassava, yuca, manioc or tapioca root and is the third most important source of starch for human consumption in the tropics after rice and maize. Cassava was first domesticated c. 10,000 years ago in central Brazil, where the wild ancestor M. esculenta subsp. flabellifolia still occurs. By the time of the Spanish conquest of tropical America, it was the staple food of native peoples in the Caribbean and Central and South America. The roots are toxic and need to be properly prepared to detoxify them before consumption. The purified starch of cassava is called tapioca, which has a great number of culinary uses. Tapioca starch is also distilled into alcoholic beverages such as cauim (kauim), chichi de yuca, masato, nihamanchi, parakari, sakura and tiquira, to name a few. Rich in protein, the leaves can also be eaten after cooking. China is investigating the use of cassava as a biofuel, and the starch is used in commercial laundry products. Consumption of cassava is dependent on people (mostly women) being able to devote long periods of time to its preparation, and as societies become more developed other less time-consuming staples are adopted. From the Peruvian Amazon comes sacha inchi (Plukenetia volubilis), a vine with edible seeds cultivated by indigenous people for centuries. The seeds are high in protein and omega-3 fatty acids, which resulted in its promotion as a health food. Its oil is now internationally recognised for its excellent flavour and health benefits (a good alternative to peanuts), and production at a larger scale has been initiated with the aid of humanitarian fair trade organisations to benefit the local Amazonian peoples. The Pará rubber tree, Hevea brasiliensis, was the most important commercial crop in the 19th century, first harvested from the wild in Amazonia and providing important financial development for the region. Later, when seeds were smuggled out of Brazil to Malaya by famous botanical explorer Richard Spruce, large plantations were set up there, and these are still the most important source of commercial non-synthetic rubber, used especially for aircraft tyres. Cultivation in plantations is

only possible outside its native range because no natural pests are present there. Acalypha bipartita is occasionally eaten in Africa, and its stems can be used for basketry. Azafancillo (Ditaxis heterantha) is used as a saffron substitute in Mexico. The candlenut tree or kemiri nut (Aleurites moluccana) has oil-rich seeds used as a condiment or spice and to make a sauce in Indonesian and Malaysian cuisine. It has been used as an alternative to castor oil, paraffin to make candles, an industrial lubricant and a component of varnishes, paint and soap. The seeds are mildly toxic when eaten raw, but this disappears after cooking. Castor oil is pressed from the seeds of Ricinus communis. This slowly burning oil has been used since ancient times in lamps, and seeds have been found in 4,000-year-old Egyptian tombs. It is also used for various cosmetic and medicinal purposes, although it contains ricin, which is highly toxic and normally removed during the processing of seeds. Castor oil now finds use as biofuel in Brazil, and in its altered form it finds its way into many food and cosmetic products, even as a substitute for cocoa butter in cheap chocolate. Inchi tree or tacay nut oil (Caryodendron orinocense) is superior to palm oil and has various culinary uses, especially in Latin America. Purging nut (Jatropha curcas), formerly used medicinally, has oil-rich seeds that can be processed into high-quality biodiesel, combustible in diesel and jet engines. Its latex inhibits the watermelon mosaic virus and can thus be used for biological control of this disease in commercial melon cultivation. Bomahnut (Pycnocoma macrophylla) produces an inedible oil used in tanning. Minor commercial waxes were harvested from various Euphorbia and Sapium species, especially candelilla (E. antisyphilitica), although these are now largely replaced by cheaper petroleum-based waxes. The pencil tree, Euphorbia tirucalli, produces a flammable latex, and they are sometimes called gasoline trees. Commercially, cultivation of this has never taken off. The plant is widely cultivated but is also linked to

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MALPIGHIALES

Hevea brasiliensis, being tapped in a Malaysian plantation [243]

Jatropha multifida, Ecuador [243]

Cnidoscolus souzae, Yucatan, Mexico [243]

Croton mauritianus, Réunion [243]

skin conditions in areas where it is frequently planted. The sandbox tree (Hura crepitans) is poisonous, and its sap is sometimes used to poison fish and coat “poison” darts. The unripe fruits were formerly used to make decorative pen sandboxes to dry ink and flatten the paper before writing. Chinese tallow tree (Triadica sebifera; formerly known as Sapium sebiferum) has a waxy aril on the seeds that is used to make soap. It also is a major honey plant for beekeepers. It has become a problematic invasive in forests and grasslands in the USA. The kamala tree (Mallotus philippensis, M. discolor) yields a yellow dye from its hairy seed capsules. A red dye was formerly extracted from Jatropha podagrica. A red gum is obtained from Macaranga indica (called macaranga gum), which is used to take impressions of 336

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coins, medallions and jewellery etc. Despite many species being highly poisonous, several species are commonly grown as garden or house plant ornamentals, especially members of Euphorbia in its many incarnations, but other frequently grown plants are cat’s tail (Acalypha hispida, A. hispaniolae), Australian holly (Alchornea illicifolia), garden croton (Codiaeum variegatum), butterf ly vine (Dalechampia aristolochiifolia), blindness tree (Excoecaria cochinchinensis), Buddha belly plant (Jatropha podagrica), spicy jatropha (J. integerrima) etc. Poinsettia (Euphorbia pulcherrima) is a ornamental especially popular at Christmas time. A 16th century legend tells that a child, too poor to provide a gift for Jesus at Christmas, collected wild flowers, and these sprouted into poinsettias on the altar, resulting in

a religious association and the millions of poinsettia plants now grown worldwide. Several succulent euphorbias are commonly grown as ornamentals and also often used as living fences, especially E. neriifolia, E. trigona and E. tirucalli. In addition, a good number of Euphorbiaceae become large trees valued for their timber. Many species have unique alkaloids and are used locally for medicinal purposes. Macaranga and Mallotus are indicators for disturbance in tropical Asia, similar to Cecropia (Urticaceae) in the Neotropics. Etymology: Euphorbia is the classical name for a spurge, named for Euphorbus, physician to King Juba II of Numidia (50 BC–23 AD, now Algeria). The physician’s name in turn is Greek (ευφορβος) for ‘good pasture’.

MALPIGHIALES

EUDICOTS

there are carpels, which can be free or fused at their base and of different lengths on different plants (heterostylous). Stigmas are capitate to filiform. Fruits are septicidal capsules, sometimes a drupe, the seeds sometimes with arils.

244. LINACEAE Flax family

Distribution: Linaceae are pantropical, with the herbaceous Linoideae extending into the temperate regions of both hemispheres and in steppe vegetation of the arid zones. This is a family of annual and perennial herbs, shrubs, trees and woody vines, sometimes with climbing hooks. Leaves are alternate, rarely opposite, sessile or petiolate, with stipules that are sometimes reduced or occasionally toothed or incised. Blades are simple, with an entire margin and pinnate venation. Inflorescences are terminal or axillary thyrses or bunches (cymes or racemes), rarely flowers solitary. The bisexual flowers are actinomorphic. The (four or) five sepals are free or fused at the base and persistent in fruit. Petals are as many as sepals and are often clawed, usually free or nearly so. Stamens are usually twice as many as petals, sometimes four, five or 15, in a single whorl and often some stamens reduced to staminodes. Filaments are basally fused into a tube, surrounded often by two to five nectary glands or a disk outside the tube. Anthers are dorsifixed and open by longitudinal slits. The superior ovary is composed of two to five (rarely eight) carpels forming twice as many locules. There are as many filiform styles as Linum dolomiticum, Helsinki Botanical Garden, Finland [244]

Phylogeny and evolution: Linaceae were always associated with Malpighiales, although they have in the past been expanded to include a number of families that are now recognised separately. Molecular phylogenetic analysis place Linaceae as sister to Ixonanthaceae, with which they share many characters. The family is readily divided into two subfamilies, with different habits, fruit types and stamen number. Linoideae are poorly resolved, and Linum may have to include genera like Rhadiola, Hesperolinon, Cliococca and Sclerolinon, pending further studies. Fossil pollen is known from Miocene deposits in Spain and Pliocene in Germany. Subfossil linseed is frequently found in archaeological sites. Genera and species: Linaceae have 13 genera and c. 255 species in two subfamilies: Hugonioideae – Hebepetalum (3), Hugonia (c. 40), Indorouchera (2), Philbornea (1) and Roucheria (7); Linoideae – Anisadenia (2),

Linum austriacum, Transylvania, Romania [244]

Cliococca (1), Hesperolinon (13), Linum (c. 180), Radiola (1), Reinwardtia (1), Sclerolinon (1) and Tirpitzia (3). Uses: For more than 7,000 years, flax (Linum usitatissimum) has been cultivated in Europe, the Middle East and Anatolia for its fibre (flax, linen), seed (linseed) and oil (linseed oil). Linen is flexible, strong and durable, and absorbs water, making it suitable for table cloths, napkins and towels. Flax was traditionally grown in Northern Ireland, and Irish linen table cloths and napkins are often treated as family heirlooms. Linen is also popular for clothes and was the main plant fibre used in Europe before cotton was imported, but in modern clothes it is often mixed with other fibres to prevent wrinkling. Linseed oil has many applications, as it is originally liquid, but dries sturdily, is water repellent and does not become brittle. It has thus been used as a base (mixed with pigments) for oil paint, in most classical art works by famous painters like Chagall, Da Vinci, Kandinsky, Monet, Rembrandt, Renoir, Rubens, Titian, Van Eyck, Van Gogh and Vermeer, among others. It is also used for putty to seal windows in place, as a wood finish, for guilding (to fix sheet gold) and as a natural food supplement, but it has a short shelf-life. Mixed with wood dust and cork particles, linseed oil can be manufactured into a durable flooring called linoleum, commonly used between 1870 and 1970, particularly in industrial settings. Additionally some species

Indorouchera griffithiana, Singapore (WA) [244]

Hugonia micans, Gabon (AP) [244]

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MALPIGHIALES of Linum are occasionally grown as rock garden or bedding ornamentals. Etymology: Linum is the classical Latin name for flax. It is also the origin of the words linnen, linseed and line. It originated from Proto-Indo-European lin-, a cord, a line or flax. The epithet usitatissimum refers to the many uses of the plant.

245. IXONANTHACEAE Twentymen-tree family

Trees and shrubs make up this family. Their simple leaves are alternate and spirally arranged and have small lateral or interpetiolar stipules at the base of the petiole.

EUDICOTS

Blades have an entire or glandular serrate margin and pinnate-reticulate venation. Inflorescences are lateral corymbs composed of bisexual, (nearly) actinomorphic flowers. The five sepals are free or fused at their base, the five petals are free. The five, ten or 20 stamens are found in one, two or three whorls, which have filaments that widen at the base or fuse with the cup-shaped nectary disk. The superior ovary is composed of five fused carpels, topped with a single, slender style and a capitate or disc-shaped stigma. Fruits are septicidal (sometimes secondarily loculicidal) capsules often with a persistent columella. Seeds are usually winged at the base or have an aril.

molecular analysis has placed Ixonanthaceae as sister to Linaceae, whereas Irvingiaceae are found elsewhere in Malpighiales, close to Pandaceae. Fossil pollen of Ixonanthaceae is known from the Palaeocene of India, along with other African elements, and it has been suggested that the family ‘drifted’ along with continental India and subsequently dispersed and diversified in tropical East Asia. Allantospermum fits morphologically and molecularly better in Irvingiaceae, where it is now placed. The genus is distinct from both families in wood anatomical characters. Phyllocosmus is merged with Ochthocosmus because the genera are indistinguishable.

Distribution: This family has a pantropical distribution, extending into the subtropical Himalayas.

Genera and species: This family is composed of three genera and c. 20 species: Cyrillopsis (2), Ixonanthes (3) and Ochthocosmus (c. 16).

Phylogeny and evolution: Ixonanthaceae are similar to Irvingiaceae in several characters, and together they were often placed in or close to Linaceae. These three families share many characters, and indeed

Etymology: Ixonanthes is composed of the Greek ήχων (ixon), a sound or tune, and άνθη (anthe), a flower. Ixonanthes icosandra is called the twenty-men tree in Singapore because the flower has twenty stamens.

Ixonanthes icosandra in fruit, Singapore (WA) [245]

Ixonanthes reticulata, Bako National Park, Sarawak, Malaysia (CD) [245]

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Ixonanthes icosandra, Singapore (WA) [245]

Ochthocosmus congolensis, Ogooue River, Gabon (CD) [245]

MALPIGHIALES

EUDICOTS

Picrodendron macrocarpum, Fairchild Tropical Botanical Garden, Florida (CD) [246]

Tetracoccus dioicus, Regional Parks Botanic Garden, Berkeley, California, USA (SS) [246]

246. PICRODENDRACEAE Hollyspurge family

This family includes unisexual and bisexual trees and shrubs. They have alternate, opposite or whorled, simple or palmately compound leaves that are petiolate. Stipules are absent or fall off early. Leaf blades have entire or toothed margins, and venation is pinnate. Inflorescences are axillary racemes, glomerules, heads or panicles with bracts that lack glands. Flowers are unisexual and actinomorphic, with usually four to eight (sometimes three to 13) free sepals and no petals. A disk is annular to lobed or absent. Male flowers have two to 30 stamens with free or fused filaments, and the basifixed to dorsifixed anthers open

Tetracoccus hallii, Palm Springs, California, USA [246]

by slits. A pistil may also be present. Female flowers have a superior ovary with two to four (sometimes five) locules, each topped with a style that can be partially fused. The fruit is a loculicidal capsule, sometimes a drupe, with carunculate seeds. Distribution: This is a pantropical family with a somewhat patchy distribution in the Neotropics. It extends from southern California and the Caribbean to Amazonia and a few isolated areas further south. In Africa they are restricted to equatorial regions and Madagascar, and in Asia they occur in Malesia, extending to New Guinea, Fiji, New Caledonia, eastern Australia and New Zealand. Phylogeny and evolution: These used to be treated as Euphorbiaceae subfamily Oldfieldioideae, but Euphorbiaceae were found to be polyphyletic in their traditional sense. Thus this subfamily was recognised as a family. They are most closely related to Phyllanthaceae, another family previously considered a subfamily of Euphorbiaceae. Paradrypetes, sometimes associated with this subfamily is

Austrobuxus montis-do, Mont Do, New Caledonia (CD) [246]

now placed in Rhizophoraceae on the basis of molecular results, even though the structure of its pollen makes it a much better fit here. Genera and species: This family has 25 genera and 96 species: Androstachys (1), Aristogeitonia (7), Austrobuxus (22), Choriceras (2), Dissiliaria (6), Hyaenanche (1), Kairothamnus (1), Longetia (1), Micrantheum (4), Mischodon (1), Neoroepera (2), Oldfieldia (4), Parodiodendron (1), Petalostigma (5), Picrodendron (1), Piranhea (4), Podocalyx (1), Pseudanthus (9), Sankowskya (1), Scagea (2), Stachyandra (4), Stachystemon (9), Tetracoccus (4), Voatamalo (2) and Whyanbeelia (1). Uses: Lebombo ironwood (Androstachys johnsonii) is hard and durable, good for floors, pillars and beams, especially used in coastal dwellings in Mozambique. It is traded as mecrussé, although it is of limited supply. Tammana (Mischodon zeylanicus) is used for construction in Sri Lanka. Etymology: Picrodendron is composed of the Greek pikra, bitter, and dendron, a tree. Plants of the World

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247. PHYLLANTHACEAE Leafflower family

These are unisexual and bisexual trees, shrubs and herbs, rarely climbers, succulents and aquatics without latex. Leaves are alternate (rarely opposite), spirally arranged or in a plane (distichous), the petioles without glands and with free or interpetiolar stipules that are usually entire and scale-like, sometimes leaf-like, rarely spinose. Blades are normally simple, margins entire (trifoliate and toothed in Bischofia), and venation is pinnate, rarely somewhat palmate at base, usually without leaf glands or domatia. Stems are sometimes broad and leaf-like (phylloclades), and then the leaves absent or reduced. Inflorescences are usually axillary

EUDICOTS

raceme- or spike-like thyrses, sometimes in glomerules or the flowers solitary; pseudanthia occur in Uapaca. Inflorescence bracts lack glands, and flowers are usually unisexual and actinomorphic. The usually three to six sepals are more or less free, rarely fused. Petals are sometimes present and then (two to) four to six or absent. A nectary disk is often present, and in male flowers these form distinct glands. Male flowers have three to ten (sometimes up to 19) stamens with free or variously fused filaments. Anthers are basifixed to dorsifixed and open with horizontal or lengthwise slits, rarely pores. A pistil is sometimes present in the middle but frequently absent. Female flowers usually lack staminodes and have a superior ovary composed of two to five (rarely one to 15) locules, each tipped with a usually free, once forked or unlobed style or styles fused, rarely absent. The fruit is usually an explosively dehiscent capsule with a persistent columella, sometimes a berry or drupe. Distribution: The family occurs in warm temperate and tropical regions around the world.

Poranthera microphylla, Perth, Western Australia [247]

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Phylogeny and evolution: Phyllanthaceae were formerly treated as a subfamily in Euphorbiaceae, but relative to the position of Linaceae, some subfamilies of Euphorbiaceae needed to be treated at the family level. They are closely related to Picrodendraceae, another former subfamily of Euphorbiaceae, together sister to Linaceae plus Ixonanthaceae. The family is estimated to have diversified c. 81 million years ago. Lachnostylis is a relic in the Cape region and is dated to c. 90 million years old. Molecular phylogenetic studies have shown that Phyllanthus should be expanded to include Reverchonia, Glochidion, Sauropus and Breynia. Genera and species: Phyllanthaceae include 57 genera and c. 2,050 species, in two subfamilies; Antidesmatoideae – Antidesma (101), Apodiscus (1), Aporosa (82), Ashtonia (2), Baccaurea (51), Bischofia (2), Celianella (1), Didymocistus (1), Distichirhops (3), Hieronyma (21), Hymenocardia (6), Jablonskia (1), Leptonema (2), Maesobotrya (18), Martretia (1), Nothobaccaurea (2), Protomegabaria (3), Richeria (5), Spondianthus (1), Thecacoris

Baccaurea bracteata, in fruit, Singapore (WA) [247]

MALPIGHIALES

EUDICOTS

Phyllanthus androgynus, Fairchild Tropical Botanical Garden, Florida, USA [247]

Phyllanthus mimosoides, Guadeloupe [247]

(16) and Uapaca (27); Phyllanthoideae – Actephila (30), Amanoa (16), Andrachne (22), Astrocasia (6), Bridelia (50), Chascotheca (2), Chonocentrum (1), Chorisandrachne (1), Cleistanthus (134), Croizatia (5), Dicoelia (2), Discocarpus (4), Flueggea (16), Gonatogyne (1), Heterosavia (4), Heywoodia (1), Keayodendron (1), Lachnostylis (2), Leptopus (10), Lingelsheimia (6), Margaritaria (13), Meineckia (30), Notoleptopus (1), Pentabrachion (1), Phyllanthopsis (2), Phyllanthus (c. 1,302),

Phyllanthus calycinus, male flowers, Western Australia [247]

Phyllanthus peltatus, New Caledonia [247]

Plagiocladus (1), Poranthera (15), Pseudolachnostylis (1), Pseudophyllanthus (1), Savia (2), Securinega (5), Tacarcuna (3) and Wielandia (13). Uses: Several species have edible fruits that are consumed locally, e.g. the currant tree (Antidesma bunius), tampoi (Baccaurea macrocarpa), rambai (B. motleyana), Burmese grape (B. ramiflora), Otaheite gooseberry (Phyllanthus acidus), Jamaican gooseberry

Phyllanthus superbus, Singapore [247]

tree (P. acuminatus), Indian gooseberry (P. emblica) and sugarplum (Uapaca kirkiana). Occasionally some species of Andrachne, Antidesma, Margaritaria and Phyllanthus are grown as garden ornamentals. Etymology: Phyllanthus is composed of the Greek words φύλλων ( fyllon), a leaf and άνθη (anthe), a flower, in reference to the flowers appearing to emerge from the leaf in some species.

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GERANIALES

EUDICOTS

GERANIALES Families 248 and 249 comprise the order Geraniales, which evolved c. 86–80 million years ago. Although a small group, relationships of Geraniales are still poorly understood because they are morphologically heterogeneous, and the phylogenetic tree produced for the order is not yet fully resolved. Geraniaceae together with Hypseocharis (formerly in Oxalidaceae) are sister to a clade combining the former families Melanthiaceae and Ledocarpaceae. The positions of Francoa and Greyia are still uncertain, but it is likely that inclusion of these in Melanthiaceae where they have been placed in former treatments, will make that family polyphyletic if Ledocarpaceae are maintained separate. It is possible to erect Greyiaceae and Francoaceae, but the families share many characters and are therefore united under their oldest name, Francoaceae.

Geraniaceae include annual and perennial herbs and shrubs that occasionally have succulent stems or underground corms. They are usually hairy with simple or glandular hairs, sometimes fragrant. Leaves are alternate or opposite, petiolate and flanked by two paired stipules (absent in Hypseocharis). Blades are simple or palmately or pinnately compound, the margins lobed or toothed and venation pinnate or palmate. Inflorescences are bracteate, often umbellate cymes, or flowers are produced singly or in pairs in the axils of the leaves. The bisexual flowers are

actinomorphic or zygomorphic with five free sepals that often have a narrow tip or awn or with a nectar spur that is fused with the pedicel. The five (rarely two, four or absent) free petals are sometimes shortly clawed and sometimes emarginate at the tip, rarely deeply split, usually entire. Nectaries are five, situated basally on the stamens or one at the tip of the nectar spur. There are five stamens in one whorl or ten or 15 in two whorls, all fertile or some staminodial, sometimes reduced to scales. Filaments are free or shortly fused at the base, the anthers dorsifixed and versatile and opening longitudinally. The superior ovary is composed of five (rarely four) carpels that are fused by their styles to a central column that elongates in fruit. The style is apically divided into five linear stigmas bearing papillae on the inwardfacing side. Fruits are schizocarps that split into five one-seeded or indehiscent parts that are attached to long awns formed from elongate styles, usually catapulted outwards, or a loculicidal capsule (Hypseocharis). In some species the styles form corkscrew-like

Erodium cygnorum, near Coober Pedy, South Australia [248]

Pelargonium zonale, Royal Botanic Gardens, Kew, UK [248]

248. GERANIACEAE Crane’s-bill family

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structures that allow the seeds to bury themselves in the soil under the influence of wind. Distribution: This family has a predominantly temperate and tropical montane distribution, the diversity greatest in Mediterranean climates, culminating in great diversity of Pelargonium in southern Africa. They are remarkably absent from the hot tropics, and despite adaptations to arid environments, they are absent from the great deserts. Phylogeny and evolution: Miocene pollen of Geranium is known from the Miocene of Spain, and Pliocene pollen of Pelargonium is found in Australia. South American Hypseocharis diverged from the rest of the family c. 62–55 million years ago and has been considered a Gondwanan relic. Genera and species: Geraniaceae include five genera and c. 650 species: California (1), Erodium (c. 60), Geranium (260), Hypseocharis (8), Monsonia (40) and Pelargonium (280). Geranium robertianum, Kingston upon Thames, UK [248]

GERANIALES

EUDICOTS

Hypseocharis biloba, Royal Botanic Gardens, Kew, UK [248]

Pelargonium ignescens, Royal Botanic Gardens, Kew, UK [248]

Pelargonium carnosum, Royal Botanic Gardens, Kew, UK [248]

Uses: Geranium oil is used as a flavour and aroma in the food industry, in drinks, candy, ice-cream, puddings, baked goods and jams, and essential oils of scented pelargoniums are important in the perfume industry, especially citronella-scented geranium, Pelargonium graveolens. However, economically most important are the thousands of hybrids and cultivars of Pelargonium, derived from just five species. Crosses of P. inquinans and P. zonale give ‘zonal geraniums’; hybrids of P. cucullatum and P. grandiflorum produce the ‘regal geraniums’ and ‘ivy-leaf geraniums’ are derived from P. peltatum. Numerous cultivars and species of Geranium (crane’s bill) and Erodium (stork’s bill) are also common in horticulture.

Etymology: Geranium is derived from Greek γερανός (geranos), a crane, in reference to the beak-like fruits. Herb Robert (Geranium robertianum) was allegedly named by Linnaeus for one of his pupils who did not wash properly and released a similarly foul odour. However, local names in various languages refer to Robert or Ruprecht predating Linnaeus by many years, and the plant is probably named for Saint Ruprecht of Salzburg (died 710) in reference to its medicinal properties.

absent but can be present and are then lateral or median, sometimes persistent. Inf lorescences are terminal or axillary, simple or compound racemes, cymes or thyrses, sometimes reduced to one, two or three f lowers, sometimes subtended by prophylls or bracts. The bisexual or functionally unisexual (in Bersama) flowers are actinomorphic or zygomorphic and sometimes resupinate (upside-down). Sepals are four or five and petaloid and prominent or not, often basally fused and persistent in fruit. The five (sometimes four or absent) petals are free, but with hairs along the edge that fuse together in Melianthus, and are smaller or larger than the sepals, often showy. There are one or two whorls of stamens, as many as or twice as many as petals (usually ten, sometimes eight, five or four), typically five long and five short, sometimes with one missing in zygomorphic species. Filaments are sometimes fused basally and occasionally hairy, often with a pair of basal appendages. Anthers are dorsifixed or basifixed and open laterally by lengthwise slits. The semiinferior or superior ovary is composed of three to five fused carpels forming as many locules and topped with a single simple style and three to five stigmatic branches or small stigmas. Fruits are papery, unevenly opening capsules, or leathery to woody septicidal or loculicidal capsules.

Carnivory: It has been demonstrated that Geranium viscosissimum has the potential to be a carnivorous plant. The leaves are sticky and catch many insects. The glandular hairs appear to secrete digestive enzymes and are capable of absorbing amino acids. Pelargonium zonale has also been suggested to be carnivorous, but this has not been investigated further. Given the habitat preferences of these two species, they seem unlikely to be truly carnivorous, and these traits exist as protection against predation with the added nutritional benefits of decaying insect carcasses.

249. FRANCOACEAE Bridal-wreath family

Perennial and annual herbs, large shrubs and small trees make up this family. Their leaves are alternate, opposite or in whorls, simple and lobed, pinnatifid, odd-pinnate or trifoliate, with an entire or serrate margin, petiolate or sessile, with the leaf bases sometimes amplexicaul. Stipules are usually

Plants of the World

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EUDICOTS

Viviania marifolia, Los Andes, Chile (MW) [249]

Greyia sutherlandii, Adelaide Bersama abyssinica, Kenya [249] Botanic Garden, South Australia [249]

Rhynchotheca spinosa, Chile (MW) [249]

Melianthus major, planted in a garden in Mount Barker, South Australia [249]

Distribution: Francoaceae are distributed mostly in Andean and southern South America, southern Chile and the tropics and subtropics of Sub-Saharan Africa and southern Chile. Phylogeny and evolution: Fossils belonging to this clade are known from the Miocene. Francoa and Tetilla were previously placed in the distantly related Saxifragaceae (Saxifragales), but they differ markedly from that family in pollen and seed morphology. Melianthus and Greyia were originally associated with Sapindaceae. Most genera have been placed in their own families or even their own orders by some authors, 344

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Francoa appendiculata var. sonchifolia, Royal Botanic Gardens, Kew, UK [249]

and only since the application of molecular phylogenetics has the placement of these genera in one family in Geraniales been suggested, although classification is still disputed. In APG III, Melianthaceae was used for this family, but the name Francoaceae has nomenclatural priority, which is fortunate as the former caused confusion with the monocot family Melanthiaceae (Liliales). The former family Ledocarpaceae (= Vivianiaceae) are embedded in this clade. They are best treated as one family Francoaceae. Genera and species: A family of nine genera and 33 species: Balbisia (8), Bersama (2), Francoa (1), Greyia (3), Melianthus (8),

Balbisia verticillata, Chile (MW) [249]

Rhynchotheca (1), Tetilla (1), Viviania (6) and Wendtia (3). Uses: The dried shoots of Viviania marifolia and several species of Balbisia are made into a herbal infusion in Chile and Argentina called ‘té de burro’, freely translated as donkey-tea. The plants have ornamental potential but are rarely cultivated. Francoa (bridal wreath), Greyia and Melianthus are occasionally grown as garden ornamentals. Melianthus major is a fashionable addition to dry gardens. Etymology: Francoa is named for Francisco Franco, a 16th century Spanish physician.

MYRTALES

EUDICOTS

MYRTALES Families 250 to 258 comprise the order Myrtales, an order that can be recognised by their often flaky bark, included phloem (sandwiched between layers of xylem), opposite leaves with strong intramarginal veins, hypanthium and nectaries, typically clawed petals and an inferior ovary. Myrtales are dated to be c. 79–75 million years old, with the oldest fossils dating back to the late Cretaceous/early Tertiary (65 million years old). These are trees, shrubs, lianas and a few mangroves. Stems are sometimes armed with spines, and the indument is of combretaceous hairs (unicellular thick-walled hairs with a distinctive basal compartment), sometimes replaced by glandular hairs or both types present. The opposite, rarely whorled, spirally arranged leaves are petiolate and simple, the blades with pinnate venation and entire margins. Petioles often have two glands or domatia, and stipules are usually absent or rudimentary. Inf lorescences are axillary

250. COMBRETACEAE Bushwillow family

Lumnitzera littorea, Singapore (WA) [250]

Terminalia rubricarpa, New Caledonia [250]

Laguncularia racemosa, Galápagos Islands [250]

or terminal clusters or heads composed of paniculate spikes or racemes. The bisexual or sometimes unisexual flowers are subtended by deciduous bracts and usually actinomorphic, rarely slightly zygomorphic, with two prophylls fused to the hypanthium (ovary) in some mangrove species. A hypanthium (receptacle) surrounds the ovary and is saucer- to tube-shaped. The four or five (to eight) sepals are formed at the rim of the hypanthium, and the four or five petals are also fused to the hypanthium and often small

Terminalia catappa, habit, Singapore [250]

Combretum indicum, Helsinki Botanical Garden, Finland [250]

Terminalia catappa in fruit, Réunion [250]

Conocarpus erectus var. sericeus, St Petersburg, Florida, USA [250]

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MYRTALES and not showy or absent. Stamens are usually twice as many as sepals, eight, ten (or up to 16), fused to the inside of the hypanthium, often in two whorls, rarely one whorl reduced to staminodes or the second whorl absent. Anthers are dorsifixed and usually versatile, opening by slits. A nectar disc is often basally present in the hypanthium. The unilocular inferior or half inferior ovary is topped with a single, simple style with a pointed stigma. The fruit is a one-seeded indehiscent, often winged capsule. Distribution: This pantropical family is found from Florida and Bermuda throughout tropical America to northern Argentina, Sub-Saharan Africa, Madagascar, the Seychelles and tropical Asia, northern Australia, Micronesia and Melanesia. Phylogeny and evolut ion: Stem Combretaceae are c. 90 million years old. Late Cretaceous fossils have been found widely across the Northern Hemisphere. They are sister to the rest of Myrtales, in which they are morphologically difficult to define. Several genera are polyphyletic, and some of these problems can be solved by expansion of, for instance, Terminalia (including Bucida, Ramatuellea and Terminaliopsis) and Combretum (including Calopyxis, Meiostemon, Quisqualis, Poivrea and Thiloa). A revision of all genera in a phylogenetic framework is underway. Genera and species: Combretaceae include c. 13–17 genera and c. 530 species: Anogeissus (8), Buchenavia (24), Calopyxis (23), Calycopteris (1), Combretum (c. 240), Conocarpus (2), Dansiea (2), Guiera (1), Laguncularia (1), Lumnitzera (2), Macropteranthes (5), Meiostemon (2), Pteleopsis (9), Strephonema (3); Terminalia (c. 200), Terminaliopsis (2) and Thiloa (3). Uses: The Indian almond (Terminalia catappa) is widely planted in the tropics, especially in coastal settings, beaches, streets and parks; it has edible kernels, used as a locally harvested snack. Several larger species of Terminalia are regionally harvested for timber, occasionally

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marketed in Europe and the USA. Buttonwood (Conocarpus erectus) is valued for charcoal. Rangoon creeper, Combretum indicum (perhaps better known as Quisqualis indica), is commonly grown as a garden ornamental. Etymology: Combretum was a name used by Pliny to describe a medicinal herb. It was applied by Pehr Loefling to this group of shrubs, trees and lianas.

251. LYTHRACEAE Pomegranate family

A family of trees, shrubs and herbs, sometimes aquatics and mangroves, they frequently have square stems and sometimes form knee roots (Sonneratia). Their leaves are opposite (rarely whorled or alternate) and simple, with or without stipules. Most genera have glands in the axil at the base of the petiole. Blades have entire margins (toothed in Trapa), and venation is pinnate with the secondary veins arching to form a submarginal vein. Inflorescences are axillary or terminal cymes, racemes, thyrses or spikes, sometimes indeterminate or flowers solitary. The bisexual flowers (unisexual in Capuronia) are actinomorphic to zygomorphic and are mono-, di-, or trimorphic. The usually four to six sepals form a floral tube (hypanthium) that is often strongly veined and usually persistent in fruit, often with alternating external appendages (an epicalyx). The usually four to six petals (rarely many or absent) are inserted on the inner rim of the hypanthium and crumpled in bud and frequently clawed. Two to 50 or numerous stamens are often of two or three lengths, filaments are placed on the hypanthium at the base or at different levels and anthers are dorsifixed, versatile, rarely basifixed, and opening longitudinally. A nectar disc is

often present inside the inner stamen whorl, sometimes unilateral and freestanding or absent. The superior, sometimes semi-inferior or inferior ovary is fused and has two to four or many locules. Fruits are usually dry capsules that open along the locules (or not), sometimes a berry-like splitting capsule with a leathery skin (Punica). Seeds are usually angular and sometimes winged. Distribution: This family has a nearly worldwide distribution. Phylogeny and evolution: Lythraceae are closely related to Onagraceae, forming an early branching clade in Myrtales. The two families separated c. 94 million years ago. The oldest fossils are known from the Deccan Intertrappean Beds that are of Late Cretaceous age. Fossil pollen, fruit, seeds and wood have been found in numerous Palaeocene, Miocene and Eocene deposits around the world. Sonneratia pollen is known from the Oligocene. North American Decodon has a fossil record throughout the Northern Hemisphere, so it was much more widely distributed during the Eocene. The genera Trapa (aquatic) and Sonneratia (mangrove), previously placed in their own families, are sister genera and are, like Punica (formerly Punicaceae), deeply embedded in Lythraceae. Genera and species: There are 30 genera and c. 600 species: Adenaria (1), Ammannia (25), Capuronia (1), Crenea (2), Cuphea (260), Decodon (1), Diplusodon (c. 65), Duabanga (2), Galpinia (1), Ginoria (14), Haitia (1), Heimia (3), Hionanthera (2), Koehneria (1), Lafoensia (6), Lagerstroemia (53), Lawsonia (1), Lourtella (1), Lythrum (36), Nesaea (56), Pehria (1), Pemphis (1), Physocalymma (1), Pleurophora (7), Punica (2), Rotala (44), Sonneratia (6), Tetrataxis (1), Trapa (1–3) and Woodfordia (1). Uses: Pomegranate (Punica granatum) originated in ancient Persia and has been cultivated around the Middle East and Mediterranean for millennia. Subfossil remains of fruits have been found in Bronze Age archaeological sites in Palestine

MYRTALES

EUDICOTS

Sonneratia alba, Singapore (WA) [251]

Heimia myrtifolia, private garden, Kingston upon Thames, UK [251]

Lagerstroemia indica, garden in Curitiba, Brazil [251]

Punica granatum ‘Nana’ in fruit, Helsinki Botanical Garden, Finland [251]

Trapa natans, Dali, Yunnan, China [251]

Cuphea lanceolata, Bergius Botanical Garden, Stockholm [251]

and Cyprus, and a fruit was preserved in the tomb of Queen Hatshepsut in Egypt. Pomegranates have become a symbol of fertility, hope and immortality and were therefore often used as a decorative motif in artwork, ceramics and religious buildings. Seeds are covered with a juicy outer layer, which can be eaten or made into juice, the original grenadine. Pomegranate juice is highly desirable due to the high levels of antioxidants, vitamins C and K and other beneficial compounds. Water caltrop and water chestnut (Trapa bicornis and T. natans) have been cultivated for at least 3,000 years but were once widespread across Eurasia and Africa,

and wild water chestnuts were a significant part of the prehistoric human diet in Europe, where they are now extinct or rare. Until the late 19th century, water chestnut was common in markets around Europe, but it is currently mostly used in Asian cuisine. It is invasive in eastern North America. Lawsonia inermis is the commercial source of henna, the cosmetic dye used in hair dyes and temporary tattoos. Heimia salicifolia has hallucinogenic alkaloids (colouring the world yellow) that also have anti-inf lammatory properties. Several species of Lythraceae produce good timber. Some species are grown as ornamentals. Crepe myrtle (Lagerstroemia indica) is

Lythrum salicaria, Windermere, Lake District, UK [251]

a common street tree in warm temperate and tropical regions, and several species of Cuphea are popular bedding plants, especially Mexican heather (C. hyssopifolia), cigar flower (C. ignea) and tiny mice (C. llavea). Originally introduced as a medicinal plant, purple loosestrife (Lythrum salicaria) became a serious invasive along waterways in North America (although an introduced native pest of the species is now bringing it under control). In its native Eurasia, the species is neither aggressive nor invasive. Etymology: Lythrum is the Latinised form of Ancient Greek λυθρον (lythron), blood, in reference to the colour of the flowers.

Plants of the World

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MYRTALES

EUDICOTS

Clarkia pulchella, Royal Botanic Gardens, Kew, UK [252]

Epilobium angustifolium, Turku, Finland [252]

Circaea lutetiana, Lake District, UK [252]

252. ONAGRACEAE Fuchsia family

These are annual and perennial shrubs, rarely trees to 30 m tall or woody vines. Simple leaves are alternate and spirally arranged, opposite or whorled and often with epidermal oil cells; petioles sometimes have basal stipules, but these usually fall off early or are absent altogether. Blades have entire or toothed to pinnatifid margins and pinnate venation with an intramarginal vein. Inflorescences are axillary in leafy spikes or racemes, occasionally in panicles; 348

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Oenothera biennis, Twickel Estate, Delden, the Netherlands [252]

Lopezia racemosa, Chelsea Physic Garden, London, UK [252]

the f lowers are sometimes solitary. The usually actinomorphic, sometimes somewhat zygomorphic f lowers are bisexual or occasionally unisexual, often with a distinct floral tube (hypanthium) that has nectar glands within. The usually four (sometimes two to five) sepals are fused to the hypanthium and can be green or coloured. The two to four (or five) petals can be clawed or not, or rarely petals are absent. The four or eight (sometimes one or two plus a staminode in Lopezia) dorsifixed and versatile or basifixed anthers open by lengthwise slits. The inferior (rarely semi-inferior) ovary is composed of as many carpels as sepals forming two to four (rarely six) locules with the septa sometimes thin or absent at maturity and topped by a single style and as many stigma lobes as sepals (usually four). Fruits are generally loculicidal capsules, nutlets or berries with numerous seeds. Distribution: A nearly worldwide family, which is absent from dry regions of the

Fuchsia canescens, Ecuador [252]

Ludwigia sedioides, Royal Botanic Gardens, Kew, UK [252]

globe. It is well represented in western North America and the Andes. Phylogeny and evolution: Onagraceae are a well-defined family in the order Myrtales with a close relationship to Lythraceae. Onagraceae are distinguished from other Myrtales by their distinctive four-nucleate embryo sac, and abundant raphides in vegetative cells and pollen grains that are united with viscid threads. Ludwigia (including Jussiaea) is sister to the rest of the family and sometimes placed in its own subfamily, Jussiaeoideae. Within Ludwigia, the taxa with five versus ten stamens form independent clades. Many genera have been recircumscribed following recent molecular findings, resulting in Oenothera being expanded (to include Calylophus, Gaura and Stenosiphon) and Chamissonia split into several genera (Camissoniopsis, Chylisma, Chylismiella, Eremothera, Eulobus, Holmgrenia, Taraxia and Tetrapteron).

MYRTALES

EUDICOTS

Founding father of families — Antoine Laurent de Jussieu (1748–1836) French botanist Antoine de Jussieu was the first to attempt a “natural” classification of flowering plants. In his Genera plantarum (1789), he grouped plants by their overall similarities, and created families and genera based on multiple morphological characters, rather than the single flower characteristics used in Linnaeus’s Sexual System. Many of his family names are still in use, and he can be considered the founding father of the concept of plant families. He nevertheless did not believe in sharp discontinuities between the groups, but rather believed they formed a circular continuum. He was the Professor of Botany at the Jardin des Plantes in Paris, where he reorganised the flower beds by family, an arrangement still found there today. By maintaining the Linnaean binomial system and organising genera in a more natural system of higher ranks, his work assumed enormous importance in the

establishment of a family classification and the continued use of binomials, which greatly benefited scientific communication. Linnaeus named a genus of Onagraceae for him, which is now synonymised under Ludwigia, but his name is maintained at the subfamilial level, Jussieuoideae. Over 60 species are named for him.

Epilobium now includes Boisduvalia, Chamerion and Zauschneria.

many Fuchsia ×hybrida cultivars (old hybrids involving F. fulgens and F. magellanica, and modern ones including other species such as F. coccinea, F. microphylla, F. splendens, F. thymifolia, etc.). Frost-hardy F. magellanica (especially ‘Riccartonii’) is often grown as hedging and locally naturalises, for example in western Ireland. Other frequently grown ornamentals are the New Zealand creeping fuchsia (Fuchsia procumbens), zauschneria (Epilobium canum), fireweed (E. angustifolium), gaura (Oenothera lindheimeri) and several evening primroses (e.g. O. biennis, O. glazioviana, O. macrocarpa, O. salicifolia, O. speciosa). Ludwigia species (e.g. L. hyssopifolia, L. sedioides) are sometimes grown as pondplants and have escaped to become noxious invasives of aquatic habitats outside their natural range.

Genera and species: Onagraceae include 22 genera and c. 656 species: Camissonia (12), Camissoniopsis (14), Chylismia (16), Chylismiella (1), Circaea (10), Clarkia (41), Epilobium (c. 170), Eremothera (7), Eulobus (4), Fuchsia (106), Gayophytum (9), Gongylocarpus (2), Hauya (2), Holmgrenia (2), Lopezia (22), Ludwigia (82), Megacorax (1), Neoholmgrenia (2), Oenothera (152), Taraxia (4), Tetrapteron (2) and Xylonagra (1). Uses: Flowers and fruit of Fuchsia are sometimes eaten in salads, and the roots and leaves of most Oenothera species (evening primrose) are edible. Some species of Oenothera are grown for their medicinal seed oil. Several species of Onagraceae are important horticultural plants. Bedding plants include godetia (Clarkia amoena), scented clarkia (C. breweri), garden clarkia (C. unguiculata), and

Engraving of Antoine de Jussieu by Charles Thévenin in 1803

Etymology: Onagra was assumed to be the food of the onager (Equus hemionus). Onagra is a later synonym of Oenothera, the evening primrose.

Onagraceae in the family beds in the Jardin des Plantes, Paris (FT)

253. VOCHYSIACEAE Quaruba family

These shrubs to tall trees accumulate aluminium and often have stellate hairs. Leaves are opposite or whorled and simple, petiolate with entire margins and pinnate venation with a submarginal vein. Stipules are often modified into glands or are associated with extrafloral nectaries. Inflorescences are terminal or axillary thyrses, cincinni or racemes, the flowers with prophylls. The bisexual flowers are strongly zygomorphic and have five basally fused sepals that are often unequal in size, the largest usually Plants of the World

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MYRTALES

Vochysia tomentosa, Montagne de Kaw, French Guiana (AP) [253]

Erisma uncinatum, Brazil (IP) [253]

Ruizterania esmeraldae, Colombia (MF) [253]

Qualea cordata, Tobati, Paraguay (CD) [253]

Qualea cordata, fruit, Tobati, Paraguay (CD) [253]

with a spur. The one, three or five petals are free and clawed (rarely absent). The single fertile stamen is free and has a dorsifixed or basifixed anther that opens by longitudinal slits. There are often also up to four staminodes. The superior or inferior ovary is uni- to trilocular and topped with a simple style and a terminal or lateral stigma. Fruits are loculicidal capsules or samaras that become four- or five-winged due to enlarged, persistent calyx lobes. Seeds are often winged and hairy.

dispersal. They were originally associated with Malpighiales, due to their zygomorphic flowers, but molecular analyses placed them as sister to Myrtaceae in Myrtales.

254. MYRTACEAE

Distribution: This family is most commonly found in the Neotropics, but three species occur in tropical West Africa. Phylogeny and evolution: The family originated in South America, with the two African genera having been established there some 30 million years ago via long-distance

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EUDICOTS

Christenhusz, Fay & Chase

Myrtle family

Genera and species: Vochysiaceae include seven genera and 218 species: Erisma (20), Erismadelphus (2), Korupodendron (1), Callisthene (10), Qualea (60), Salvertia (1), Ruizterania (19) and Vochysia (105). Uses: The seeds of Erisma calcaratum are locally used to manufacture soap. The wood of quaruba (Vochysia hondurensis) is locally used as timber. Generally, wood of this family is of low quality. Several species are grown as garden ornamentals in Latin America. Etymology: Vochysia is derived from vochy, the Galibi name for a tree of this genus in French Guiana.

Myrtaceae are a family of evergreen, bisexual trees and shrubs that usually have obvious glands in cavities producing ethereal oils (usually terpenes) on foliage, branchlets and flowers. The opposite or alternate, sometimes whorled, leaves are simple and all similar or dimorphic, with juvenile ones of different size or shape than those of older plants. Stipules are absent or small and unnoticeable and in

MYRTALES

EUDICOTS

that case also fall off early. Blades have entire margins, and venation is pinnate or with three main veins in some genera, often with veins running along the margin. Inflorescences are axillary or terminal cymes variously arranged into panicles, thyrses, botryoids and dichasial cymes or reduced to one flower. Flowers are bisexual (occasionally unisexual) and mostly actinomorphic with a hypanthium that is fused to the ovary and prolonged above it. The four or five (rarely three) sepals are free or fused into a cap (calyptra), the margins entire to deeply dissected. The four or five petals are free or also fused into a cap, sometimes petals absent, and occasionally the fused petals fall as an entity when the flower opens. Stamens are usually numerous in one to several whorls. Usually filaments are free, sometimes fused into clusters or fascicles that are opposite the petals. Anthers are dorsifixed or basifixed and open usually by lengthwise, sometimes terminal, slits with connectives bearing one or more glands at the tip. The inferior, semi-inferior or rarely superior (subfamily Psiloxyloideae) ovary is composed of (one or) two to five, rarely more, carpels forming one to many locules. A terminal style is set into a pit on the ovary tip or at the base of the ovary (rarely), usually with a capitate or lobed stigma at the top. Fruits are variable, usually a dry loculicidal or indehiscent capsule, or a one- to many-seeded berry, rarely a drupe. Distribution: This family has a pantropical and southern temperate distribution. It is found in the Neotropics and temperate

Heteropyxis natalensis, South Africa (AD) [254]

South America, the Mediterranean region, Sub-Saharan Africa, Madagascar, the Mascarenes, tropical and (warm) temperate Asia, Australia, New Zealand and throughout the Pacific islands. It is an important component of the Australian and Brazilian floras, forming the dominant species in most plant communities, with many endemics. Phylogeny and evolution: The oldest fossil record of Myrtaceae is from the upper Cretaceous, from Argentina, tropical Africa, Borneo, China and Australia, which corresponds to the estimated molecular age of the family (c. 84–90 million years). Even though pollen of Myrtaceae is distinctive, fossil pollen can rarely be placed below the level of family and is therefore not useful as calibrations to enhance estimates of divergence times within the family. The application of names of extant genera to leaf fossils, sometimes erroneously so, confuses the picture of fossil evolution, but Eucalyptus is known from the Miocene of New Zealand and the Eocene of Argentina. Xanthomyrtus fossils from the Eocene of Tasmania are similar to the leaves of extant taxa from Malesia and New Caledonia. Myrtaceae were traditionally divided into two subfamilies based on fruits being dry or fleshy, resulting in an unnatural classification; the fleshy-fruited species mostly forming a clade that is derived from a dry fruited grade. The two subfamilies accepted here are based on molecular evidence and morphological characters (Psiloxyloideae have spirally arranged leaves, are dioecious

Eucalyptus macrocarpa near Geraldton, Western Australia [254]

and have a different pollen type). Their rapid divergence during the Oligocene and Miocene is likely to have been a response to aridification. Psiloxylon may have hopped around Indian Ocean islands for some 40 million years, but it is now only found on Mauritius. Generic delimitations are unclear, and many issues need to be addressed. Syzygium has been sometimes included in Eugenia, although they are not exclusively related. Syzygium has been expanded to include Acmena, Acmenosperma, Anetholea, Piliocalyx and Waterhousea. Other genera that need attention are Eucalyptus, Eugenia, Melaleuca and Myrcia, all of which are more likely to be expanded to include genera rather than being dismembered to maintain smaller entities. Genera and species: Myrtaceae include 140 genera and 5,828 species in two subfamilies: Psiloxyloideae – Heteropyxis (3) and Psiloxylon (1); Myrtoideae – Acca (3), Accara (1), Actinodium (1), Agonis (5), Algrizea (2), Allosyncarpia (1), Aluta (5), Amomyrtella (2), Amomyrtus (2), Angophora (14), Archirhodomyrtus (5), Arillastrum (1), Astartea (9), Asteromyrtus (7), Astus (4), Austromyrtus (8), Babingtonia (2), Backhousia (9), Baeckea (47), Balaustion (1), Barongia (1), Basisperma (1), Beaufortia (20), Blepharocalyx (4), Callistemon (46), Calothamnus (43), Calycolpus (15), Calycorectes (28), Calyptranthes (268), Calyptrogenia (6), Calytrix (79), Campomanesia (39), Chamelaucium

Eucalyptus jacksonii, Walpole, Western Australia [254]

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EUDICOTS

Pimenta dioica, cultivated in Réunion [254]

Darwinia masonii, Australian National Botanical Garden, Canberra [254]

Calothamnus quadrifidus, Mt Benia, Western Australia [254]

(13), Chamguava (3), Cheyniana (2), Choricarpia (2), Cloezia (5), Conothamnus (3), Corymbia (92), Corynanthera (1), Curitiba (1), Cyathostemon (1), Darwinia (53), Decaspermum (34), Enekbatus (10), Eremaea (14), Eucalyptopsis (2), Eucalyptus (754), Eugenia (1,044), Euryomyrtus (7), Gossia (37), Harmogia (1), Homalocalyx (11), Homalospermum (1), Homoranthus (31), Hottea (9), Hypocalymma (23), Kanakomyrtus (6), Kania (6), Kardomia (6), Kjellbergiodendron (1), Kunzea (41), Lamarchea (2), Legrandia (1), Lenwebbia (2), Leptospermum (87), Lindsayomyrtus (1), Lithomyrtus (11), Lophomyrtus (2), Lophostemon (4), Luma (2), Lysicarpus (1), Malleostemon (6), Marlierea (90), Melaleuca (256), Meteomyrtus (1), Metrosideros (54), Micromyrtus (50), Mitranthes (7), Mitrantia (1), Mosiera (32), Myrceugenia (44), Myrcia (396), Myrcianthes (37), Myrciaria (26), Myrrhinium (1), Myrtastrum (1), Myrtella (2), Myrteola (3), Myrtus (2), Neofabricia (3), Neomitranthes (15), Neomyrtus (1), Ochrosperma (6), Octamyrtus (6), Osbornia (1), Oxymyrrhine (4), Pericalymma (4), Petraeomyrtus (1), Phymatocarpus (3), Pileanthus (8), Pilidiostigma (6), Piliocalyx (8), Pimenta (15), Pleurocalyptus 352

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(2), Plinia (68), Pseudanamomis (1), Psidium (95), Purpureostemon (1), Regelia (1), Rhodamnia (40), Rhodomyrtus (21), Rinzia (12), Ristantia (3), Sannantha (15), Scholtzia (13), Seorsus (4), Siphoneugena (11), Sphaerantia (2), Stenostegia (1), Stereocaryum (3), Stockwellia (1), Syncarpia (3), Syzygium (1,136), Taxandria (11), Tepualia (1), Thaleropia (3), Thryptomene (32), Triplarina (7), Tristania (1), Tristaniopsis (39), Ugni (4), Uromyrtus (23), Verticordia (101), Welchiodendron (1), Whiteodendron (1), Xanthomyrtus (23) and Xanthostemon (48). Uses: Myrtaceae include many fruit species, of which the guava (Psidium guajava), originally from South America, is the most commonly cultivated. Guava is eaten fresh or pressed into juice, sometimes on a commercial scale. Strawberry guava (P. cattleyanum) is locally grown and has been introduced as a fruitcrop in many places, often with disastrous results. Psidium species have become devastating invasives in many areas, especially tropical islands where they crowd out the local flora. Other minor fruit crops are feijoa or pineapple guava (Acca sellowiana), grumichama (Eugenia brasiliensis), rainforest

Ugni candollei, Royal Botanic Gardens, Kew, UK [254]

plum (E. candolleana), Rio Grande cherry (E. involucrata), pitomba (E. luschnathiana), Cedar Bay cherry (E. reinwardtiana), Suriname cherry or pitanga (E. uniflora), Chilean murta (Ugni molinae), jabuticaba (Plinia cauliflora), myrtle berry (Myrtus communis), water apple (Syzygium aqueum), waterberry (S. cordatum), jambolan plum (S. cumini), roseapple (S. jambos), riberry (S. luehmannii), Malay apple or pommerac (S. malaccense), blue lilly pilly (S. oleosum), magenta lilly pilly (S. paniculatum) and Java apple (S. samarangense). Economically most important are the various Eucalyptus species grown for timber in plantations around the world. The most widely grown species is the blue gum (E. globulus) from Tasmania, although others are also frequently grown, depending on their climatic tolerance. Eucalyptus plantations create serious ecological problems outside Australia because their leaves often decay poorly and flamable oils can be a fire hazard that can then spread into surrounding native vegetation. They can also be a disperal barrier for native species to spread, due to the inhospitable vegetation that a Eucalyptus plantation provides. Essential oils from

MYRTALES

EUDICOTS

Eucalyptus (eucalyptus oil) are used medicinally and frequently as flavourings of sweets and candy, cough medication and a scent oil in cosmetics, perfumes and steam rooms. Several culinary spices are represented in Myrtaceae. These include clove (flower buds of Syzygium aromaticum) from India, allspice (berries of Pimenta dioica) from the Caribbean and to a lesser extent the Australian lemon myrtle (Backhousia citriodora) and aniseed myrtle (Syzygium anisatum), all used as food flavourings. 
The second most important species economically is the spice clove (Syzygium aromaticum). Clove oil was the earliest widely used essential oil, which contains eugenol, a natural analgesic. It is also antibacterial and was used to treat toothache; it is still used in modern dentistry. The dried pedicels and flower buds of cloves are frequently used as a spice in cooking or as a scent and have been known to be used as a spice or medicine outside its native range since the 2nd and 3rd centuries BC in Syria

and China, respectively. During the Middle Ages, cloves were the main commodity traded across the Indian Ocean by Muslim traders, leading to stories like the Seven Voyages of Sindbad the Sailor of the Arabian Nights. Tea-tree oil is extracted from Melaleuca alternifolia, which is highly antimicrobial and anti-inflammatory. Other essential oils are niaouli (M. quinquenervia), lemon-scented oil (from Backhousia citriodora, Corymbia citriodora, Leptospermum petersonii) and bay rum (Pimenta racemosa). Berries and leaves of myrtle (Myrtus communis) are made into an aromatic liqueur called mirto in Sardinia and Corsica. Myrtle was a sacred plant in Greek and Roman mythology, being associated with the goddesses Aphrodite and Venus, respectively. It is the symbol of love and immortality, and hence they are commonly used in wedding bouquets and wreaths. A sprig from Queen Victoria’s wedding bouquet was rooted and sprigs of this same plant have featured in several British royal wedding bouquets since. Crowns of myrtle are used in

Calytrix leschenaultii, Australian National Botanical Garden, Canberra [254]

Psidium guajava, Easter Island [254]

some wedding rituals, especially in Ukraine. A number of species are valuable garden ornamentals and street trees, especially species of Callistemon, Calothamnus, Chamelaucium, Corymbia, Eucalyptus, Eugenia, Leptospermum, Luma, Melaleuca, Metrosideros, Myrtus, Rhodomyrtus, Syncarpia, Verticordia and Xanthostemon. Geraldton wax (Chamelaucium uncinatum) is commonly cultivated in the cut-flower industry. Tallest flowering plant: The giant ash, Eucalyptus regnans, is the tallest known angiosperm, with the current tallest specimen 101 metres tall, but before logging there may have been trees of over 122 metres. Etymology: Myrtus or murtus is classical Latin for a myrtle bush. This is in turn derived from μυρτιά (myrtia) or μυρσίνη (myrsine), the Greek name for the plant, probably derived from murdinnu or murdunnu, Sumerian or Akkadian for plant. Myrtus communis, in fruit, Royal Botanic Gardens, Kew, UK [254]

Corymbia calophylla, Mt Benia, Western Australia [254] Xanthostemon ruber, New Caledonia [254] Verticordia ovalifolia, Mt Benia, Western Australia [254]

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MYRTALES

or truncate stigma. The fruit is a loculicidally dehiscent or indehiscent, dry or fleshy capsule or a berry.

This is a family of annual and perennial herbs, shrubs and trees (up to 20 m tall), sometimes climbing, usually terrestrial, rarely epiphytic or aquatic. Simple leaves are usually opposite, rarely whorled, and lack stipules. Blades are equal in size or with one of a pair slightly smaller than the other, rarely one or a pair aborted. Margins are entire or serrate, with palmately pinnate venation and usually one to four (or five) secondary veins on each side of the midvein originating at or near the base and converging at the apex, the tertiary veins numerous, parallel, and connecting secondary veins with the midvein (Melastomatoideae); in Olisbeoideae, the secondary veins are pinnate, and tertiary veins are reticulate. Domatia when present are

represented by pits or pockets. Inflorescences are basically cymes, composed of umbels, corymbs or panicles, or reduced to fascicles, cincinni, spikes, single cymes, clusters or rarely solitary f lowers. Inf lorescence bracts are sometimes brightly coloured and persistent, and flowers are subtended by two opposite bracteoles. The bisexual flowers are actinomorphic, but stamens and associated structures sometimes on one side, making the flower slightly zygomorphic. The flower base forms a funnel-, bell-, cup- or urn-shaped hypanthium topped with the usually four or five (sometimes three to six) free or fused sepals. Petals are free, usually four or five, usually equal the number of sepals. Stamens are usually twice as many as petals (usually eight or ten, rarely up to 96) and in two whorls, rarely as many as petals by loss of a whorl, sometimes one whorl staminodial. Filaments are free, often jointed, and anthers are typically basifixed with a variously appendaged or ornamented connective that is often colourful. Anthers open by one or two apical pores or longitudinal slits. The inferior or semi-inferior (rarely superior) ovary is composed of usually four or five (rarely three to 14) carpels fused into as many locules. It is topped by a single style and a minute capitate

Brachyotum jamesonii, Ecuador [255]

Miconia calvescens, Tahiti [255]

Tibouchina pulchra, São Paulo, Brazil [255]

255. MELASTOMATACEAE Senduduk family

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EUDICOTS

Christenhusz, Fay & Chase

Distribution: This is a predominantly pantropical family extending into the subtropics and temperate zones in eastern North America, southern Brazil, the Middle East, China and Japan. Phylogeny and evolution: The family is estimated to have diversified c. 41–53 million years ago. Even though the current distribution sometimes appears to be linked to continental drift, it seems that more recent long-distance dispersal between the continents during the Miocene is more likely. Rhexia, now restricted to North America, is found in Tertairy fossil deposits throughout Eurasia, which has implications for biogeographical scenarios and phylogenetic dating of the tree. Olisbeoideae are sister to Melastomatoideae, and the former are sometimes treated as a separate family, Memecylaceae. Pternandra is sister to the rest of Melastomatoideae and has in the past also been treated at subfamilial level (Kibessioideae). Crypteroniaceae, formerly included in Melastomataceae, are more closely related to Alzateaceae and

MYRTALES

EUDICOTS

Memecylon elaeagni, Seychelles [255]

Medinilla intermedia, Royal Botanic Gardens, Kew, UK [255]

Melastoma malabathricum, Singapore Botanical Garden [255]

Penaeaceae. Leandra is polyphyletic and needs to be recircumscribed on the basis of seed morphology.

Heterotrichum (10), Huberia (14), Kendrickia (1), Kerriothyrsus (1), Killipia (5), Kirkbridea (2), Lavoisiera (46), Leandra (c. 200), Lithobium (1), Loreya (14), Loricalepis (1), Macairea (22), Macrocentrum (15), Macrolenes (15), Maguireanthus (1), Maieta (3), Mallophyton (1), Marcetia (23), Mecranium (24), Medinilla (c. 375), Melastoma (22), Melastomastrum (6), Meriania (74), Merianthera (3), Miconia (c. 1,000), Microlepis (4), Microlicia (170), Mommsenia (1), Monochaetum (45), Monolena (9), Myriaspora (1), Neblinanthera (1), Neodriessenia (6), Nepsera (1), Ochthocharis (2), Omphalopus (1), Opisthocentra (1), Orthogoneuron (1), Osbeckia (50), Ossaea (c. 80), Oxyspora (24), Pachyanthus (16), Pachycentria (8), Pachyloma (6), Phainantha (4), Phyllagathis (56), Physeterostemon (3), Pilocosta (5), Pleiochiton (7), Plethiandra (7), Podocaelia (1), Poikilogyne (20), Poilannammia (4), Poteranthera (2), Preussiella (2), Pseudodissochaeta (5), Pseudoernestia (1), Pseudosbeckia (1), Pternandra (15), Pterogastra (2), Pterolepis (14), Quipuanthus (1), Rhexia (11), Rhynchanthera (15), Rousseauxia (13), Rupestrea (2), Salpinga (8), Sandemania (1), Sarcopyramis (3), Schwackaea (1), Scorpiothyrsus (6), Siphanthera (16), Sonerila (c. 175), Sporoxeia (6), Stanmarkia (2), Stenodon (1), Stussenia (1), Svitramia (6), Tateanthus (1), Tayloriophyton (2), Tessmannianthus (6), Tetraphyllaster (1), Tetrazygia (25), Tibouchina (c. 350), Tibouchinopsis (2), Tococa (47), Topobea (c. 50), Trembleya (11), Triolena (22), Tristemma (15), Tryssophyton (1), Vietsenia (4) and Wurdastom (8).

are some species with edible fruit such as senduduk or Malabar melastome (Melastoma malabathricum), which stains the mouth when eaten (hence the name, see Etymology). Other species with edible fruit include Bellucia grossularioides, B. pentamera, Blakea spp., bush currants (some Clidemia and Miconia spp.), Conostegia xalapensis, Heterotrichum spp., Loreya collata, Macrolenes muscosa, korakaha (Memecylon umbellatum), Mouriri spp. etc. Some species provide reasonable timber, especially Astronia, Calycogonium and Memecylon, and Dionycha boyeri and Memecylon umbellatum are used locally for dyes. Melastomataceae have also been used as soil indicators to classify Amazonian rain forests. Many species are grown as ornamental plants, especially Bertolonia maculata and hybrids, B. mosenii, Centradenia inaequilateralis, Clidemia rubra, Dissotis rotundifolia, Medinilla magnifica, Melastoma malabathricum, Memecylon umbellatum, Monolena primuliflora, Oxyspora paniculata, Rhexia virginica, Sonerila margaritacea, Tetrazygia bicolor, Tibouchina urvilleana and Triolena pustulata and many others are potentially good garden and park ornamentals. However, some naturalise easily, such as Koster’s curse (Clidemia hirta) and Miconia calvescens, which have become noxious weeds in many tropical countries, especially on Pacific and Indian Ocean islands.

Genera and species: Melastomataceae have two subfamilies with 178 genera and c. 5,000 species: Olisbeoideae – Lijndenia (10), Memecylon (c. 150), Mouriri (81), Spathandra (6), Votomita (6) and Warneckea (31); Melastomatoideae – Acanthella (2), Aciotis (13), Acisanthera (c. 20), Adelobotrys (31), Allomaieta (8), Allomorphia (25), Alloneuron (4), Amphiblemma (13), Amphorocalyx (5), Anaectocalyx (3), Anerincleistus (30), Antherotoma (2), Appendicularia (1), Arthrostemma (4), Aschistanthera (1), Astrocalyx (1), Astronia (59), Astronidium (67), Axinaea (c. 25), Barthea (1), Behuria (14), Bellucia (8), Benevidesia (2), Bertolonia (17), Bisglaziovia (1), Blakea (c. 100), Blastus (12), Boerlagea (1), Boyania (1), Brachyotum (50), Brasilianthus (1), Bredia (30), Bucquetia (3), Cailliella (1), Calvoa (19), Calycogonium (36), Cambessedesia (21), Campimia (1), Castratella (2), Catanthera (17), Catocoryne (1), Centradenia (6), Centradeniastrum (2), Centronia (16), Chaetolepis (1), Chaetostoma (11), Charianthus (6), Cincinnobotrys (7), Clidemia (117), Comolia (22), Comoliopsis (1), Conostegia (43), Creochiton (6), Cyphostyla (3), Cyphotheca (1), Dalenia (3), Desmoscelis (1), Dicellandra (3), Dichaetanthera (34), Dinophora (2), Dionycha (3), Dionychastrum (1), Diplarpea (1), Diplectria (11), Dissochaeta (20), Dissotis (c. 120), Dolichoura (1), Driessenia (14), Enaulophyton (2), Eriocnema (1), Ernestia (16), Feliciadamia (1), Fordiophyton (14), Fritzschia (1), Graffenrieda (44), Gravesia (110), Guyonia (2), Henriettea (67), Heterocentron (28),

Uses: Although this is a large family, there are few species of economic interest. There

Etymology: Melastoma is composed of the Greek words μέλανος (melanos), black, and στόμα (stoma), mouth, because when the berries are eaten, they stain the mouth black. Plants of the World

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Crypteronia paniculata, Minang Kob, Sabah, Malaysia (CD) [256]

Crypteronia paniculata, Minang Kob, Sabah, Malaysia (CD) [256]

Dactylocladus stenostachys, Klias Forest Reserve, Sabah, Malaysia (CD) [256]

256. CRYPTERONIACEAE

Distribution: This family is found in Southeast Asia, Sri Lanka and throughout the Malesian Archipelago east to New Guinea and Melanesia.

Etymology: Crypteronia is composed of the Greek word κρυπτώ (krypto), secret, and έρως (eros), love, in reference to the small flowers.

Bekoi family

These evergreen trees have simple opposite leaves with an entire margin and pinnate venation. They often have small stipules, but these may be absent. Inflorescences are spikes or racemes bearing many bisexual or unisexual flowers. The four or five fused sepals (calyx) are minute or nearly absent. The minute four or five petals can also be absent, the perianth with one or two whorls, sometimes the corolla falling off as a calyptra when the flower opens (Axinandra). Stamens are bent downwards in bud, pushing away the corolla when becoming upright. The connective is often enlarged, and anthers open by longitudinal slits. The inferior or semi-inferior ovary is bi- to hexalocular and topped with a style terminated by a capitate stigma. Fruits are capsules with few to many flat, winged seeds.

356

EUDICOTS

Christenhusz, Fay & Chase

Phylogeny and evolution: This family probably evolved during the Late Cretaceous and is at most 65 million years old. The only fossil evidence is of Miocene pollen from Borneo. The family has been associated with Melastomataceae in the past on the basis of anatomical characters, but this has since been disputed, although placement in Myrtales is undoubtedly correct. Molecular evidence places them close to Alzateaceae and Penaeaceae. It has been suggested that Crypteroniaceae originated in Africa and reached Southeast Asia by drifting on the Indian subcontinent, a calculation based on a molecular clock, but long-distance dispersal is an equally likely explanation. Dactylocladus is sister to the rest with weak support. Genera and species: This poorly known family consists of three genera and c. 13 species: Axinandra (5), Crypteronia (7) and Dactylocladus (1). Uses: The young shoots of Crypteronia paniculata are eaten with rice in Malaysia.

257. ALZATEACEAE Wantsum family

These small evergreen trees and shrubs grow sometimes partly epiphytically and have quadrangular stems. Leaves are opposite or whorled, the blades simple and margins entire with pinnate venation and parallel secondary venation. Stipules are two or a few surrounding the node. Inf lorescences are axillary thyrses formed at the tips of branches. The actinomorphic, bisexual flowers lack petals or petals are rudimentary. The five (or six) sepals are thick, fleshy and fused at the base forming a bell-shaped floral tube. Petals are absent, rarely rudimentary and deliquescing. The five stamens are fleshy and inserted between the

MYRTALES

EUDICOTS

sepals at the margin of a broad, lobed nectary disc. The short filaments are tipped with a large heart-shaped, pinkish connective, and anthers are dorsifixed and open by a lateral slit. The superior ovary is composed of two locules, flattened from the sides, and topped by a short, stout style. Fruits are flattened dry capsules that open along the locules, and the flattened seeds are encircled by a thin wing.

Genera and species: Alzateaceae consist of the single species Alzatea verticillata. Two subspecies that are probably not distinct have sometimes been accepted at species level.

Distribution: Alzatea is found in small populations along the eastern slopes of the Andes in Bolivia, Peru and Colombia and on mountain slopes in Panama and Costa Rica.

258. PENAEACEAE

Etymology: Alzatea is named for Mexican astronomer and geographer José Antonio de Alzate y Ramírez (died 1795).

Cape-fellwort family

Phylogeny and evolution: Alzateaceae are most closely related to Penaeaceae with which they form a clade that evolved 53–92 million years ago. These families share stout styles, quadrangular stems, opposite or whorled, simple leaves and reduced petals. Alzatea verticillata, Suiza Nueva, Oxapampa, Peru (CD) [257]

Penaea cneorum, Langeberg Mountains, South Africa (CD) [258]

Alzatea verticillata, habit, Panama (PM) [257]

These are (heather-like) shrubs, shrublets and large trees, the last often with fluted or buttressed stems (Olinia). The young stems are square, and stem nodes are swollen, the ridges ending often in four tooth-like processes on each side of the leaf base or young shoots terete (Rhynchocalyx). Opposite or whorled leaves are petiolate and simple, the blade entire (irregularly toothed in Sonderothamnus) and pinnately veined. Stipules are rudimentary or withered. Inflorescences are variable, determinate or not, terminal or axillary panicles, thyrses or spikes, usually composed of three-flowered cymules with opposite bracts, or flowers are solitary. The bisexual flowers are actinomorphic with a tubular hypanthium (or disc-shaped in Rhynchocalyx) that has (four or) five teeth in Olinia. The four to six sepals are showy and petal-like and inserted on the hypanthium rim. Olinia has (four or) five petals on the inner side of Saltera sarcocolla, Fernkloof, Western Cape, South Africa (CD) [258]

Sonderothamnus speciosus, South Africa (JA) [258]

Rhynchocalyx lawsonioides, Umtamvuna Nature Reserve, Eastern Cape, South Africa (CD) [258]

Plants of the World

357

MYRTALES the rim that are much shorter than the sepals and scale-like and hairy; Rhynchocalyx has six narrowly clawed petals with a stalk and a round, hood-like lamina, but in other genera petals are absent. The four to six stamens are inserted below the sepals on the hypanthium rim and often incurved with short filaments and dorsifixed or basifixed, longitudinally opening anthers. The inferior (or superior in Rhynchocalyx) ovary is composed of two to five fused carpels forming as many locules. The stout style usually remains inside the floral tube, is often angular or winged and has a capitate stigma. The fruit is a drupe with a thin flesh, pink or red when ripe (Olinia), or a

EUDICOTS

loculicidal capsule, dorsiventrally compressed in Rhynchocalyx. Distribution: This family is found in southern and eastern Africa and on Saint Helena, with the greatest diversity in the Cape region. Phylogeny and evolution: Olinia has been difficult to place on morphological grounds, but molecular studies placed it sister to Penaeaceae with Rhynchocalyx sister to the rest. The two genera (Rhynchocalyx and Olinia) have in the past been treated as separate families (Rhynchocalycaceae and Oliniaceae), but they share many characters.

Genera and species: Penaeaceae include nine genera with 32 species: Brachysiphon (5), Endonema (2), Glischrocolla (1), Olinia (8), Penaea (4), Rhynchocalyx (1), Saltera (1), Sonderothamnus (2) and Stylapterus (8). Uses: Penaea is a source of edible gum that taste like liquorice. Some species of Olinia have in the past been exploited for their timber. Etymology: Penaea is named in honour of French physician and botanist Pierre Pena (c. 1520–1600), who was an assistant to Matthieu de l’Obel (Matthias Lobelius).

CROSSOSOMATALES Families 259 to 265 form the order Crossosomatales, on morphological grounds an unanticipated assembly of genera that were previously placed in other families, although often they were anomalous in these. As families, they share the presence of a hypanthium, free styles and seed coats with much-thickened outer layers. The order has two clades that are split between the Northern (Crossosomataceae, Guamatelaceae, Stachyuraceae, Staphyleaceae) and Southern Hemispheres (Aphloiaceae, Geissolomataceae, Strasburgeriaceae), a result of a combination of vicariance and long-distance dispersal.

259. APHLOIACEAE Mountain-peach family

Aphloia theiformis, Réunion [259]

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Evergreen shrubs and slender trees make up this family. Leaves are alternate, in one plane, and the blades are simple with serrate or serrulate (rarely nearly entire) leaf margins. The stipules are small and fall off early. Inflorescences are few-flowered axillary racemes or fascicles, flowers are solitary. The bisexual, actinomorphic flowers are subtended by minute scale-like bracts. The tepals are usually four or five (rarely six), spirally

arranged and white, the inner ones more petallike. Stamens are numerous, the filaments free and inserted on the outer edge of the nectary disc. Small globose anthers are basifixed and open by lateral slits. The superior ovary is composed of a single carpel with one locule. The stigma is sessile on the ovary and large, bilobed-capitate. The fruit is a fleshy white berry surrounded by persistent sepals. Distribution: The family is restricted to East Africa (from Kenya to Mozambique), Madagascar, the Mascarene Islands, Comoros and Seychelles. Phylogeny and evolution: Previously placed in the now dismembered Flacourtiaceae, from which they differ anatomically, molecular evidence placed them in the polymorphic order Crossosomatales. There is substantial morphological and sequence variation in Aphloia.

CROSSOSOMATALES

EUDICOTS

Geissoloma marginatum, Marloth Nature Reserve, Langeberg Mountains, South Africa (CD) [260]

Ixerba brexioides, fruit, New Zealand (JC) [261]

Geissoloma marginatum, Marloth Nature Reserve, Langeberg Mountains, South Africa (CD [260])

Ixerba brexioides, New Zealand (JC) [261]

Strasburgeria robusta, New Caledonia (JA) [261]

Genera and species: The sole genus and species in this family is Aphloia theiformis, which is polymorphic and probably consists of about eight rather than a single species.

cordate and has a thickened margin. Stipules are subulate and sit on either side of the petiole-like leaf base. Inflorescences are uniflorous, terminating axillary bracteate shoots, the bracts increasing in size, the upper ones petal-like, often bearing vestigial flower buds. The bisexual flowers are actinomorphic and have four tepals that are basally fused and persistent in fruit. The eight stamens in two whorls are attached at the base to the floral tube with free slender filaments and dorsifixed, longitudinally opening anthers. A nectary disc is present at the base of the floral tube. The superior ovary is composed of four partially fused sessile carpels, each with a locule and tipped with four free styles. Fruits are four-lobed, hard capsules enclosed in the flower tube and opening along the locules, each locule with a single seed.

with either of these. Molecular analyses placed it as sister to Strasburgeriaceae, and hence it is placed in Crossosomatales.

Uses: Tea from the leaves of Aphloia theiformis is used in local medicine. It has a beneficial effect on the urinary tract. Etymology: Aphloia is composed of the Greek prefix a- (a-), without, and φλοιός ( floios), bark.

260. GEISSOLOMATACEAE Cape-cups family

Genera and species: The family consists of a single genus and species: Geissoloma marginatum. Etymology: Geissoloma is composed of the Greek words γείσος (geisos), the eaves of a house, and λωμα (loma), a fringe, in reference to the imbricate aestivation of the petals.

261. STRASBURGERIACEAE Tawari family

Distribution: This family is restricted to the Langeberg Mountains in the Cape Province of South Africa, where they grow in fynbos.

Densely branched, leafy, low shrubs that accumulate aluminium make up this family. The opposite leaves are leathery, nearly sessile, with a simple blade that is basally

Phylogeny and evolution: Originally described as a species of Penaea (Penaeaceae, Myrtales), Geissoloma marginatum was placed in that family or Celastraceae by early botanists, although it has little in common

A family of evergreen trees, glabrous or in Ixerba with unicellular lignified T-shaped hairs. Leaves are alternate, opposite or whorled and lack stipules. Blades are simple, Plants of the World

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CROSSOSOMATALES

Staphylea colchica ‘Hessei’ in fruit, Royal Botanic Gardens, Kew, UK [262]

Staphylea colchica, Royal Botanic Gardens, Kew, UK [262]

Staphylea trifolia forma pyriformis, Royal Botanic Gardens, Kew, UK [262]

with close or widely set serrate (sometimes gland-tipped) margins and pinnate venation, the petioles sometimes with narrow wings. Inflorescences are terminal, few-flowered cor ymbose (umbel-like) panicles, or flowers are solitary in the leaf axils. The bisexual f lowers are actinomorphic with a short hypanthium fused to the base of the ovary. The five (Ixerba) or eight to ten (Strasburgeria) free sepals are persistent in fruit, the outer ones shorter, the inner enclosing the bud. The five (rarely six) petals are free, clawed or not, and inserted on the hypanthium. The five or ten stamens are free and alternate with the nectary disc lobes, and anthers are dorsifixed and open by longitudinal slits. The superior (or semiinferior) ovary is composed of five (Ixerba) or four to seven (Strasburgeria) fused carpels, each forming a locule topped by a five-grooved style and a lobed stigma. The fruit is a few-seeded, five-lobed loculicidal capsule, or indehiscent, the seeds large, sometimes with a rudimentary aril.

or Celastraceae, and Strasburgeria was often placed in or near Ochnaceae. Neither placement was sustained by morphological similarities. Molecular analyses placed the two genera together as sister to Geissolomataceae, which is also supported by morphological characters like wood anatomy and floral structure. Zealandia broke away from Australia 60–85 million years ago, and fossil pollen of Strasburgeria is known from Tertiary deposits in Antarctica (Wilkes Land), southern and southwestern Australia, Tasmania and New Zealand, suggesting that its current restriction to New Caledonia is relictual.

262. STAPHYLEACEAE

Distribution: Strasburgeriaceae are a family distributed in the continental fragment called Zealandia. Ixerba occurs in the North Island of New Zealand, and Strasburgeria is endemic to mountain forests on ultrabasic soil in New Caledonia. Phylogeny and evolution: Ixerba was often considered to be a member of Saxifragaceae

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Genera and species: A family of two genera, each with a single species: Ixerba brexioides and Strasburgeria robusta. They are sometimes placed in their own families but share numerous morphological characters. Uses: Tawari (Ixerba brexioides) is occasionally used as bee fodder for honey production in New Zealand. Etymology: Strasburgeria is named for Polish-German botany professor Eduard Adolf Strasburger (1844–1912), who established the modern laws of plant cytology. Ixerba is an anagram of Brexia (Celastraceae), to which it was thought to be related. Although Ixerbaceae is the older name, Strasburgeriaceae has been nomenclaturally conserved.

Bladdernut family

Staphyleaceae are deciduous and evergreen trees and shrubs. Leaves are opposite, odd-pinnately compound or trifoliolate, rarely simple, with or without stipules and the leaflets stalked and pinnately veined, the margins serrate. Inflorescences are terminal or axillary panicles near the end of branches. The bisexual (rarely unisexual) flowers are actinomorphic and have five usually petal-like sepals and five free or basally fused petals. The five stamens have free filaments emerging from the disc, and their dorsifixed anthers open by longitudinal slits. The superior ovary is composed of two, three or four carpels that are nearly free, composed of a weakly united, lobed ovary with as many locules as carpels. Styles are free or slightly united at the capitate stigma. Fruits are inflated capsules, dry follicles or berry-like drupes. Distribution: A family distributed across the Northern Hemisphere, in temperate regions

CROSSOSOMATALES

EUDICOTS

(Mexico, eastern North America, southeastern Europe, the Black Sea Region, Himalayas and East Asia) and tropical Central and northern South America, the Caribbean, southern India, Sri Lanka, tropical East Asia, Malesia and New Guinea.

Uses: The fermented flower buds of bladdernut (Staphylea colchica) are eaten in the Caucasus. The wood is not durable but has been used for timber. Rapidly growing species can be used for reforestation and prevention of erosion. Several species are grown as ornamentals.

Phylogeny and evolution: The fossil record of Staphyleaceae corresponds more or less to its current distribution, and seeds of several extant taxa have been found in Tertiary deposits. Relationships of Staphyleaceae (often tentatively placed in Sapindales) could not be unravelled on morphological grounds. Molecular studies placed the family with Crossosomataceae and Stachyuraceae in Crossosomatales. The genera traditionally recognised were based on fruit morphology and turned out not to be monophyletic. Turpinia was, for instance, found to be biphyletic, the type species being part of the Staphylea clade, the remainder part of Dalrympelea, the two genera now accepted in this family. Tapiscia and Huertea, previously placed here, have always been thought to be different from other members of the family, and they are unrelated and now placed in Tapisciaceae (Huerteales).

Etymology: Staphylea is derived from the Greek staphyle, a cluster.

Genera and species: There are only two genera with c. 45 species: Dalrympelea (c. 23) and Staphylea (23).

263. GUAMATELACEAE

ovary is composed of three sessile carpels that are free below, tipped with fused styles that separate after the flower opens. Each style is tipped with a capitate stigma. Ovaries are sparsely hairy with unicellular hairs. The fruit is a follicle, generally three per flower, opening along the ventral suture, exposing numerous seeds with a membranous aril. Distribution: Southern Mexico (Oaxaca, Chiapas), Guatemala, Honduras.

Guatemalan-bramble family Phylogeny and evolution: Guamatela has traditionally been included as an isolated genus in Rosaceae, where it was unique among the Mesoamerican species with its opposite leaves. Molecular studies indicated that the genus should be excluded from Rosaceae and placed in Crossosomatales in which it is treated as a separate family.

These are branched shrubs with opposite petiolate leaves and stipules. Simple, ovate blades have serrate margins and are palmately veined. Inflorescences are terminal racemes or panicles with actinomorphic flowers. The five free, persistent sepals, five free petals and ten distinct stamens are attached to a shallow hypanthium. Anthers are ovate and dorsifixed and open longitudinally. The connective has an apiculum at the tip. The semi-inferior

Genera and species: This monotypic family consists only of Guamatela tuerckheimii. Etymology: Guamatela is an anagram of the country name Guatemala, where the type was collected. The country in turn is derived from Nahuatl Cuauhtēmallān, meaning ‘place of many trees’.

264. STACHYURACEAE Spiketail family

Guamatela tuerckheimii, La Selva Negra, Chiapas, Mexico (CD) [263]

These shrubs and small trees, sometimes vines, are deciduous and evergreen. Alternate, simple leaves are petiolate and have linear-lanceolate stipules. Leaf blades have serrate to serrulate margins and pinnate venation. Inflorescences are axillary racemes or spikes that are nodding or sometimes erect, with basally fused bracteoles below each f lower. The actinomorphic f lowers

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CROSSOSOMATALES are bisexual or unisexual with dioecious plants; bracteoles are connate at base. The four sepals are in two whorls, the outer two smaller. The four petals are free, yellow, greenish, pink or white. The eight stamens form two whorls, with free filaments and versatile, dorsifixed, sagittate anthers that open by longitudinal slits. The superior ovary is composed of four fused carpels forming four incomplete locules. The style is simple, short, with a shallowly four-lobed capitate stigma. Fruits are berries with leathery pericarp and numerous seeds with soft arils.

EUDICOTS

generally accepted, their systematic position has long been controversial. They were considered related to Actinidiaceae, Clethraceae, Flacourtiaceae, Hamamelidaceae, Theaceae or Violaceae by different authors based on evidence from embryology, seed anatomy, wood anatomy and palynology. DNA analysis placed the family as sister to Crossosomataceae.

265. CROSSOSOMATACEAE Rockflower family

Genera and species: The sole genus in this family, Stachyurus, has about eight species.

Phylogeny and evolution: Although the familial status of Stachyuraceae has been

Etymology: Stachyurus is composed of the Greek words stachys, a spike and oura, a tail, referring to the slender racemes of these shrubs.

This is a family of shrubs, often microphyllous, rarely trees. Stems are spiny or have black hairs. Leaves are alternate or opposite, scattered along stems or clustered in fascicles. Stipules are usually absent or minute. The bisexual or rarely unisexual flowers occur solitarily in the leaf axils or terminate short shoots and are actinomorphic. The usually four or five (rarely three or six)

Stachyurus himalaicus in fruit, Royal Botanic Gardens, Kew, UK [264]

Stachyurus praecox, Caerhays Estate, Cornwall, UK [264]

Crossosoma californicum in fruit, Rancho Santa Ana Botanical Garden, California, USA [265]

Stachyurus praecox var. leucotrichus, Royal Botanic Gardens, Kew, UK [264]

Glossopetalon spinescens var. aridum, Nevada, USA (SS) [265]

Distribution: This family is distributed across East Asia: Japan, the Ryukyu Islands, Taiwan and much of China.

Uses: Some species are occasionally grown as garden ornamentals, especially Stachyurus praecox.

Crossosoma californicum (JG) [265]

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EUDICOTS

sepals are equal or unequal and persistent on the hypanthium rim. The free petals are the same number as and alternating with the sepals, often shortly clawed. There are four to 50 stamens that are sometimes of unequal length; anthers are basifixed and open by lateral slits. The superior ovary is composed of one to five (to nine) free carpels that are stalked or sessile, each with a single locule and capitate stigma. Fruits are ventrally opening dry follicles containing disc-shaped seeds with a fimbriate aril.

Distribution: This family occurs in xeric habitats in the southwestern USA and northwestern Mexico. Phylogeny and evolution: The position of Crossosomataceae remained controversial until molecular analysis indicated that they form an order with, among others, Stachyuraceae and Staphyleaceae, families that also were difficult to place on morphological grounds. Crossosoma is sister to the remainder of the family.

Genera and species: This small family includes four genera and c. ten species: Apacheria (1), Crossosoma (3), Glossopetalon (5) and Velascoa (1). Etymology: Crossosoma is composed of the Greek words κρόσσος (krossos), fringe, and σώμα (soma), a body, in reference to the fringed aril of the seed.

PICRAMNIALES This recently recognised order diverged c. 108 million years ago. They are morphologically superficially similar to Simaroubaceae, in which they were formerly placed.

266. PICRAMNIACEAE Bitterbush family

Picramnia pentandra, infructescence (WJ) [266]

This is a family of unisexual trees and shrubs. The spirally arranged leaves are odd-pinnate and lack stipules. Leaflets are alternate or opposite and stalked. Inf lorescences are terminal or axillary, long slender thyrses or racemes. The unisexual flowers are small and actinomorphic. The three to five (or six) sepals are fused at their base and persistent in fruit. The petals are small, reduced in female flowers and sometimes absent in male flowers. In male flowers there are as many stamens as there are Alvaradoa amorphoides, Mary Selby Botanical Garden, Florida, USA [266]

sepals (three to five), sometimes fused into a column, and a pistillode is sometimes present. In female flowers there are often three to five staminodes. The superior ovary is composed of two or three fused carpels that are borne on a small disc or stalk (gynophore) and is uni- to trilocular. The stylodia are short, one per carpel, strongly recurved with a papillose stigma on the upper side. Fruits are berries or compressed winged capsules (samara), the calyx and styles persistent.

Alvaradoa amorphoides, Mary Selby Botanical Garden, Florida, USA

[266]

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PICRAMNIALES Distribution: Picramniaceae are restricted to the New World subtropics and tropics, where they occur from southern Florida and central Mexico to Paraguay and northwestern Argentina. Phylogeny and evolution: This family was previously placed in Simaroubaceae, but molecular studies have shown that they are not related, placing Picramniaceae in an isolated position between the rosid I and II clades. Because of their isolated position of uncertain affinity, the family is in its own order Picramniales. Tariric acid, a fatty acid, is rare outside this family. The family is dated to an estimated 108 million years old. Genera and species: This is a family of three genera and 49 species: Alvaradoa (5), Nothotalisia (3) and Picramnia (41). Uses: The fruits of some Picramnia are edible, and the wood of some species is locally used. Etymology: Picramnia is a contraction of the Greek words πικρός (pikros), bitter, and θάμνος (thamnos), a shrub.

EUDICOTS

GUMILLEA (UNPLACED) Gumillea auriculata, previously placed in Cunoniaceae and unplaced in APG IV, may belong to Crossosomatales or Picramniales. As its position remains uncertain, we mention it here separately. They are branched shrubs with oddpinnate alternate leaves, basally attached with pinnule-like sessile stipules, branched spikelike inflorescences with sessile, bisexual, actinomorphic flowers that have five fused sepals, five stamens and free bicarpellate, superior ovaries with many ovules topped with recurved stigmas. It has not been recollected after its initial discovery in Peru, and herbarium specimens on which the illustration and description must have been based could not be located. Therefore no molecular data are available for this species, and placement has to be based on just morphological characters from the original description and associated illustration. Stipules (even though they may be basal leaflets) are absent in Picramniaceae, with which it shares many characters (the recurved styles, pinnate alternate leaves, elongate spikes, lack of petals), but the flowers

are apparently bisexual and not unisexual as is the case in Picramniaceae. It also shares characters with Staphyleaceae, but if placed in that family it will be unique with sessile flowers and no petals. The clade to which this plant belongs remains a mystery.

Gumillea auriculata and Hydrocotyle triflora (Araliaceae) by Jose Brunete (1777–1816), original drawing (MA), used for the basis of the publication by Ruiz & Pavón, in which this taxon was described

Biebersteinia multifida, Iraq (TM) [267]

SAPINDALES Families 271 to 279 comprise the order Sapindales, which include c. 3% of eudicot diversity and have relatively high diversification rates. The order emerged 90.5–177.4 million years ago and is united in having free styles with fused stigmatic heads. Most species are woody, apart from the herbaceous Biebersteiniaceae, which are sister to all other members of this order. Many species produce unusual compounds, resulting in many medicinal applications.

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267. BIEBERSTEINIACEAE Khardug family

This family comprises perennial, foetid herbs with woody, tuberous rhizomes. Leaves alternate along the stem or are nearly all basal. The (false) stipules are fused to the petiole. Leaf blades are pinnate with the pinnae one- to three-times pinnatisect. Inflorescences are bracteate panicles or spikes, sometimes condensed into heads. The actinomorphic, bisexual, flowers are subtended by two bracteoles. The five sepals are free and persistent, sometimes enlarged in fruit. The five free petals are yellow, orange or white, often toothed at the apex. The five fleshy nectary glands sit inbetween the petals. The ten stamens have filaments that are fused at the base and have dorsifixed anthers that open longitudinally and inwards. The superior, shortly stalked ovary is composed of partially fused carpels forming a deeply five-lobed structure with five styles free at the base that are connected at their tips in a capitate stigma. Fruits are schizocarps consisting of five indehiscent dry nutlets, each with a single seed.

Phylogeny and evolution: Biebersteinia was originally compared with Suriana (Surianaceae) and Grielum (Neuradaceae), and aff inites with Zygophyllaceae, Rosaceae, Rutaceae and especially Geraniaceae were later suggested, mostly due to superficial similarities. Molecular analysis has shown it to be an independent herbaceous lineage in Sapindales, in which it may be sister to all other families of this order, a placement that agrees with its phytochemistry and f lower morphology. The origin of Biebersteinia can be traced to China, and its age has been estimated to be c. 60 million years. Genera and species: The sole genus in the family is Biebersteinia with four species. Uses: Root stocks of some species were collected from the wild in the past and eaten, but today are considered a famine food. Biebersteinia heterostemon has immuneregulatory properties and is widely used in Chinese traditional medicine. The leaves have a foetid smell.

Distribution: This family is distributed from the eastern Mediterranean to Central Asia, western China and western Siberia usually in montane, semi-arid or forested habitats.

Etymology: Biebersteinia is named for German botanist Baron Friederich August Marschall von Bieberstein (1768–1826). As secretary for a Russian general, the Count of Kochovski, he was posted in the Crimea in 1793. Inf luenced by German naturalist P. S. Pallas, he started collecting plant material from Crimea and throughout the Caucasus when he accompanied the Russian forces invading Persia. He is famed for his Flora Taurico Caucasica (1808– 1819), a comprehensive floristic account of the region.

Nitraria billardieri, fruiting, Dalhousie, South Australia [268]

Nitraria billardieri, Dalhousie, South Australia [268]

268. NITRARIACEAE Nitrebush family

Perennial and annual herbs and shrubs to 2 m tall that are creeping or upright and often succulent make up this family. Branches are often spiny at the tips, and leaves are alternate (rarely opposite in Tetradiclis), arranged spirally along the stem or in fascicles with intrapetiolar, free stipules, sometimes Peganum harmala, Kars, Turkey [268]

Peganum harmala, Harran, Turkey [268]

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SAPINDALES bristle-like or absent (Peganum). Blades are simple or irregularly pinnatifid (Peganum, Malacocarpus), petiolate or nearly sessile, usually succulent and deciduous. There is a single midvein or the venation is palmate. The margins are entire and pinnatifid or have two or three teeth at the tip. Inflorescences are spike-like cymes, or flowers are solitary, subtended by bracts. The actinomorphic flowers are bisexual. The four or five fleshy sepals are persistent in fruit. The four or five white or yellowish petals are free and usually cucullate. Stamens are four or ten to 15 and alternating and opposing the petals in one or three whorls, the filaments free, often dilated below, and the anthers dorsifixed and opening by lengthwise slits. A sessile, inferior ovary is composed of two to six fused carpels each forming a locule. The single style is topped with an ovoid, lobed stigma. Fruit is a purple, red or yellow, fleshy drupe with a bony seed (Nitraria), berry (Malacocarpus) or dry, tri- to tetralocular capsule (Peganum, Tetradiclis). Distribution: Nitrariaceae can be found in arid and semi-arid regions of eastern Mexico (along the Gulf), North Africa, southern Europe, Arabia, southern Siberia, warm temperate and boreal Asia to western China and North Korea and southwestern and southern Australia. Phylogeny and evolution: This family was previously included in Zygophyllaceae, but they differ in many morphological, chemical and anatomical characters. Molecular studies place them in Sapindales. The two entities formerly called Nitrariaceae and Tetradiclidaceae/Peganaceae are molecularly divergent and also differ in several characters. However, they are closely related and share many characters of floral structure and are thus now united into one family. Genera and species: Nitrariaceae include four genera and 19 species: Malacocarpus (1), Nitraria (12), Peganum (5) and Tetradiclis (1). Uses: Fruits of Nitraria schoberi and others are eaten locally (e.g. Siberia, Australia

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EUDICOTS

etc.), but those of N. retusa are used as an intoxicant in Arabia. Plants accumulate salt and can be burned to extract it from the ashes. Woody species are also sometimes used as firewood, and Nitraria is sometimes planted to prevent erosion. A dye called Turkey red is produced from the fatty oils of the seeds of harmal, Peganum harmala, which is used to dye tarbooshes, a kind of hat, and wool for the red in carpets and traditional clothing. Peganum harmala contains harmine, a narcotic drug, which in small doses is used in the treatment of asthma, hysteria and hiccups and was applied as a ‘truth drug’ by the Nazis in World War II. This species is used in traditional folk medicine and witchcraft and as an aphrodisiac (carbolines are sexual stimulants). Fruits are also made into amulets to fend off evil spirits. The plants were burned as an intoxicant in the Middle East and Central Asia for some 7,000 years and are probably the source of hallucinations such as flying carpets. It is a dominant species in areas with heavy grazing by goats, which do not eat this plant due to its toxicity (harmalol). Etymology: This genus was named Nitraria by J. C. D. von Schreber because he first found it at Siberian nitre works (saltpetre mines), together with other salt-loving plants.

269. KIRKIACEAE White-seringa family

Kirkiaceae are shrubs and trees to 20 m tall, often with a thickened base below ground. Deciduous leaves with no stipules are alternate, usually crowded at the branch tips, leaving scars on the bark after falling. The unevenly pinnate leaf blades have up to 30 lateral pinna pairs that are pinnately

veined with a serrate to crenate margin. Leaf axes are winged or not, and leaflets are sometimes stalked. Inflorescences are axillary or cauliflorous compound thyrses (dichasial and monochasial cymes). Morphologically bisexual, but functionally unisexual, flowers are actinomorphic. Generally male flowers have enlarged anthers and reduced ovaries, whereas in female flowers stamens are shorter and sterile (staminodia). Numbers of flower parts vary; although they are generally fourparted, in lower parts of the inflorescence they can be five- or six-parted and in upper parts they are sometimes three-parted. Thus, the three to six free (or shortly, basally fused) sepals are mirrored by the same number of linear, free petals. Stamens are free, alternating with petals, and inserted on the outside of a swollen, fleshy disk. Filaments are often broader towards the base and topped by dorsifixed anthers that open by lateral slits. The superior, shortly stalked ovary is composed of four or eight fused carpels, and situated atop the disk. Free styles are four or eight, apical on the ovary and fused just at the capitate stigmas. Fruits are woody schizocarps that break into four or eight single-seeded nutlets. Distribution: This family is restricted to tropical East Africa, from Ethiopia and Somalia to South Africa, and in eastern Namibia and western Madagascar, usually in dry habitats often on limestone hills or in wooded savannas. Phylogeny and evolution: Kirkia was in some morphology-based classifications placed in Simaroubaceae, but molecular studies have shown that when broadly circumscribed Simaroubaceae were polyphyletic, comprising of elements now in Malpighiales, Picramniales and various families in Sapindales. Kirkia lacks the chemistry (quassinoids, limonoids) typical of Simaroubaceae, and molecular analyses place Kirkiaceae close to the Burseraceae–Anacardiaceae clade. The segregate genus Pleiokirkia from Madagascar, with double the number of carpels, has been merged with Kirkia, with which it shares all other characters.

SAPINDALES

EUDICOTS

Kirkia acuminata, inflorescence, Malawi (GB) [269]

Genera and species: This family includes a single genus, Kirkia, with six species. Uses: Kirkia acuminata provides good quality wood, used for a great variety of purposes from building materials to utensils, arts and crafts, furniture, cabinetry, carts, agricultural tools, packaging material and toys. This and other species are also used to make charcoal, and bark can be used to make string or woven into cloth. In times of drought, the tuberous roots of these trees can be chewed to quench thirst. There are also medicinal applications of the bark and fruit. Etymology: Kirkia was named in honour of Scottish physician, naturalist and explorer Sir John Kirk (1832–1922). He accompanied explorer David Livingstone on the Zambesi Expedition. He was also instrumental in banning the slave trade in East Africa.

270. BURSERACEAE Frankincense-and-myrrh family

Kirkia acuminata, fruit, South Africa (OM) [269]

This is a family of commonly unisexual trees and shrubs, sometimes epiphytic or scandent, which can be recognised by their resin canals (schizogenous canals) in the vascular tissues. The sap is colourless or white and usually highly scented. They have alternate, pinnate leaves that are sometimes reduced to a single terminal leaflet, appearing simple or bipinnate or trifoliolate, the petioles often with a swollen pulvinus at the base and the rachis sometimes winged. Stipules are rarely present and often false. Leaf blades are often pelluciddotted, and margins of the opposite (alternate in Boswellia) pinnae are often toothed. Inf lorescences are axillary or (pseudo-) terminal, rarely cauliflorous thyrses that often appear as spikes, fascicles or racemes. The actinomorphic flowers are usually unisexual but occasionally bisexual, and a hypanthium is sometimes present. The three to five partially fused sepals are persistent and sometimes accrescent in fruit. The three to five (or six) petals are free or basally fused. The one or two whorls of stamens are as many or twice as many as the petals, with the filaments usually basally fused and anthers sometimes confluent with the filaments, or arrow-shaped and basifixed, opening with lengthwise slits. Stamens are sterile (staminodia) in female flowers. An intrastaminal disk is usually present, but it can be extrastaminal or adnate to the receptacle. Female flowers have a superior (or half inferior) ovary composed of

two to five (rarely to 12) fused carpels, each forming a locule, which is reduced or absent in male flowers. A single style is shortly branched near the apex, and stigmas have as many lobes as there are carpels or are capitate. Fruits are dry or fleshy compound drupes or false capsules. Seeds are inside a pyrene, which is enveloped in a berry-like fruit or separated by a columella, the pyrene winged or arillate. Distribution: This is a pantropical family found from southern North America (southwestern USA and Mexico), throughout the Caribbean and Central America to southern Brazil, Sub-Saharan Africa, Madagascar and the Mascarenes, southern Arabia, tropical Asia into warm-temperate China and south to northern Australia and southern Pacific islands. Phylogeny and evolution: The crown group of Burseraceae is dated to the early Palaeocene (c. 60–65 million years ago). The family exhibits a boreotropical distribution, and all lineages were widespread across the Northern Hemisphere by the end of the Late Oligocene (23 million years ago). Fossil fruits and pyrenes have been found, for instance, in the Eocene and Oligocene deposits of the London Clay. Several genera have recently been recircumscribed. Tapirocarpus, previously

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Bursera microphylla, San Antonio Botanical Garden, Texas, USA [270]

Boswellia sacra, Royal Botanic Gardens, Kew, UK

Commiphora opobalsamum, Ruissalo Botanical Garden, Turku, Finland [270]

Canarium vulgare, Singapore (WA) [270]

placed here, is a synonym of Talisia (Sapindaceae). Ambilobea is segregated from Boswellia because it is not immediately related to the Boswellia alliance, but rather closer to Canarium. Paraprotium is a segregate from Protium but may be part of that genus. Several genera have not yet been (fully) sampled in molecular phylogenetic studies. Pseudodacryodes, Rosselia and Scutinanthe have not yet been included, and sampling of these in future studies may result in further realignments of the genera.

(C. odontophyllum) and the Chinese olive (C. album) are also consumed locally, and all are high in protein, fats and carbohydrates. Eben or safu nuts (Dacryodes edulis) are also eaten locally. Many species are of local cultural importance and especially well known are the species responsible for products of Biblical fame. Frankincense or olibanum (Boswellia sacra) is the dried sap harvested from this and other Boswellia species native to the southern Arabian Peninsula and the Horn of Africa. It has been traded in Arabia for some 5,000 years and has traditionally been used in many religious ceremonies by Greeks, Romans and Jews since early times because it was a symbol of divinity and an emblem of prayer. This pure white fragrant substance was considered an appropriate gift for the Biblical Magi (wisemen from the East) to present to the infant Jesus Christ. Frankincense is still used in Christian religious ceremonies (and the smoke is lightly hallucinogenic), but olibanum is also highly valued by the perfume industry. Myrrh, a natural gum from the resin of Commiphora myrrha, C. abyssinica and other species, was also offered to Christ by the Magi and has a similarly long tradition. Apart

from incense in religious ceremonies, myrrh is used as an antiseptic in mouthwashes and toothpaste and in some healing salves. The resin of copperwood (Bursera simaruba) can be used as an incense, glue or varnish. The wood of this species was traditionally used to make carousel horses in the United States. Tabonuco or candlewood (Dacryodes excelsa) from the Caribbean produces a hard aromatic waxy resin that can be used to make candles and incense and was used by native Tainos in Puerto Rico to make torches. The wood is comparable to mahogany and can be used for construction, cabinetry, wood carving etc. Many other species of Dacryodes and also some Aucoumea, Bursera, Canarium, Protium and Santiria produce similarly valuable timber.

[270]

Genera and species: A family with c. 20 genera and c. 760 species: Ambilobea (1), Aucoumea (1), Beiselia (1), Boswellia (c. 20), Bursera (c. 100), Canarium (c. 120), Commiphora (c. 185), Crepidospermum (7), Dacryodes (c. 70), Garuga (4), Haplolobus (16), Paraprotium (4), Protium (c. 180), Pseudodacryodes (1), Rosselia (1), Santiria (22), Scutinanthe (2), Tetragastris (9), Trattinnickia (14) and Triommia (1). Uses: Several species of Canarium have edible nuts, the most important being the galip nut (C. indicum) and the pili nut (C. ovatum). Fleshy avocado-like fruits of dabai 368

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Etymology: Bursera is named in honour of German physician and botanist Joachim Burser (1583–1639), who travelled extensively around Europe, collecting many plants. He became Professor of Botany at Sorø Academy in Denmark in 1625, from where his collection found its way to the Botanical Museum in Uppsala and was a great help in the botanical studies of Carolus Linnaeus.

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anther that usually opens longitudinally. The superior ovary is composed of a single or up to five fused carpels, each forming a locule, bearing two to five free styles or a simple style with a capitate, disc-shaped, lobed or spathe-like stigma. Fruits are usually drupes, sometimes an achene or samara, rarely syncarps, utricles or berries.

These are trees, shrubs and vines that often have resinous bark and sap that turns black upon exposure to air. Resin canals are present in most parts of these plants. Leaves are usually alternate, rarely opposite, and simple, trifoliate or pinnately compound, rarely palmate or bipinnate, and lack stipules. Leaflets are opposite or alternate with pinnate

venation and an entire, toothed or crenate margin. Inflorescences are axillary or terminal panicles, thyrses, or spikes, rarely cauliflorous or flowers solitary. The actinomorphic (rarely slightly zygomorphic) flowers are unisexual (and then plants often unisexual as well) or bisexual, often with an articulate pedicel. The three to five sepals are fused at least at the base, sometimes forming a cap (calyptra), persistent or not, sometimes enlarged in fruit. Usually the four to five (rarely three, up to eight or absent) petals are free, sometimes fused at the base, surrounding an annular disk. Stamens are inserted on the disk margin and usually as many or twice as many as petals, normally five to ten, sometimes one or up to >100 and placed in one or two whorls; in some genera there are only one or two fertile stamens, the others staminodial. Filaments are free or fused at the base and support a dorsifixed or basifixed

Rhus typhina, female, Mackinac Island, Michigan, USA [271]

Rhus typhina, male, Mackinac Island, Michigan, USA [271]

Toxicodendron rydbergii, Muir Woods, California, USA [271]

Mangifera odorata, Singapore (WA) [271]

Cotinus coggygria, East Bergholt Place Arboretum, Suffolk, UK [271]

271. ANACARDIACEAE Cashew family

Distribution: This family has a primarily pantropical distribution, with a few species extending into the temperate zones and greater species richness in the Palaeotropics. It is found from southern Canada to Patagonia, in southern Europe, northern Africa, Sub-Saharan Africa, Madagascar, southern Arabia, western and southern Asia, Southeast and East Asia to Japan and Korea, and throughout the Malesian and Pacific islands south to northern and eastern Australia.

Anacardium occidentale, planted near Malindi, Kenya [271]

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Sclerocarya birrea, Hluhluwe Game Reserve, KwaZulu Natal, South Africa (CD) [271]

EUDICOTS

Mangifera indica, Ecuador [271]

Phylogeny and evolution: Anacardiaceae diverged from Burseraceae 51–64 million years ago. Many Late Tertiary fossil taxa are known from genera that are now restricted to East Asia, and Anacardium (now restricted to the Neotropics) is known from Mid Eocene deposits in Germany. The distinctive fruits of the Old World Dracontomelon are known from Late Eocene deposits in Panama. Amphipterygium and Orthopterygium were segregated into Julianiaceae in some morphology-based classifications, but these two genera are embedded in Anacardiaceae, with which they share many characters. Similarly, Campylopetalum and Dobinea were sometimes segregated as Podoaceae, which have also been demonstrated to belong here. There are two subfamilies recognised, although Spondiadoideae are probably not monophyletic in their current circumscription. Generic delimitation is poorly investigated in many cases and needs to be revised in light of new molecular findings; it seems unnecessary that there should be so many small unclearly delimited genera. Genera and species: Anacardiaceae have two subfamilies with a total of c. 83 genera and c. 860 species: Anacardioideae (60 genera) – Abrahamia (19), Actinocheita (1), Amphipterygium (5), Anacardium (11), Apterokarpos (1), Astronium (7), Baronia (3), Blepharocarya (2), Bonetiella (1), Bouea (3), Campnosperma (13), Campylopetalum (1), Cardenasiodendron (1), Comocladia (16), Cotinus (4), Dobinea (2),

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Drimycarpus (3), Euleria (1), Euroschinus (9), Faguetia (1), Fegimanra (3), Gluta (30), Haplorhus (1), Heeria (1), Holigarna (7), Laurophyllus (1), Lithraea (3), Loxopterygium (3), Loxostylis (1), Malosma (1), Mangifera (69), Mauria (15), Melanochyla (30), Melanococca (1), Metopium (3), Micronychia (10), Mosquitoxylum (1), Myracrodruon (2), Nothopegia (10), Ochoterenaea (1), Orthopterygium (1), Ozoroa (40), Pachycormus (1), Parishia (4), Pentaspadon (6), Pistacia (12), Protorhus (1), Pseudosmodingium (4), Rhodosphaera (1), Rhus (35), Schinopsis (7), Schinus (30), Searsia (c. 120), Semecarpus (75), Smodingium (1), Sorindeia (9), Swintonia (12), Thyrsodium (7), Toxicodendron (22) and Trichoscypha (32); Spondiadoideae (23 genera) – Allospondias (2), Androtium (1), Antrocaryon (3), Attilaea (1), Buchanania (30), Choerospondias (1), Cyrtocarpa (5), Dracontomelon (8), Haematostaphis (1), Haplospondias (1), Harpephyllum (1), Koordersiodendron (1), Lannea (c. 40), Operculicarya (8), Pegia (2), Pleiogynium (2), Poupartia (7), Poupartiopsis (1), Pseudospondias (2), Sclerocarya (3), Solenocarpus (2), Spondias (16) and Tapirira (8). Uses: The cashew tree (Anacardium occidentale), originally from northeastern Brazil, is widely grown commercially for several economically important products. The seeds, cashew nuts, are roasted as a popular snack or added to curries and sauces. They are also ground into a peanut-butter-like spread, powdered to make a beverage or pressed to

Malosma laurina, Santa Catalina Island, California, USA [271]

produce a cooking oil. The swollen pedicel above the fruit is called a cashew apple and of a more local importance because it transports poorly. Unripe cashew apples are too astringent to consume. It can be eaten fresh, made into juice or fermented into alcoholic beverages. Shells of cashew nuts contain a resin called cashew nutshell liquid, which is used industrially in the manufacture of plastics, glues, lubricants etc. Other species of Anacardium also produce nuts that are edible after roasting. Mango (Mangifera indica) is an important tropical fruit crop worldwide, which was domesticated in India c. 5,000 years ago, probably from wild M. sylvatica. Mangoes can be eaten fresh and are juiced or pickled as a chutney. Fruit skins contain allergens that can cause dermatitis. The culprit, mangiferin, is an antimicrobial compound used pharmaceutically and extracted from leaves and bark. A dye, India yellow, is made from urine of cows that have been fed on mango leaves. Marula fruit (Sclerocarya birrea) has become economically more important due to the recent export of the liqueur Amarula Cream, which is made of this East African fruit. Pistachio (Pistacia vera), a species domesticated in dry regions of Central Asia, is roasted to make a popular snack and used in confectionery, baked goods, desserts and ice-cream. Bombay mastic (P. atlantica) produces, apart from gum mastic, edible fruits that have been eaten since the Neolithic. Young shoots of P. chinensis are used as a vegetable

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EUDICOTS

in Chinese cuisine. Mastic tree (P. lentiscus) produces a gum that has been chewed in the Mediterranean since ancient times. Currently, it is most commonly harvested and chewed on the Greek island of Chios, from where it is also exported for use as varnish, dental filler and treatment for halitosis. It is also used as a flavouring for food and drink. Rose pepper (Schinus terebinthifolius) originated in Brazil but is commonly grown throughout the tropics as a citrus-peppery spice. It is invasive in Florida, Australia, South Africa and oceanic islands (Mascarenes, Hawaii). The related Peruvian pepper tree or mulli (S. molle) produces similar fruit that are sometimes used to adulterate pepper. Seeds of Rhus coriaria are used commonly in the Middle East as the spice sumac, and leaves of this species can be used for tanning leather and were important in the production of parchment. Fruits of smooth sumac (Rhus glabra) and velvet sumac (R. typhina) are used to make drinks called sumacade in North America. Seeds of true sumac (R. coriaria) are ground and added to flavour salads in the Mediterranean. Other species of locally important fruits and nuts are the little gooseberry tree (Buchanania arborescens), the charoli nut or almondettes (B. lanzan), wild mango (B. obovata), lapsi or Nepali hog plum (Choerospondias axillaris), chupadilla (Cyrtocarpa procera), dao (Dracontomelon dao), sao (D. duperreanum), kaffir plum (Harpephyllum caffrum), bastard marula (Lannea stuhlmannii),

bauno (Mangifera caesia), horse mango (M. foetida), bambangan (M. pajang), apple mango or kuwini (M. ×odorata = M. foetida × M. indica), raisin bush (Ozoroa insignis), desert date or busanguli (Searsia natalensis), ambarella (Spondias dulcis), yellow mombin (S. mombin), red mombin or jocote (S. purpurea), warmia berry (Tapirira guianensis) and African dao (Trichoscypha arborea). Anacardiaceae are infamous for their allergenic properties. Many species can cause contact dermatitis in sensitive people (including cashews, mangoes and rose pepper), but the most problematic are poison oak (Toxicodendron diversilobum), poison ivy (T. radicans, T. rydbergii), poison sumac (T. vernix) and related species abundant in North America, although there are many other species with allergenic compounds in other parts of the world. The allergens are catechols, resorcinols and other phenolic compounds. Resins of Toxicodendron vernicifluum, T. succedaneum and some species of Gluta are used in East Asia as a lacquer to decorate wooden art, picture frames, jewellery boxes and furniture. This increases the durability and water and heat resistance but also can cause allergic reactions. The galls created by aphids on Rhus chinensis are used as a blue dye for silk in China and Japan. The fruits of R. copallina and R. glabra produce a red dye, and the roots of the latter make a yellow dye. Many species are large trees that make good timber, especially Astronium

fraxinifolium, Myracrodruon urundeuva and Schinopsis species, which have commercial value. Some species are commonly cultivated as ornamentals, especially Cotinus coggygria, Harpephyllum caffrum, Pistacia chinensis, Rhodosphaera rhodanthema, Rhus typhina, Schinus molle and Searsia lancea.

Xanthoceras sorbifolium, Royal Botanic Gardens, Kew, UK [272]

Aesculus indica, Royal Botanic Gardens, Kew, UK [272]

Acer shirasawanum ‘Aconitifolium’, private garden, Kingston upon Thames, Surrey, UK [272]

Etymology: Anacardium is the Latinised form of Ancient Greek ανακαρδιών (anacardion), a mulberry twig, originally from Sanskrit vranakart, to make a wound. This alludes to the eating of an astringent fresh ‘cashew apple’ being dangerous for the mouth. It has also been suggested that the word is composed of Greek ανά (ana), to return, and καρδιά (kardia), a heart, in reference to the shape of the fruit, but this is a less likely derivation.

272. SAPINDACEAE Maple family

This is a variable family including trees, shrubs, woody and herbaceous climbers and perennial herbs. Leaves are usually alternate

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or opposite and simple to pinnately, ternately or palmately compound (up to tripinnate and triternate, or a combination of divisions), the basal leaflets sometimes reduced and covering the stem to resemble stipules (true stipules are only present in climbing species), the terminal leaflet reduced in most shrubby and tree-like species with pinnate leaves, hence the leaves appearing evenly pinnate. Leaflets have pinnate venation and entire, crenate, serrate or toothed margins. In simple leaves, venation may be palmate (e.g. many Acer) or pinnate. Inflorescences are axillary, terminal or cauliflorous thyrsoid panicles, racemes, spikes or fascicles or the flowers solitary. Coiling tendrils are sometimes formed in the inflorescences, replacing a pair of cincinni in the thyrse, often in the tip of the inflorescence. Flowers are pentamerous and actinomorphic or tetramerous and zygomorphic and bisexual or unisexual through functional reduction. The four or five petals are free or fused just at the base, the four or five (rarely absent) petals are usually coloured (often white or yellow) and usually bear a variously shaped, often petal-like appendage. These appendages can be basally fused, simple, forked or hooded structures that sometimes are mere extensions of the petal margins but frequently attached Dodonaea angustifolia, New South Wales, Australia [272]

Cardiospermum galapageium, Galápagos Islands [272]

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elsewhere on the petal and concealing the lobed, cup- or dish-shaped nectary disk that surrounds the stamens. The five to eight (sometimes three, four or up to 30) stamens have free or basally fused filaments and dorsifixed or basifixed anthers that open on the inside by lengthwise slits. Female flowers have sterile stamens (staminodes). The superior ovary is composed of usually three, sometimes fewer or more (up to eight) fused carpels, topped with a terminal (rarely gynobasic in Deinbiollia) two- or threebranched style or a two- or three-lobed stigma, which is enlarged into wings in Acer. Pistils are reduced and not functional (pistillodial) in male flowers. Fruits are septifragal or loculicidal capsules, schizocarps with winged parts, berries or (rarely) drupes. Seeds often have an aril and are usually horsechestnutor conker-like with a glossy skin and a large scar. Distribution: This is a mainly tropical and subtropical family, found nearly worldwide, and extending into temperate and even (hemi-) boreal zones. Phylogeny and evolution: Fossils that can be assigned to Sapindaceae are known from the

Koelreuteria paniculata, Royal Botanic Gardens, Kew, UK [272]

Late Cretaceous, corresponding to a molecular age estimate of 98–116 million years. Several pollen types attributed to Sapindaceae are widespread in the fossil record, including the excitingly named Wehrwolfea, described from the Eocene of Canada. The subfamilies of Sapindaceae spread in the Mid Cretaceous from Laurasia, with important diversifications in Southeast Asia. New Caledonia, which has a high diversity of Sapindaceae, has been reached by this family at least ten times from various distant parts of the world including the Mascarenes (Cossinia). The maples (Acer) separated from their closest relative (Dipteronia) c. 78 million years ago. Due to the position of Xanthoceras sorbifolium as sister to the rest of the family, the former families Aceraceae and Hippocastanaceae are included in Sapindaceae s.l., both united in subfamily Hippocastanoideae. Xanthoceras displays a mixture of characters typical of both Sapindoideae and Aceroideae and has been placed in its own family (Xanthocerataceae) to maintain Aceraceae and Hippocastanaceae, but this is an unneccesary inf lation of family names in a clade that shares many morphological similarities. Allosanthus is a synonym of Thouinia, Hypseloderma is now part of Camptolepis, Diplopeltis huegelii, City Beach, Western Australia [272]

SAPINDALES

EUDICOTS

Sapindus mukurossii, Chen Shan Botanical Garden, Shanghai [272]

Nephelium lappaceum, Singapore (KH) [272]

Neotina and Tinopsis are merged with Tina and Otonephelium may be a synonym of Dimocarpus but is tentatively accepted here. Chonopetalum from Equatorial Guinea is insufficiently known to place it in any subfamily, and Hirania from Somalia is of uncertain affinity because it differs from Sapindaceae in several technical characters. However, no placement is certain, but scanty molecular evidence does suggest Hirania may belong here. Further study of these genera is needed.

Castanospora (1), Chouxia (6), Chytranthus (c. 27), Cnesmocarpon (4), Conchopetalum (2), Cubilia (1), Cupania (c. 50), Cupaniopsis (60), Deinbollia (c. 38), Delavaya (1), Diatenopteryx (2), Dictyoneura (3), Dilodendron (3), Dimocarpus (6), Diploglottis (12), Elattostachys (c. 20), Eriocoelum (c. 10), Erythrophysa (5), Erythrophysopsis (1), Gereaua (1), Glenniea (8), Gloeocarpus (1), Gongrodiscus (3), Gongrospermum (1), Guindilia (3), Guioa (c. 64), Haplocoelopsis (1), Haplocoelum (6), Hornea (1), Houssayanthus (5), Jagera (2), Koelreuteria (4), Laccodiscus (6), Lecaniodiscus (2), Lepiderema (8), Lepidopetalum (6), Lepisanthes (c. 24), Litchi (1), Lophostigma (2), Lychnodiscus (7), Macphersonia (8), Matayba (c. 50), Melicoccus (10), Mischarytera (3), Mischocarpus (c. 15), Molinaea (10), Namataea (1), Nephelium (c. 16), Otonephelium (1), Pancovia (c. 11), Pappea (1–4), Paranephelium (4), Paullinia (c. 190), Pavieasia (1), Pentascyphus (1), Phyllotrichum (1), Placodiscus (10), Plagioscyphus (10), Podonephelium (4), Pometia (2), Porocystis (2), Pseudima (1), Pseudopancovia (1), Pseudopteris (3), Radlkofera (1), Rhysotoechia (c. 14), Sapindus (c. 10), Sarcopteryx (13), Sarcotoechia (11), Schleichera (1), Scyphonychium (1), Serjania (c. 230), Sinoradlkofera (1), Sisyrolepis (1), Smelophyllum (1), Stadmania (6), Stocksia (1), Storthocalyx (4), Synima (2), Talisia (52), Thinouia (c. 12), Thouinia (c. 30), Thouinidium (6), Tina (19),

Genera and species: Sapindaceae include c. 140 genera and about 1,860 species in four subfamilies: Xanthoceratoideae (1 genus) – Xanthoceras sorbifolium; Hippocastanoideae (5 genera) – Acer (126), Aesculus (13), Billia (2), Dipteronia (1) and Handeliodendron (1); Dodonaeoideae (22 genera) – Arfeuillea (1), Averrhoidium (4), Cossinia (3), Diplokeleba (2), Diplopeltis (5), Distichostemon (6), Dodonaea (67), Doratoxylon (6), Euchorium (1), Euphorianthus (1), Eurycorymbus (1), Exothea (3), Filicium (4), Ganophyllum (2), Harpullia (26), Hippobromus (1), Hypelate (1), Llagunoa (4), Loxodiscus (1), Magonia (1), Majidea (4) and Zanha (3); Sapindoideae (111 genera) – Alectryon (c. 25), Allophylus (c. 250), Amesiodendron (1), Aporrhiza (c. 5), Arytera (c. 28), Atalaya (12), Athyana (1), Beguea (1), Bizonula (1), Blighia (3), Blighiopsis (1), Blomia (1), Bridgesia (1), Camptolepis (4), Cardiospermum (c. 15),

Hirania rosea, holotype, collected by Kuchar (no 17237) in Somalia (Herbarium Royal Botanic Gardens, Kew, UK) [272]

Toechima (8), Toulicia (c. 12), Trigonachras (8), Tripterodendron (1), Tristira (1), Tristiropsis (3), Tsingya (1), Ungnadia (1), Urvillea (c. 15), Vouarana (2), Xerospermum (2) and Zollingeria (4). Unplaced – Chonopetalum (1) and Hirania (1). Uses: A number of important fruit crops are found in Sapindaceae. These include the longan (Dimocarpus longan), lychee (Litchi chinensis), keneep or mamoncillo (Melicoccus bijugatus), rambutan (Nephelium lappaceum), pulasan (N. ramboutan-ake), jacket plum (Pappea capensis), pitomba (Talisia esculenta), guaya (T. oliviformis) and velvetfruited zanha (Zanha suaveolens). Some, like lychee and rambutan, are grown on a large commercial scale and are frequently found on fruit stalls and in supermarkets. Others are enjoyed more locally, although they may be canned and then found in exotic grocery stores. Arillodes of Blighia sapida are poisonous when unripe, causing blood sugar lows that can result in coma or death. It is nutritous when ripe, cooked and prepared as ackee, a staple food in Jamaica, but it is surprisingly not consumed in its native Africa. Seeds of Blighia must have reached Jamaica through the slave trade, and the species was discovered there by William Bligh, captain of the Bounty, infamous for the mutiny. He brought material of ackee back to England, where it was named in his honour. Plants of the World

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SAPINDALES The emblem of Canada, the sugar maple (Acer saccharum), is of commercial importance. The sap of this and other maples (especially also red maple, A. rubrum, and black maple, A. nigrum) is drained and boiled down into a sweet liquid known as maple syrup, a delight on pancakes, waffles, cakes, porridge etc. Maple species store starch in their trunks during winter, which is converted to sugar when the sap starts flowing in spring, and the trees are bored and tapped, a custom of native Americans adopted by European settlers. Currently a vast proportion of maple syrup is produced in Quebec. The unique flavour is highly valued, but its chemistry is not fully understood. Seeds of Brazilian Paullinia cupana are the source of caffeine-rich stimulant guaraná, now commonly used in energy and soft drinks. Other species of Paullinia are also used for the high caffeine content of their seeds. Paullinia pinnata is used as a dart or arrow poison. The fruit pulp (soapberry) and the seeds (soapnut) of Sapindus species contain saponins and have been used by native peoples in the Americas and Asia to wash the body and clothes for millennia. Nuts of Asian S. mukorossi are often used in Ayurvedic cleansers, whereas commercially the nuts of American Sapindus saponaria are more commonly on offer in commercial cosmetics and detergents from bio-dynamic stores. Its reputed spermacidal effect has proven not to be sufficient for it to be used as a commercial contraceptive. Oils are pressed from the seeds of Pappea capensis and the kusum tree (Schleichera oleosa). Pometia pinnata is used for firewood in the Pacific islands. A number of tree species produce good timber, and especially Euphorianthus, Harpullia, Schleichera and Acer are frequently used for construction. Ornamentals are numerous and especially members of the genera Acer (maple), Aesculus (horse chestnut), Arfeuillea (hop-tree), Allophylus (titberry), Blighia (ackee), Cardiospermum (love-in-a-puff), Dipteronia (money maple), Diplopeltis (pepperf lower), Dodonaea (hopbush), Filicium (ferntree), Harpullia (tulipwood), Koelreuteria (goldenrain tree), Sapindus

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(soapberry) and Xanthoceras (yellowhorn) are frequently cultivated in gardens. There are hundreds of Acer cultivars, especially in the group of Japanese maples (A. palmatum, A. shirasawanum), which are also popular in East Asia as subjects for bonsai. Etymology: Sapindus is composed of Latin sapo, soap, and indicus, from India.

273. RUTACEAE Citrus family

These trees, shrubs and perennial herbs are sometimes spiny or scandent and usually aromatic due to (often translucent) glands that contain volatile oils in many parts and oil cells in the plant tissue (parenchyma). These glands are absent in some Cneoroideae. Leaves are alternate, opposite or whorled without stipules. Blades are simple or variously compound, often palmately so, sometimes with a single leaflet (unifoliolate) and often with a winged rachis. Venation is pinnate, and blade margins are entire, crenate or toothed. Inflorescences are axillary or terminal racemes, thyrses, panicles, spikes, bunches, heads, umbels or cincinni, or flowers are solitary. The usually actinomorphic or somewhat zygomorphic flowers are bisexual or unisexual. The three, four or five sepals are free or completely fused. Petals are equal in number to sepals and free or fused for part of their length. Stamens are twice as many as petals, rarely more numerous, and sometimes a whorl of stamens is reduced to staminodes or absent. Filaments are free or fused partially into a corona-like tube, with basifixed or dorsifixed anthers that open sideways or inside via lengthwise slits. The connective is often glandular at the tip. A nectariferous disk is usually present at the base of the stamens. The superior ovary

is composed of one to five (to many) carpels that are nearly free or completely fused, often on a short stalk (gynophore). Styles are free or fused, the stigmas often joined, even in species with free carpels. Fruits are variable and can be in a cluster of one to five free or apically fused follicles, drupes (sometimes winged) or samaras, or in species with fused carpels becoming a septicidal capsule or berry, sometimes with a hard shell and the seeds often ejected. In some special cases, the fruit is a sarcocarp or hesperidium (in Citrus, an allusion to the mythological golden apples of the Hesperides), in which the berry is composed of a leathery outer part (exocarp) covered with oil glands, a spongy thick middle part (mesocarp) and an inner membranaceous part (endocarp) with juicy, thin-stalked, pulp vesicles (modified hairs) among which the seeds are embedded. Distribution: Rutaceae are a nearly cosmopolitan family but are most diverse in the tropics and subtropics, with a few taxa extending into temperate North and South America, Central Europe, Central and boreal Asia and New Zealand. Citrus has one of its centres of diversity in Australia. Phylogeny and evolution: A number of early Tertiary fossils are known from genera that are now restricted to East Asia, and fossil citrus relatives (Citrophyllum) are known from the Cretaceous and Eocene of North America, the Palaeocene and Eocene of China and Italy and the Oligocene of the Caucasus. Rutaceae are most closely related to Simaroubaceae and Meliaceae in Sapindales. Two of the subfamilies (Aurantioideae and Rutoideae) are closely linked morphologically and chemically, but the third subfamily Cneoroideae were often considered a separate family, Cneoraceae, allied to Rutaceae. However, the single genus included in Cneoracaeae, Cneorum, was found to be related to several genera of Spathelioideae (Rutaceae) in molecular studies, which also showed that they are sister to Aurantioideae/ Rutoideae. Also included here were the genera previously recognised as Ptaeroxylaceae and Harrisoniaceae; these were included

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in the family because they share some morphological characters (including oil cells in their tissues) and some chemical traits with Rutoideae. They do stand apart due to some differences in chemistry, which are more similar to that of Simaroubaceae. The overall most satisfactory treatment of these is as a subfamily of Rutaceae. Molecular and morphological studies have resulted in the recircumscription of a number of genera. Several new genera were proposed, but many of these genera have been included in others treated in a broader sense, for instance Citrus now includes the former segregates Clymenia, Eremocitrus, Feroniella, Fortunella, Microcitrus, Oxanthera, Pleurocitrus and Poncirus, names that are still sometimes used in the horticultural trade. Vepris was expanded to include Araliopsis, Diphasia, Diphasiopsis, Humblotiodendron, Oricia, Oriciopsis, Teclea, Tecleopsis and Toddaliopsis. Severinia is tentatively placed as a synonym of Atalantia. Monnieria has some nomenclatural confusion and various spellings (Moniera, Monniera) but is now considered a synonym of Ertela. Achuaria is a synonym of Raputia. Boninia, Evodiella, Platydesma and Tractocopevodia are part of an expanded Melicope. Cyanothamnus is now a section of Boronia. Echinocitrus is included in Triphasia. Kodalyodendron belongs to Amyris. Taravalia is now included in Ptelea. Urocarpus belongs to Asterolasia. Genera and species: Rutaceae include 161 genera and c. 2,070 species in three subfamilies: Cneoroideae (7 genera) – Bottegoa (1), Cedrelopsis (8), Cneorum (2), Dictyoloma (1), Harrisonia (4), Ptaeroxylon (1) and Spathelia (c. 18); Aurantioideae (26 genera) – Aegle (3), Aeglopsis (5), Afraegle (4), Atalantia (17), Balsamocitrus (2), Bergera (5), Burkillanthus (1), Citropsis (8), Citrus (c. 30 wild species and some 145 apomictic and cultivated ‘species’), Clausena (15), Glycosmis (c. 50), Limnocitrus (1), Limonia (1), Luvunga (10), Merope (1), Merrillia (1), Micromelum (10), Monanthocitrus (4), Murraya (5), Naringi (2), Pamburus (1), Paramignya (c. 12), Pleiospermium (5), Swinglea (1), Triphasia (3) and Wenzelia (9); Rutoideae (122 genera) – Acmadenia (32),

Cneorum tricoccon, Helsinki Botanical Garden, Finland [273]

Acradenia (2), Acronychia (48), Adenandra (18), Adiscanthus (1), Agathosma (>150), Almeidea (5), Amyris (c. 40), Andreadoxa (1), Angostura (8), Apocaulon (1), Asterolasia (18), Balfourodendron (2), Boenninghausenia (1), Boronella (6), Boronia (148), Bosistoa (4), Bouchardatia (1), Brombya (2), Calodendrum (2), Casimiroa (c. 10), Chloroxylon (3), Choisya (6), Chorilaena (1), Cneoridium (1), Coatesia (1), Coleonema (8), Comptonella (8), Conchocarpus (c. 45), Coombea (1), Correa (11), Crossosperma (2), Crowea (3), Decagonocarpus (2), Decatropsis (2), Decazyx (2), Dendrosma (1), Desmotes (1), Dictamnus (1), Dinosperma (4), Diosma (28), Diplolaena (15), Drummondita (9), Dutailliopsis (1), Dutaillyea (2), Empleurum (2), Eriostemon (2), Ertela (2), Erythrochiton (7), Esenbeckia (30), Euchaetis (23), Euodia (7), Euxylophora (1), Fagaropsis (4), Flindersia (17), Galipea (15), Geijera (6), Geleznowia (1), Halfordia (1), Haplophyllum (c. 66), Helietta (8), Hortia (10), Ivodea (10), Leionema (27), Leptothyrsa (1), Lubaria (1), Lunasia (1), Maclurodendron (6), Macrostylis (10), Medicosma (25), Megastigma (2), Melicope (238), Metrodorea (5), Microcybe (4), Muiriantha (1), Myrtopsis (9), Naudinia (1), Nematolepis (7), Neobyrnesia (1), Neoraputia (6), Neoschmidea (2), Nycticalanthus (1), Orixa (1), Peltostigma (2), Pentaceras (1), Perryodendron (1), Phebalium

(28), Phellodendron (2), Philotheca (53), Phyllosma (2), Picrella (3), Pilocarpus (c. 17), Pitavia (1), Pitaviaster (1), Plethadenia (2), Polyaster (1), Psilopeganum (1), Ptelea (3), Raputia (11), Raputiarana (2), Rauia (c. 10), Raulinoa (1), Ravenia (11), Raveniopsis (19), Rhadinothamnus (3), Ruta (7), Rutaneblina (1), Sarcomelicope (9), Sheilanthera (1), Sigmatanthus (1), Skimmia (4), Spiranthera (4), Stauranthus (1), Tetractomia (6), Tetradium (9), Thamnosma (8), Ticorea (5), Toddalia (1), Toxosiphon (4), Vepris (c. 80), Zanthoxylum (c. 225) and Zieria (c. 60). Uses: Economically, the most important genus of the family is Citrus. Citrus growing is the most important fruit industry in warm countries, contributing significantly to the economy of many countries including Spain, Portugal, South Africa, Israel, Australia, USA (California and Florida) and Brazil. This industry is more than fruit and juice; citrus trees produce many products, such as perfumes, essential oils, spices, preservatives, marmalades, curds, liqueurs, vitamin C, sun-tan oil etc. Five millennia ago, citrus fruits were already cultivated and eaten in China. The first citrus fruit to reach Europe was the citron (C. medica), which was originally cultivated for the fragrance of its fruit and flowers, popular in the perfume industry,

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Murraya paniculata, Réunion [273]

Citrus ×sinensis, Sicily, Italy [273]

Choisya ternata, private garden, Hengelo, the Netherlands [273]

and giving towns like Orange in France their names. Citrons had probably been brought from India by Alexander the Great and soon became important in Jewish traditions, in which the cultivar ‘Etrog’ is used for a ritual during the Feast of Tabernacles. The Romans later introduced lemons and Seville oranges, to which the Arabs added lime and pomelo. Portuguese traders introduced the mandarin and sweet orange from China in the early 19th century, and the grapefruit is an accidental hybrid that arose in the Caribbean. During the Renaissance, Citrus became horticulturally important and fashionable in colder countries, where special houses with south-facing windows were built to overwinter citrons, bitter oranges, lemons, kumquats, limes and others in so-called ‘orangeries’, which from the early 17th century remained the way of protecting plants against winter weather until they were replaced by glass and iron greenhouses in the 1840s. The taxonomy of Citrus is extremely difficult due to frequent hybridisation, backcrossing, apomictic lineages and multiple embryos (sexual and asexual) in a seed. We make an attempt here to list the most commonly encountered kinds, but there are more, and new cultivars are likely to appear in supermarkets at regular intervals. Finger lime (Citrus australasica) is a species from northern Australia with a fruit that contains

small vesicles, now dubbed ‘lime caviar’, a popular ‘bushfood’ used in fashionable cuisine. A marmalade or pickle can also be made from the fruit, and the peel can be dried to make a pleasant spice. The finger lime has been crossed with calamondin to form the sunrise lime (C. ×olivieri; C. australasica × C. ×microcarpa), a common ‘bush tucker’ crop in Australia. Australian lime (C. australis) is another wild lime from northeastern Australia, sometimes valued as a bushfruit, and this has been crossed with the finger lime to create Sydney hybrid limes (C. ×virgata; C. australasica × C. australis). Dessert kumquat (C. glauca) is cold- and drought-tolerant, and its fruit can be used in drinks and preserves, usually only locally in the interior of Australia. Kaffir lime (C. hystrix) is commonly used in Southeast Asian cuisine, the rind in curry paste, and fruits can be candied. Especially the leaves are a popular ingredient in many traditional dishes of the region. The zest of this species is used in ‘arranged’ rums in Martinique, La Réunion and Madagascar. The acidic juice makes a good cleanser, and the oils are an effective leech repellent, which is why it is also called the leech lime. Citrus cavaleriei or Ichang papeda, is a native species of western China, which is famed for its frost hardiness and has yielded locally important crosses, such as the Ichang lemon (C. cavaleriei × C. maxima), the

yuzu (C. ×junos; C. cavaleriei × C. reticulata) and kabosu (“C. sphaerocarpa”). New summer orange or hyuganatsu (C. ×tamurana; C. ×junos × C. maxima) is another orangelike citrus fruit from Japan, known for its hardiness, light yellow skin and juicy, sweet flesh. Kumquat (C. japonica) when hybridised with a mandarin results in the calamondin (C. ×microcarpa; C. reticulata × C. japonica), which is commonly grown as ornamental pot plants because it is highly fructiferous. The kumquat is also crossed with the orange resulting in the orangequat (C. ×aurantium × C. japonica), the limequat (C. ×floridana; C. ×aurantiifolia × C. japonica) and the complex cold-hardy citrangequat (C. ×georgiana; C. ×insitorum × C. japonica). Pomelo (C. maxima) is a tropical species that is grown for its melon-sized fruits that have a good flavour. It is involved in a number of hybrids, including the orange and grapefruit. Hybrids of pomelo and mandarin give oranges and grapefruits. Original hybrids with more features of C. maxima (sour orange group) give Seville and bitter oranges (C. ×aurantium; C. maxima × C. reticulata) which are used for marmalade and candied peel or in liqueurs like Blue Curaçao or Cointreau. Sweet oranges and blood oranges (C. ×sinensis; C. maxima × C. reticulata) are orginal hybrids with more features of C. reticulata (sweet orange group), which are popular as a hand fruit or for juice.

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Myrtle-leaved orange (C. ×aurantium var. myrtifolia) is a selection of the orange with small fruit and leaves, popular as a decorative potplant. The grapefruit group (C. ×paradisi; C. ×aurantium × C. maxima) includes backcrosses between an orange and pomelo made in Barbados in the 18th century. Many cultivars are now known, and the fruit is Philotheca spicata, Mt Benia, Western Australia

[273]

popular for its juice and fresh for breakfast or dessert. Sweetie or oroblanco (C. maxima × C. ×paradisi) and tangors or ortaniques (C. ×nobilis; C. reticulata × C. ×paradisi) are further backcrosses with one parent. Tangors have been crossed with mandarin again to produce the tangelo (C. ×tangelo) and ugli fruit, among others. Many further

backcrosses with C. reticulata are now sold as the ‘mandarines’ of commerce. Bergamot orange (C. ×bergamia; C. medica × C. ×aurantium) has a fragrant fruit from which bergamot oil is extracted as a perfume (of Eau de Cologne fame) and flavouring (e.g. for Earl Grey tea). Key lime (C. ×aurantiifolia; C. maxima × C. medica/C. hystrix) is a popular

Zanthoxylum simulans, Jardin des Plantes, Paris

Geleznowia verrucosa, near Geraldton, Western Australia [273]

[273]

Correa reflexa, Kangaroo Island form, Australian National Botanic Garden, Canberra [273]

Ruta chalepensis, Sicily, Italy [273]

Dictamnus albus, Helsinki Botanical Garden, Finland [273]

Diplolaena ferruginea, Mt Benia, Western Australia [273] Halfordia kendac, New Caledonia [273]

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SAPINDALES fruit for spice and to add additional flavour to a gin and tonic. It is especially popular for key lime pies. Persian or Tahiti lime (C. ×latifolia; C. ×aurantiifolia × C. ×limon) is a seedless hybrid, which is commonly used as a replacement for key lime. More complex hybrids are still being made to obtain new fruit crops such as the seedless limelo (C. ×aurantiifolia × C. ×limon). Lemons (C. ×limon; C. medica × C. ×aurantium) have been in cultivation for such a long time that their exact parentage is unknown, although citron (C. medica) is a certain ancestor. They are used for juice, which is commonly added to food as a preservative and flavouring. It is also a key ingredient of lemonade and many other drinks, some alcoholic like the liqueur limoncello. Rough lemons (C. ×taitensis; C. medica ×C. reticulata) are often used as a stock for grafting. A cultivar ‘Otaheite’ is used as an ornamental potplant, popular at Christmas. Meyer lemon (C. ×meyeri; probably a cross between C. ×limon and C. reticulata) is sweeter than the common lemon and a popular fruit in Californian cuisine. Sweet lemon (C. limetta; possibly a cultivar of C. ×aurantium) has a sweetish acidic taste and is cultivated in the Mediterranean. Ponderosa lemon (C. ×limon × C. medica) is a backcross with citron with the same flavour as a lemon but much larger size. Buddha’s hand (C. medica ‘Sarcodactyla’) is a many-fingered form of the citron, and is normally only grown as a curiosity. Mandarins (C. reticulata) and their cultivars, clementines (a possible backcross with C. ×aurantium), satsumas and tangerines are hand fruits that peel easily and have a sweet taste. Mandarins were introduced into Europe via Kew in 1805 and are now commonly grown around the world. Satsumas are hardy, possibly among the hardiest of edible citrus. Amanatsu (‘Natsudaidai’) is a cultivar of mandarin the size of a grapefruit (thus possibly a hybrid with C. maxima) which is popular in Japan for juice and ice-cream. Kiyomi (C. reticulata × C. ×aurantium) is a mandarin-orange cross and an easily peeled, sweet, seedless fruit. Poncirus or trifoliate orange (C. trifoliata) is a cold-hardy deciduous species and makes a good barrier hedge. It is often used as a rootstock for grafting. The

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fruit has little flesh, but the rind can be made into marmalade. This spiny temperate species has been crossed to get cold-hardy citrange (C. ×insitorum; C. trifoliata × C. ×aurantium), which makes a good, virus-resistant rootstock for grafted citrus. A good number of citrus species, hybrids and cultivars are now cultivated. Fresh fruit, juice and marmalade as well as orange blossom and citrus peel oil are now driving an immense industry. White zapote or matasano (Casimiroa edulis) has a tasty, sweet, creamy flesh, but when consumed in considerable amounts may induce sleep because it contains a glucoside called casimirosine that has a hypnotic and sedative effect. It is not related to other species of zapote (Ebenaceae, Sapotaceae). Triphasia trifolia also has edible fruit. Several species are used as spices due to their fragrant leaves or fruit. Garden rue or herb of grace, Ruta graveolens, has a strong aromatic scent and a bittersweet flavour. A popular herb in some Mediterranean cuisines, it should be used sparingly because it is mildly poisonous. It should also be avoided by pregnant women because it reportedly can act as an abortifacient, and the leaves can cause severe dermatitis upon touch in sensitive people. Sichuan pepper is a spice commonly used in East Asian cuisine, usually derived from Zanthoxylum bungeanum or Z. simulans. The seeds and husks are used whole or are ground into a fine powder, which has a slight lemony aroma and causes a tingling numbness in the mouth that sets the palate for hot spices such as ginger, star anise and chillies that are often combined with it. The Japanese pepper (Z. piperitum) is a similar spice frequently used in Japanese cuisine. The curry tree (Bergera koenigii, formerly Murraya koenigii) has aromatic leaves that are an essential ingredient of Indian and Sri Lankan curries. Its essential oil also has antifungal properties and is added to soaps and hair tonics and used in aromatherapy. The leaves of several Australian Rutaceae are also distilled for their essential oils, including brown boronia (Boronia megastigma), which is important in the perfume industry. Torchwood (Amyris species) is high in resin that can be used in lacquers, varnishes, perfumes, cosmetics,

soap and incense. The wood burns easily and is therefore often gathered for firewood. Few species are large enough to provide much timber, but some, such as balsam torchwood (Amyris balsamifera), Ceylon satinwood (Chloroxylon swietenia) and West Indian sandalwood (Zanthoxylum flavum) have hard, scented and easily workable wood. Other genera such as Flindersia, Euxylophora, Phellodendron, Tetradium and Zanthoxylum are locally used for timber. A number of genera are grown as garden ornamentals: Boronia, Choisya, Citrus, Cneorum, Correa, Dictamnus, Diosma, Murraya, Phellodendron, Ptelea, Ruta, Skimmia, Tetradium, Toddalia, Triphasia, Zanthoxylum and many others locally. Some Boronia are commonly grown for the cut-flower industry. Etymology: Ruta is the classical Latin name of garden rue (Ruta graveolens). The origin of the word is uncertain, but it is probably from the ancient Pelopponesian rhyte, a bitter herb.

274. SIMAROUBACEAE Tree-of-heaven family

Leitneria floridana in fruit, Missouri Botanical Garden, USA (CD) [274]

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This is a family of trees and shrubs that usually have bitter-tasting bark. Leaves are alternate and spirally arranged without stipules (except in Picrasma, which has triangular pseudostipules at the base of the petiole). Blades are unevenly pinnate, rarely trifoliate or unifoliolate (simple), with entire, toothed, serrate or lobed leaflet margins. Leaflets have pinnate or reticulate venation and sometimes flat or sunken glands. Inflorescences are terminal or axillary thyrses that sometimes appear raceme-like, catkinlike or umbel-like or are clustered in the leaf axils. Flowers are bisexual or unisexual and actinomorphic, usually subtended by bracts (which become large in Leitneria). The four or five fused sepals are sometimes split unevenly and may bear glands. The four or five petals are free, but both sepals and petals are absent in Leitneria. Stamens are the same number

as petals or twice as many (four, five, eight or ten, rarely to 18), with free filaments that usually have a hairy appendage at the base and dorsifixed, basifixed or versatile anthers that open by two lengthwise slits. A disk is usually present, and sometimes the ovary is stalked (gynophore) or male and female parts stalked (androgynophore). The superior ovary is composed of (one to) two to five carpels that are free or fused at the base, the styles free or fused into a single style, the stigmas spreading like a star or capitate and lobed. The fruit is composed of one to five separate fruitlets that are drupe-like or samara-like. Distribution: Simaroubaceae are a mainly pantropical family extending into the temperate zones of southeastern North America, the Middle East and the Asian Far East.

Simarouba glauca in fruit, Fairchild Tropical Botanical Garden, Florida, USA (CD) [274]

Quassia amara, Royal Botanic Gardens, Kew, UK [274]

Phylogeny and evolution: Simaroubaceae have been dated to first diversify in the Late Cretaceous (Maastrichtian), c. 65 million years ago, but the greatest diversification occurred during the Caenozoic. The family has an excellent fossil record, many are found in temperate areas where they do not currently occur, and to explain the fossil and current-day distribution, the biogeographical history is complex involving many dispersals and extinctions. The fossil record of Leitneria extends to the Mid Oligocene (30–32 million years ago). Leitneria was often placed in its own family of uncertain association and due to its lack of perianth it had been associated with Hamamelidales, but biochemical studies suggested an affiliation with Simaroubaceae in Sapindales, which has now been confirmed by molecular analyses. Castela, Holacantha

Ailanthus altissima, street tree in Bucharest, Romania [274]

Soulamea terminalioides, Seychelles [274]

Picrasma quassioides, Royal Botanic Gardens, Kew, UK [274]

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SAPINDALES and Picrasma are sister to the rest. Pleiokirkia, previously placed here, is now a synonym of Kirkia (Kirkiaceae). Genera and species: Simaroubaceae include 22 genera and c. 108 species: Ailanthus (5), Amaroria (1), Brucea (7), Castela (12), Eurycoma (3), Gymnostemon (1), Hannoa (c. 6), Holacantha (2), Iridosma (1), Laumoniera (1), Leitneria (1), Nothospondias (1), Odyendea (1), Perriera (2), Picrasma (8), Picrolemma (2), Pierreodendron (2), Quassia (2), Samadera (6), Simaba (25), Simarouba (6) and Soulamea (13).

275. MELIACEAE Neem family

Etymology: Simarouba is a Latinised form of the local Galibi name for S. amara in French Guiana.

These are usually unisexual, sometimes bisexual trees, shrubs and shrublets, the stems sometimes pachycaul, and male trees sometimes produce bisexual flowers. The usually alternate, spirally arranged leaves, rarely opposite, lack stipules, but false stipules (formed from the basal pair of leaflets) are often present. Leaves are usually once pinnate, sometimes bipinnate (Melia), trifoliolate or unifoliolate, sometimes the blade terminated by a bud. The rachis is rarely winged, and leaflets are somewhat oblique at the base and margins usually entire, sometimes lobed or serrate. Venation is pinnate. Inflorescences are axillary or cauliflorous, rarely epiphyllous (Chisocheton) thyrses, racemes or spikes, which are sometimes reduced to fascicles or solitary flowers. Actinomorphic flowers are usually unisexual or occasionally bisexual, and rudiments of the opposite sex are clearly present. There are three to six fused or free sepals that are usually cup-shaped or tubular, sometimes circumscissile when the flower opens. The three to seven (or up to 14)

Mahogany, Swietenia mahagoni, Yucatán, Mexico [275]

Azadirachta indica, Aurangabad, India [275]

Uses: Simaroubaceae contain quassinoids, which have antiviral, antimalarial and insecticidal properties and are used commonly to cure these and other ailments related to digestion in traditional medicine. Corkwood (Leitneria floridana) has one of the lightest woods known and is traditionally used for fishing-net floats. Several species are widely grown garden ornamentals, especially bitterwood (Quassia amara), paradise tree (Simarouba glauca) and, unfortunately, the tree-of-heaven (Ailanthus altissima), which is an aggressive invasive in temperate regions where it is not native. It was introduced to eastern North American in hopes of starting a silk industry based on the silk of the Ailanthus silkmoth, without success.

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petals are in a single whorl or (Chisocheton) spirally arranged. Petals are free or fused at the base of the staminal tube. The three to ten (or more) stamens are usually fused into a staminal tube that is lobed or not, but stamens are free in Cedrela and Toona. Anthers are usually basifixed, sessile on the staminal tube and opening by lengthwise slits. A nectary disk is formed around the ovary or absent. The staminal tube is sterile in female flowers. The superior ovary is composed of two to six (to 20) fused carpels, each forming a locule. The style is fused, single with a disc- or head-shaped stigma that is sessile on the ovary in some taxa. The fruit is a capsule, a berry (often dry and indehiscent), or, rarely, a drupe. Seeds are winged or embedded in a fleshy aril or sarcotesta that wholly or partly covers the seed. Distribution: This is a pantropical family, with a few temperate representatives in China and South Africa. Phylogeny and evolution: Meliaceae are closely related to Simaroubaceae and Rutaceae. It is estimated that the two subfamilies diverged c. 39 million years ago, although diversification of the family probably happened some 50 million years or so earlier, fossils of undoubted Meliaceae are known from the Upper Cretaceous. There is a good fossil record, which helps with the complicated biogeography of this family. This suggests an origin in Africa and subsequent dispersal to other continents, possibly through Turraea obtusa, Royal Botanic Gardens, Kew, UK [275]

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Trichilia havanensis, Royal Botanic Gardens, Kew, UK [275]

Cedrela odorata, capsules, Turbaco, Bolívar, Colombia [275]

Sandoricum koetjape, Singapore (WA) [275]

the boreotropics. Molecular studies have resulted in some generic realignments. Asian Trichilia species are now treated as Heynea. Pseudobersama is tentatively accepted here as separate from Trichilia because the species is morphologically distinct. Aglaia is paraphyletic with regard to Lansium and Reinwardtiodendron, which probably should be included. Megaphyllaea is a synonym of Chisocheton.

Uses: The most important fruit crop in this family is the langsat (Lansium parasiticum; synonym: L. domesticum), which is probably native to southern Thailand but is now cultivated pantropically. The fleshy aril around the bitter inedible seeds tastes somewhat like sweet grapefruit. Fruits of Sandoricum species are also edible, and santol (S. koetjape) has some cultivated forms with improved fruits. Fruits are also cooked, candied and made into marmalade. Seeds of both species may cause intestinal problems if swallowed. Some Aglaia species also have edible fruit but are only consumed locally. The reddish young leaves of the red toon (Toona sinensis) are commonly cooked and eaten as a vegetable in China. It has an onion-like odour but a floral flavour. Neem (Azadirachta indica) is tolerant of droughts and therefore widely cultivated in warm countries as shade and wind-breaks and for its oil-rich seeds. Neem oil is used as fuel and in soaps, toothpaste and lotions; its insecticidal properties are well known, and it can be used as a systemic pesticide on vegetables and ornamental plants (a promising alternative to neonicotinoids). Neem oil has numerous medicinal applications. Andiroba seed oil (Carapa procera and C. guianensis) is similar to neem oil, light yellow in colour and extremely bitter. When subjected to a temperature below 25°C it solidifies, taking on a consistency like that of petroleum jelly. The wood is similar to mahogany and of high quality.

Meliaceae include some important timber species. True mahogany (Swietenia mahagoni) has been reduced in quality and quantity due to overexploitation and genetic erosion, and the species is now listed under CITES to restrict international trade. It is widely planted in Latin America, but many trees lack vigour, and it will be a while before stocks will have been replenished sufficiently for commercial harvesting. It is now widely replaced by Spanish mahogany (S. macrophylla), although this species may also be under threat of overexploitation. Another important timber is the fragrant wood of Spanish cedar (Cedrela odorata), which is often adulterated with C. fissilis of lesser quality. Other important timbers are chickrassy (Chukrasia tabularis), sapele (Entandrophagma cylindricum), sipo (E. utile), African mahogany (Khaya), Nigerian golden walnut (Lovoa trichilioides), Persian lilac or white cedar (Melia azedarach) and red cedar (Toona ciliata), and many other species produce valuable timbers of local use. Fruits of Melia azedarach have been used as beads, but these are toxic to humans. However, they can be used as an insecticide. The species is commonly planted as an ornamental in tropical and warm-temperate regions.

Genera and species: Meliaceae are a family of 51 genera and c. 600 species, in two subfamilies: Cedreloideae (14 genera) – Capuronianthus (2), Carapa (>2), Cedrela (c. 17), Chukrasia (1), Entandrophagma (11), Khaya (5), Lovoa (2), Neobeguea (3), Pseudocedrela (1), Schmardaea (1), Soymida (1), Swietenia (3), Toona (5) and Xylocarpus (3); Melioideae (37 genera) – Aglaia (c. 120), Anthocarapa (1), Aphanamixis (3), Astrotrichilia (12), Azadirachta (2), Cabralea (1), Calodecaryia (2), Chisocheton (53), Cipadessa (1), Dysoxylum (c. 80), Ekebergia (4), Guarea (c. 40), Heckeldora (6), Heynea (2), Humbertioturraea (10), Lansium (3), Lepidotrichilia (4), Malleastrum (23), Melia (3), Munronia (4), Naregamia (2), Nymania (1), Owenia (5), Pseudobersama (1), Pseudoclausenia (1), Pterorhachis (2), Quivisianthe (2), Reinwardtiodendron (7), Ruagea (5), Sandoricum (5), Sphaerosacme (1), Synoum (1), Trichilia (c. 70), Turraea (c. 60), Turraeanthus (3), Vavaea (4) and Walsura (16).

Etymology: Μέλια (Melia) is a classical Greek name for an ash tree (Fraxinus, Oleaceae), because some species yielded good honey, and the name was given to this tree because of the leaf similarity and presumably also because the tree can yield honey.

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EUDICOTS

HUERTEALES Families 267 to 270 comprise the unusual order Huerteales, which were previously not recognised; they are composed of a number of small families sharing few characters. They are sister to Brassicales+Malvales. Wood anatomy seemed the only character that united the families Dipentodontaceae (ex-Celastraceae), Gerrardinaceae (ex-Flacourtiaceae) and Tapisciaceae (ex-Staphyleaceae), until Petenaeaceae (ex-Elaeocarpaceae) were added, resulting in the loss of that last character. All have small flowers, dentate leaves and unilocular ovaries that grow into berries or drupes. Dipentodontaceae plus Tapisciaceae are sister to Gerrardinaceae plus Petenaeaceae. All families appear to be of a relictual nature.

276. PETENAEACEAE Petén-linden family

stamens are free, with nectary glands between the glabrous filaments. Anthers are dorsifixed and open with apical slits. The superior ovary is hairy and composed of four or five fused carpels. The fruit is a globose, lobed berry. Distribution: This family occurs locally in southern Mexico (mainly Chiapas), northern Guatemala and western Belize.

This is a family of small trees and large shrubs with palmately veined, cordate, minutely toothed leaves that resemble those of a linden (Tilia, Malvaceae). They have minute triangular stipules at the base of the petioles. Petioles, peduncles and pedicels are tinged pinkish red. Inflorescences are manyflowered villous-tomentose panicles. The bisexual, actinomorphic flowers lack petals, but the five sepals are white-hairy outside and the receptacle is adorned with long, showy, bright pink, multicellular hairs. The eight to 12

Petenaea cordata, Huehuetenango, Guatemala (MV) [276]

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Phylogeny and evolution: Described as a member of Elaeocarpaceae, Petenaea was later considered to belong to Tiliaceae (now Malvaceae), probably because of a superficial resemblance to members of that family. The genus was difficult to place because of its unusual combination of characters: simple and branched trichomes, cordate leaves with tiny stipules, petals replaced by (but not homologous with) moniliform hairs, receptacular glands between stamens, dorsifixed anthers dehiscing through a terminal slit that eventually opens down the full length of the anther and a large axile Petenaea cordata in fruit, Xhanil, Chiapas, Mexico (CD) [276]

placenta with numerous ovules and berries. Because of the minute stipules and a similar ovary structure, a possible relationship with Muntingiaceae was also suggested, but the lack of petals in Petenaea and the differences in wood anatomy between these made this unlikely. Molecular analyses indicate a distant relationship to other members of Huerteales, the closest family being African Gerrardinaceae. Few morphological characters support such a relationship. Genera and species: The single genus in this family is Petenaea with only one species, P. cordata, which can be locally common. Uses: The berries are edible and eaten as a treat by local children. Etymology: Petenaea is named for the Lake Petén Itza, in El Petén Department in Guatemala, where the type was collected. The name itself orginated from Nojpetén, capital of the Itza Maya Kingdom, now the city of Flores. Petenaea cordata, Huehuetenango, Guatemala (MV) [276]

HUERTEALES

EUDICOTS

Gerradina foliosa, fruit, Eastern Cape, South Africa (TD) [277]

277. GERRARDINACEAE Brown-ironwood family

This is a family of shrubs and small trees, often with pendent branches, sometimes somewhat scandent. They have simple, alternate, stipulate, petiolate, glandular toothed leaves and pinnate venation. Inflorescences are axillary, few-flowered, raceme-like cymes. The bisexual flowers are actinomorphic and minute. The five sepals are basally fused and persistent in fruit. The five fused petals are white and opposed by five stamens. Filaments are free, and anthers are basifixed and open by longitudinal slits. A cup-like, five-lobed disc is fused to the flower base (hypanthium). The superior ovary is composed of two fused carpels topped with a single style and a shallowly bilobed or capitate stigma. Fruits are red, pendent, dry or fleshy berries with up to four seeds each.

Gerradina foliosa, Eastern Cape, South Africa (TD) [277]

Distribution: A family with a patchy distribution in southern and southeastern Africa in forest margins, open hillsides and rocky outcrops in savanna and montane forest, in Malawi, Mozambique, Tanzania, Zimbabwe and South Africa. Phylogeny and evolution: Gerrardinaceae are sister to Petenaeaceae. They were previously placed in Flacourtiaceae, but the teeth are not salicoid; thus, their relationship with Salicaceae (in which Flacourtia is now placed) has been debated in the past. They share some wood-anatomical characters with other Huerteales but not with their closest relative Petenaeaceae. Genera and species: A single genus, Gerrardina, with two species: G. foliosa from South Africa with leathery leaves that are rounded to acute at the base and G. eylesiana from southeastern Africa (Tanzania, Malawi and Zimbabwe), which has soft cordate leaves. Etymology: Gerrardina is named for William Tyrer Gerrard (died 1866), an English naturalist who collected plants in South Africa and Madagascar in 1865–1866.

278. TAPISCIACEAE Silverpheasant-tree family

Tapisciaceae are evergreen and deciduous trees with fluorescing wood and alternate, petiolate leaves with soon-falling stipules. Blades are unevenly pinnate, with up to ten pairs of leaflets, the basal pairs sometimes trifoliate or pinnate (basally bipinnate). Leaflets have glands or stipels where they join the stem, venation is pinnate and margins are serrate to crenate. Inflorescences are axillary panicles with small male or bisexual, actinomorphic flowers. The five sepals are shortly basally fused and form a tube or cup with the hypanthium on which they are situated. The five petals are free and often hairy inside, white or yellow. The five stamens have filaments that are free and longer than the petals, with dorsifixed anthers that open

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EUDICOTS

petiolate. Stipules are small, distinct and soon deciduous. Blades are entire or serrulate with pinnate venation and often small domatia in the axils of larger veins on the lower side. Inflorescences are produced on the current year’s growth; they are axillary in shortened cymes in a pedunculate umbel or thyrses in a raceme or panicle. The actinomorphic flowers are small, up to 4 mm in diameter. Sepals and petals are (four or) five (to seven), free and more or less undifferentiated. Stamens are (four or) five inserted on the margin of a disk, the anthers face inward and open lengthwise. The superior ovary has two or basally four locules. The fruit is a drupe-like, tardily opening capsule with one arillate seed or a berry with two to four seeds. Tapiscia sinensis, private garden, Kingston upon Thames, Surrey, UK. [278]

Huertea glandulosa in fruit, Suiza Nueva, Oxapampa, Peru (CD) [278]

by lengthwise slits. A nectary disk is fivelobed in Huertea and absent in Tapiscia. The superior ovary is composed of two completely fused carpels that form a single locule with a single ovule. The fruits are drupes or berries that take more than a year to mature.

Eocene are known from Oregon, England and Germany.

Distribution: Tapisciaceae occur disjunctly in temperate East Asia (southern and southeastern China and northern Vietnam) and in tropical America along the Andes from Mesoamerica to Peru and the Greater Antilles.

Uses: The wood of cedro macho, Huertea glandulosa, is considered good for construction in Peru. Tapisciaceae have no growth rings.

Phylogeny and evolution: The genera of Tapisciaceae were previously placed in Staphyleaceae, to which they have a superficial resemblance. However, their placement has long been controversial, especially because they have no clear anatomical similarity with either genus of Staphyleaceae. They differ in wood anatomy, leaf arrangement and seed structure. Molecular studies have shown Staphyleaceae belong to Crossosomatales, and Tapisciaceae to a clade sister to the Malvales+Brassicales, now known as Huerteales. Tapisciaceae are characterised by alternate imparipinnate leaves with glands or stipels at the insertion point of the pinnae and small flowers in axillary panicles. Fossil fruits and seeds of Tapiscia from the

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Genera and species: Tapisciaceae include two genera with six species: Huertea (4) and Tapiscia (2).

Etymology: Tapiscia is an anagram of Pistacia (Anacardiaceae) because the original author saw some similarity to the latter.

Distribution: Perrottetia is distributed in Central and South America (Andean), southern China, Taiwan, Malesia, Macronesia, Melanesia, northeastern Australia and Hawaii. Dipentodon sinicus comprises two isolated populations in southern China and adjacent Vietnam. Phylogeny and evolution: Both Dipentodon and Perrottetia have often been treated in Celastraceae, but they were never associated with each other before molecular studies. However, morphological and molecular evidence indicates that they are closely related and should be placed in Huerteales, not Celastrales.

Shichi family

Genera and species: This small family has two genera and 20 species: Dipentodon sinicus and Perrottetia (19).

This family is composed of semi-evergreen and deciduous, unisexual and bisexual trees and shrubs. Leaves are alternate, simple and

Etymology: Dipentodon is composed of the Greek words δυο (dio), twice, πέντε (pente), five, and οδόντος (odontos), a tooth, in reference to the two whorls of five teeth on the perianth. The author (Dunn, 1911) provided the etymology as follows: “The name Dipentodon, proposed for it, refers to the most remarkable character possessed by the flowers in the exact similarity of the calyx teeth and petals (if I rightly call them so) and their insertion so nearly in one whorl that the appearance is given of a ten-toothed perianth”.

279. DIPENTODONTACEAE

MALVALES

EUDICOTS

Dipentodon sinicus in fruit, Yunnan, China (QY)

[279]

Dipentodon sinicus, Yunnan, China (QY) [279]

Perrottetia longistylis, University of California Botanical Garden, Berkeley, USA [279]

Perrottetia longistylis, University of California Botanical Garden, Berkeley, USA [279]

MALVALES Families 280 to 289 comprise the Malvales, an order that can typically be recognised by palmate venation (at least at the base of the blade), mucilage canals and fibrous bark (tough and strong in Thymelaeaceae); stellate or fascicled hairs are common. Seeds are also often hairy, to the extreme in cotton (Gossypium, Malvaceae). Malvales contain some 3.2% of eudicot diversity, and the order is estimated to have evolved c. 71–78 million years ago. Relationships of several families are well known, those at deeper levels remain obscure. Elaeocarpaceae, previously often associated with this order, are now placed in Oxalidales.

280. CYTINACEAE Rockrose-rape family

Cytinaceae are endophytic, parasitic herbs that lack chlorophyll and true roots. Infection starts inside the host roots with filamentous or plate-like structures that resemble a fungal mycelium. Leaves are reduced to spirally arranged scales surrounding the inflorescence stalk and the base of the flowers (bracts). Inflorescences are terminal spikes, reduced

to an umbellate structure or less defined cluster or the flowers solitary and terminating the inflorescence stalk. The unisexual or sometimes bisexual flowers are actinomorphic and usually subtended by two scale-like bracteoles. The four to nine fused sepals are brightly coloured and petal-like and form a tubular structure. Petals are absent. Nectaries are formed between the sepal lobes in cavities surrounded by papillae. Male flowers have eight (or up to 20) stamens fused into a column, the anthers united into a ring at the tip of this filament-column, sometimes with spurs formed from the connective. Anthers open by longitudinal slits. Female flowers lack stamens. The semi-inferior ovary is topped with a single style bearing a globose or capitate stigma. Fruits are fleshy berries that sometimes open irregularly. Many small seeds are surrounded by sweet, slimy pulp.

Distribution: Cytinaceae have a patchy distribution in Mexico, Mesoamerica, Mediterranean Europe and North Africa, Anatolia, southern Africa and Madagascar. Phylogeny and evolution: Cytinaceae were previously included in the now defunct order Raff lesiales (or as part of Rafflesiaceae) because in their vegetative stage these parasites grow as filamentous or plate-like endophytes inside the host. Inflorescences emerge from the host tissue, a similar life strategy to that in Rafflesiaceae (Malpighiales). Molecular studies have demonstrated that they are most closely related to Muntingiaceae in Malvales. Cytinaceae consist of two genera of a ch lor o phyl lou s holo p a r a sit e s , Bdallophytum from the New World, which parasitises Burseraceae (mainly Bursera)

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EUDICOTS

Cytinus ruber dissected flower, Sicily, Italy [280]

Cytinus hypocistis, Cabo de Gata, Almeria, Spain

Bdallophytum americanum, Cerro el Campanario, Jiqiuipilas, Chiapas, Mexico (CD) [280]

Muntingia calabura, Ecuador [281]

Dicraspidia donnellsmithii, Costa Rica (BH) [281]

and occasionally also Cochlospermum, Guazuma, Gyrocarpus, Ficus and Haematoxylum, and Cytinus in the Mediterranean, South Africa and Madagascar, which parasitises Cistaceae (mainly Cistus), Asteraceae (especially Helichrysum) and Hamamelidaceae (Dicoryphe).

Etymology: Κυτίνος (Kytinos) was used by Theophrastus and Pliny for the calyx of a pomegranate blossom, which bears some resemblance to this plant.

Muntingiaceae are a family of trees and shrubs covered with stellate hairs, especially on young shoots. The alternate petiolate leaves occur in a plane (distichous). The simple blades are unequally heart-shaped at the base with palmate venation and serrate margins. Stipules are filiform or leaf-like and peltate or reduced in one half. Inflorescences are formed above the leaf axes and few-flowered in clusters or the flowers are solitary. The actinomorphic flowers are bisexual and pedicellate. The usually five (sometimes four to seven) sepals are more or less fused at the base forming a saucer- or cup-shaped tube. There are as many petals as sepals (usually five), which are crumpled in bud and free, longer than the calyx with an irregular margin. The numerous stamens

(CD) [280]

Cytinus ruber on its host Cistus creticus (Cistaceae), Sicily, Italy [280]

Genera and species: Cytinaceae include two genera with ten species: Bdallophytum (2) and Cytinus (8). Uses: Young inf lorescences of Cytinus hypocistis and C. ruber are eaten locally as an asparagus substitute in the Mediterranean. They have also been used medicinally in the past. 386

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281. MUNTINGIACEAE Bajelly-tree family

MALVALES

EUDICOTS

have thin, free filaments with (near) basifixed anthers that open by longitudinal or apical slits. The superior to inferior ovary is composed of five fused carpels, with a short, thick style and thick, lobed stigma. The fruit is a juicy, sweet berry with small seeds. Distribution: This family is native to tropical America, from central Mexico through Mesoamerica and into Andean South America and also on Hispaniola and Puerto Rico. Muntingia calabura is cultivated and naturalised throughout the tropics. Phylogeny and evolution: Like Petenaea (n ow Pe t e n a e a c e a e , Hu e r t e a le s), Muntingia had formerly been placed in Elaeocarpaceae, Tiliaceae (= Malvaceae) and even Flacourtiaceae (= Salicaceae). Molecular studies have placed Muntingia and Dicraspidia in Malvales, close to Cytinaceae. Neotessmannia has only been collected once and is placed here based on morphological similarities. In Malvales, the clade of Muntingiaceae and Cytinaceae is not resolved, but it is clear they are not members of the now expanded Malvaceae.

Uses: Bajelly tree, calabura or jam tree (Muntingia calabura) has edible berries; they are rapidly growing trees, sometimes used for firewood. Etymology: Muntingia is named for Dutch botanist Abraham Munting (1626–1683), Professor of Botany at Groningen University, who succeeded his father as the director of the Hortus in Haren, which after his death was directed in turn by his son.

282. NEURADACEAE Pietsnot family

receptacle that enlarges in fruit and resembles an epicalyx sometimes bearing spines. The five, free, persistent sepals are sometimes spine-tipped, and the five petals are free. The ten stamens occur in two whorls, the inner whorl shorter than the outer whorl, with free, glabrous or hairy filaments that are basally broader. Anthers are dorsifixed and versatile, opening by lengthwise slits. The ovary is composed of a whorl of ten carpels sunken into the receptacle, each tipped with a style and a capitate stigma. Styles are persistent and harden in fruit, and the ovary may become asymmetrical due to uneven development of the carpels. The fruit is usually an indehiscent spiny disc-shaped capsule. Seeds germinate within the fruit. Distribution: This is a family of arid land in the subtropics of North Africa through Arabia, the Levant and Iran to India and South Africa and Namibia. They usually grow on sandy soil.

Genera and species: Muntingiaceae include three genera, each with a single species: Dicraspidia donnellsmithii, Muntingia calabura and Neotessmannia uniflora.

Neuradaceae are annual and perennial herbs with stems that are pressed to the ground and usually covered in woolly hairs. Leaves are alternate and petiolate, the blades simple, lobed or pinnatifid and pinnately or palmately veined with a toothed margin. Inflorescences are reduced to solitary, axillary flowers. The bisexual flowers are actinomorphic, with a

Phylogeny and evolution: Previously placed in Rosales with which they share floral anatomical characters, molecular analyses place Neuradaceae in Malvales. They differ from Rosales in pollen and seed structure and their mucilage canals, which gave them their name in Afrikaans “pietsnot”, referring to the thick, slimy sap of these plants.

Neurada procumbens, in fruit, Israel (GP) [282]

Neurada procumbens, flower, Israel (GP) [282]

Grielum grandiflorum, South Africa (FF) [282]

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MALVALES Genera and species: Neuradaceae include three genera with nine species: Grielum (5), Neurada (1) and Neuradopsis (3).

EUDICOTS

283. MALVACEAE Mallow family

Uses: Called sa’ada in Arabic, Neurada procumbens is occasionally eaten as a vegetable by Bedouins in North Africa and Arabia. The species may elevate blood pressure so care has to be taken when consuming this plant. It has recently become invasive in Australia. Etymology: Neurada is possibly derived from the Greek νεύρων (neuron), a nerve, an unclear reference. Νευρασ is also a name in Greek and it may be derived from that. Its etymology remains uncertain.

This is a diverse family of trees, shrubs and annual and perennial herbs, sometimes even climbers, but all have tufted or stellate hairs, sometimes mixed with simple hairs, glands or scales, rarely with prickles or spines. Their

Abroma angusta, Singapore (WA) [283] Melochia tomentosa, Tulum, Yucatán, Mexico [283]

Herrania balaensis, Royal Botanic Gardens, Kew, UK [283]

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Lasiopetalum lineare, Geraldton, Western Australia [283]

hairs can be irritating to the skin in some cases. Alternate, simple or palmately compound leaves have blades with palmate venation and entire to toothed, serrate, crenate, lobed or dissected margins. Petioles are usually thickened (pulvinate) at both ends and a stipule is present. However, this is sometimes reduced or lost early. Inflorescences are axillary, terminal, cauliflorous or opposite a leaf, and usually composed of cymes in various combinations, often bracteate. Flowers are bisexual or unisexual by abortion and usually actinomorphic (somewhat zygomorphic or asymmetrical in some Helicteroideae). Three or more sterile fused bracts (epicalyx) are often present below the flowers. The five sepals are free or fused and sometimes Theobroma cacao in fruit, Helsinki Botanical Garden, Finland [283]

Rulingia luteiflora, Australian National Botanic Garden, Canberra [283]

MALVALES

EUDICOTS

Corchorus olitorius, Aurangabad, India [283]

Corchorus neocaledonicus, New Caledonia [283]

separate unevenly, sometimes petal-like. If present, nectaries are composed of glandular hairs inside the sepals or rarely on petals. The five petals are free or fused at the base or with the stamen filaments, various in shape and size, sometimes reduced or lacking. The male and female organs are sessile or placed on a stalk (androgynophore). Five to >1,000 stamens are usually in groups or fascicles, with the filaments free or fused and then forming a stamen tube that surrounds the style. Anthers are basifixed or dorsifixed, the thecae opening by pores or locules. Sterile stamens (staminodes) are often present, usually fused with the filaments and sometimes petal-like. The superior ovary is usually composed of five carpels, but sometimes many more or fewer carpels are formed in a whorl. Carpels are free or fused and tipped with a simple or apically branched style, or the free carpels have free styles. Stigmas are fused or divided. Fruits are dehiscent or indehiscent capsules, sometimes splitting into many parts (mericarps) or the fruit winged. The capsules may be hairy inside or filled with a fleshy pulp. Seeds are often hairy or winged, sometimes with an aril. Distribution: Malvaceae are a nearly cosmopolitan family found on all continents and most oceanic islands, extending to northern North America, Iceland, Scandinavia and Siberia. They are absent from the Arctic and Antarctic and the Gobi Desert. Phylogeny and evolution: In most morphology-based classifications, the current members of Malvaceae were distributed in four families. Of these, Malvaceae were

Grewia occidentalis, Nairobi, Kenya [283]

found to be monophyletic, but members of Bombacaceae, Sterculiaceae and Tiliaceae were not monophyletic, all of these would have to be recircumscribed if they were to be maintained as separate families, and several additional families would have had to be accepted. To stabilise the taxonomy of this clade, it was decided in APG III (2009) to merge all into a single larger family Malvaceae, in which nine subfamilies are recognised. The relationships among these subfamilies is not entirely clear. Crown Malvaceae are estimated to have diversified c. 31–44 million years ago, although an age of 66 million years has also been suggested. Tilioideae and Helicteroideae fossils are known from the early Tertiary onwards, and these genera were then much more widely spread than is currently the case. Several genera have recently been recircumscribed. For instance, Disaster (formerly in Rhamnaceae) and Pimia are now considered synonyms of Commersonia (Byttnerioideae). Oceanopapaver has been shown to belong to the large and diverse genus Corchorus, in which Pseudocorchorus may also may belong; Diplophractum and Columbia are synonyms of Colona (Grewioideae). Hainania is merged with Pityranthe, and Tahitia belongs to Christiana (Brownlowioideae). Chorisia is merged with Ceiba, and Bombacopsis is part of Pachira (Bombacoideae). Azanza may have to be recognised as separate from Thespesia. Abelmoschus, Decaschistia, Fioria, Goethea, Kosteletzkya, Macrostelia, Malvaviscus, Papuodendron, Pavonia, Senra and Talipariti are included in Hibiscus, although not all

Clappertonia ficifolia, Singapore [283]

combinations have been made. Lavatera is a synonym of Malva (Malvoideae). Abutilon and Sida are polyphyletic, and many other genera seem to be in need of recircumscription. Genera and species: Malvaceae are a family with 244 genera and 4,225 species in nine subfamilies: Byttnerioideae (26 genera) – Abroma (1), Ayenia (>70), Byttneria (>130), Commersonia (c. 12), Dicarpidium (4), Glossostemon (1), Guazuma (3), Guichenotia (14), Hannafordia (3), Hermannia (>100), Herrania (17), Keraudrenia (9), Kleinhovia (1), Lasiopetalum (c. 35), Leptonychia (c. 45), Lysiosepalum (5), Maxwellia (1), Megatritheca (2), Melochia (c. 60), Raylaya (1), Rulingia (c. 22), Scaphopetalum (c. 20), Seringia (1), Theobroma (22), Thomasia (c. 30) and Waltheria (c. 55); Grewioideae (25 genera) – Ancistrocarpus (4), Apeiba (7), Clappertonia (3), Colona (c. 30), Corchorus (c. 70), Desplatsia (c. 5), Duboscia (3), Eleutherostylis (1), Entelea (1), Erinocarpus (1), Glyphaea (3), Goethalsia (1), Grewia (c. 290), Heliocarpus (1), Hydrogaster (1), Luehea (c. 25), Lueheopsis (7), Microcos (c. 60), Mollia (c. 18), Pseudocorchorus (6), Sparrmannia (3), Tetralix (7), Trichospermum (c. 36), Triumfetta (c. 150) and Vasivaea (2); Tilioideae (3 genera) – Craigia (2), Mortoniodendron (c. 12) and Tilia (c. 23); Brownlowioideae (8 genera) – Berrya (5), Brownlowia (17), Carpodiptera (>6), Christiana (4), Diplodiscus (8), Jarandersonia (5), Pentace (c. 25) and Pityranthe (2); Helicteroideae (12 genera) – Boschia (7), Coelostegia (5), Cullenia (3), Durio (c. 20), Helicteres (c. 60), Kostermansia (1), Mansonia (5), Neesia (8), Neoregnellia (1), Reevesia Plants of the World

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EUDICOTS

(25), Triplochiton (2) and Ungeria (1); Sterculioideae (13 genera) – Acropogon (>12), Argyrodendron (7), Brachychiton (31), Cola (10), Pterygota (>12), Scaphium (10) and Sterculia (c. 250); Dombeyoideae (21 genera) – Andringitra (6), Astiria (1 extinct), Burretiodendron (6), Cheirolaena (1), Corchoropsis (1), Dombeya (>200), Eriolaena (8), Harmsia (2), Helmiopsiella (4), Helmiopsis (9), Melhania (c. 50), Nesogordonia (18), Paradombeya (3), Paramelhania (1), Pentapetes (1), Pterospermum (c. 18), Ruizia (1), Schoutenia (9), Sicrea (1), Trochetia (5+1 extinct) and Trochetiopsis (2+1 extinct); Bombacoideae (27 genera) – Adansonia (8), Aguiaria (1), Bernouilla (4), Bombax (8), Camptostemon (2), Catostemma (>10), Cavanillesia (4), Ceiba (11–20), Chiranthodendron (1), Eriotheca (c. 20), Tilia americana, Mission Peninsula, Michigan, USA [283]

Brownlowia elata. Illustration from W. Roxburgh (1819). Plants of the coast of Coromandel, vol. 3 [283]

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Fremontodendron (3), Gyranthera (2), Huberodendron (4), Lagunaria (1), Matisia (c. 30), Neobuchia (1), Ochroma (1), Pachira (c. 50), Patinoa (4), Pentaplaris (3), Phragmotheca (11), Pseudobombax (c. 20), Quararibea (>20), Scleronema (5), Septotheca (1), Spirotheca (9) and Uladendron (1); Malvoideae (100 genera): Abutilon (c. 135), Acaulimalva (19), Akrosida (1), Alcea (c. 60), Allosidastrum (4), Allowissadula (9), Althaea (c. 12), Alyogyne (5), Anisodontea (19), Anoda (23), Anotea (1), Asterotrichion (1), Azanza (5), Bakeridesia (9), Bastardia (3), Bastardiastrum (8), Bastardiopsis (10), Batesimalva (4), Billieturnera (1), Briquetia (5), Callianthe (45), Callirhoe (9), Calyculogygas (1), Calyptraemalva (1), Cenocentrum (1), Cephalohibiscus (1), Cienfuegosia (26), Corynabutilon (6), Cristaria (c. 75), Decaschistia (17), Dendrosida (7), Dicellostyles (3), Dirhamphis (2), Eremalche (3), Fryxellia (1), Fuertesimalva (14), Gaya

Helicteres baruensis, Playa Maya, Yucatán, Mexico [283]

(33), Gossypioides (2), Gossypium (49), Gynatrix (2), Hampea (21), Helicteropsis (1), Herissantia (6), Hibiscadelphus (3+4 extinct), Hibiscus (c. 675), Hochreutinera (2), Hoheria (6), Horsfordia (4), Howittia (1), Humbertianthus (1), Humbertiella (6), Iliamna (7), Julostylis (3), Jumelleanthus (1), Kearnemalvastrum (2), Kitaibela (2), Kokia (3+1 extinct), Krapovickasia (4), Kydia (2), Lawrencia (12), Lebronnecia (1), Lecanophora (5), Malachra (c. 9), Malacothamnus (11), Malope (3), Malva (c. 25), Malvastrum (15), Malvella (4), Megistostegium (3), Meximalva (2), Modiola (1), Modiolastrum (5), Monteiroa (10), Napaea (1), Nayariophyton (1), Neobaclea (1), Neobrittonia (1), Nototriche (c. 100), Palaua (c. 15), Peltaea (16), Periptera (5), Perrierophytum (9), Phragmocarpidium (1), Phymosia (8), Plagianthus (2), Pseudabutilon (19), Radyera (2), Rhynchosida (2), Robinsonella (15), Rojasimalva (1), Sida (c. 250), Sidalcea (c. 20), Sidasodes (2), Sidastrum (>8), Sphaeralcea (c. 40), Symphyochlamys (1), Tarasa (c. 30), Tetrasida (2), Thespesia (12), Urena (c. 7), Wercklea (12) and Wissadula (c. 26). Uses: One of the most important Malvaceae crops is cocoa (Theobroma cacao). Native to tropical areas of Central and South America, the seeds have been used to prepare drinks and for ceremonial purposes in coastal Chiapas (Mexico) since at least 1900 BC. Initially it was probably related to the preparation of fermented alcoholic beverages. The Mayans believed kakaw, together with other food crops, was discovered on a mountain by the gods. To prevent counterfeiting, the Aztec empire installed cacao beans as a currency. Montezuma drank large quantities of cocoa, effectively consuming money. Spanish conquistadors brought cocoa beans to Spain in the mid 16th century, and from there it rapidly spread to France, England and other European countries, leading these nations to establish cocoa plantations in their colonies. A drink made from cocoa beans, often sweetened with milk added, became popular in the 18th century. Currently the vast majority of cocoa powder comes from plantations in Africa (especially Ivory Coast, Ghana and Nigera) and Indonesia. The seeds lie in a white, edible pulp,

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Cola nitida, Matthaei Botanical Garden, Sterculia nobilis, Beijing Botanical Ann Arbor, Michigan, USA [283] Garden, China [283]

Brachychiton populneus in fruit, Gundabooka National Park, New South Wales, Australia [283]

Trochetiopsis ebenus, Royal Botanic Gardens, Kew, UK [283]

Dombeya burgessiae, Adelaide Botanic Garden, South Australia [283]

which is sometimes juiced to make a refreshing drink. The seeds are fermented, roasted and ground to make cocoa mass. Seeds contain high percentages of fat (up to 50%), which is separated as cocoa butter. The remaining cocoa solids are ground into cocoa powder, which is then mixed with sugar and cocoa butter or other fat to make chocolate (from Nahuatl xocolatl, bitter water). Milk chocolate is dark chocolate with milk added, and white chocolate lacks the cocoa solids. Cupuaçu (Theobroma grandiflorum) is closely related, and the seeds can also be used to make chocolate, so it has high potential as a crop. However, the fruit is better known in western Amazonia for its juice that makes excellent ice-cream. Native to the rainforests of West Africa, kola nuts (Cola acuminata and C. nitida) are high in caffeine and roasted, pounded or chewed or added as a f lavouring to drinks. They are best known for flavouring caffeinated drinks such as commercial cola

and energy drinks. Seeds of the Chinese parasol tree (Firmiana simplex) also contain caffeine and can be used to make a tea. The wood of this species is valued locally for musical instruments. Durian (Durio zibethinus and other Durio species) is called the ‘king of fruits’ and is known for its large size (to 30 cm across) and distinctive, foetid odour. Some people find the flavour of the edible pulp that surrounds the seeds pleasantly sweet, whereas others find it revolting. The scent is persistent, and this has led to its banishment from flights and hotels in Southeast Asia. Chupa chupa (Quararibea cordata) has, like several other species of that genus, edible fruits that are eaten fresh or made into a soft, sweet juice. Baobab (Adansonia digitata and other species) has found increasing interest in the past few years because the powdered seeds can be used as a food thickener, sweetener or spice. It is now marketed in the EU and USA as a superfood.

Brachychiton populneus in flower, planted in Beirut, Lebanon [283]

The seeds can also be pressed into an oil for cooking. The fruit pulp of all species can be eaten fresh, but mostly A. digitata from Africa is used and even employed as an ingredient in brewing beer in Tanzania. Fruits of Thespesia garckeana are also edible. Roselle or rosa jamaica (Hibiscus sabdariffa), from tropical Africa, has fleshy calyces that are used to flavour drinks, teas, jellies and deserts. The young shoots are also eaten as a vegetable, and the stems can be used for fibre. Okra, bindi or lady’s fingers (Hibiscus esculentus) has edible young fruits that are a popular vegetable in Asian, African, Caribbean, southern USA and Middle Eastern cuisines. The closely related ambrette (H. abelmoschus) provides a seed oil, and the plants have strong fibres. Tree spinach (H. manihot) also has jute-like fibres used for rope making, but it is most commonly cultivated for its young shoots, which are cooked like spinach. West African okra (H. manihot var.

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MALVALES

Ceiba speciosa, Curitiba, Paraná, Brazil [283]

EUDICOTS

Adansonia gregorii, JudbaraGregory, Northern Territory, Australia [283]

caillei) is a cultigen derived from it, grown for the young fruits. Economically, the fibres of Malvaceae are the most lucrative, especially cotton and jute. Cotton plants (Gossypium spp.) bear hairy seeds, which have single-celled hairs that are 30 times longer than wide (a trait that has been selected by humans since prehistoric times) and can be spun into thread when dried. It has been cultivated since ancient times to make fabric, and many cultivars and hybrids are now grown, some natural, some genetically modified. The fibres are also used to make paper, twine, viscose, cellophane and photographic film. The oil of cotton seeds is used in cooking and to make soap. Flour of the seeds is high in protein and can be eaten, although it may also contain poisonous gossypol, a male contraceptive, but seeds without this compound have now been genetically engineered, allowing them to be used as a protein source. Tree cotton (G. arboreum) has been cultivated in Pakistan since 1800 BC and has been grown in the Middle East for several millennia. African cotton (G. herbaceum) was independently domesticated in East Africa and so was sea island cotton (G. barbadense), of which seeds have been found in 5,000-year-old archaeological sites in Ecuador. Gossypium hirsutum (upland or Mexican cotton) is the species most widely planted in the United States. Egyptian cotton was derived from 392

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Fremontodendron californicum ‘Butano Ridge’, University of California Botanical Garden, Berkeley, USA [283]

G. barbadense in the early 19th century. Kapok (Ceiba pentandra) originated in the Caribbean region, but because it produces a light, buoyant fibre, it has been used in the past as insulation and a filling for packaging, mattresses, pillows, furniture, life jackets and stuffed toys. It was for that reason planted around the tropics but has now largely been replaced by artificial fibres because kapok is highly flammable and its harvest is labour intensive. The seed oil of kapok is locally used to make soap and has a mild odour and taste, similar to cottonseed oil. Jute has long, glossy, soft fibres that can be spun into strong thread. It is mostly made from members of the genus Corchorus (Chinese jute, Corchorus capsularis; tossa jute, C. olitorius), but also from other specis such kenaf or Deccan jute (Hibiscus cannabinus), tree hollyhock (H. hamabo), bolo bolo (Clappertonia ficifolia), kolle (Decaschistia crotonifolia), Brazil jute (Malachra capitata), burweed or burbark (Triumfetta spp.), Congo jute (Urena lobata), maholtine (Wissadula amplissima) and pacopaco (W. spicata). Althaea cannabina fibre was formerly used for paper making. Young shoots of Corchorus and Adansonia are eaten like spinach in Africa. Lindens or limes (Tilia spp.) are important shade trees and good sources of honey, and the flowers can be harvested for herbal teas that are sweet and aromatic. They have been used for various minor ailments in herbology,

Pachira aquatica, New York Botanical Garden, USA [283]

especially in Europe where Tilia cordata is most valued. The wood of Tilia is easily worked because of its fine grain and density. It was often used to make shields in Mediaeval times and for intricate wood carvings. It is also the favoured wood for puppets and marionettes. The Ainu people in Japan weave their clothing from the bark of Tilia. Mallows (Malva spp.) have many traditional uses, but best known is the curled mallow (Malva verticillata ‘Crispa’) that is sometimes cultivated for its sweet leaves used as a salad green. Fruits of other mallows (e.g. M. parviflora, M. sylvestris) have been eaten in the past and were cultivated for this purpose; they have now spread to many areas outside their native range. Many species of Malvaceae produce mucilaginous substances, and well known for this was marshmallow (Althaea officinalis), used to make marshmallows, a spongy, sticky treat, now usually made with gelatin. Some species of Sida have hallucinogenic properties, especially S. acuta, which is used as a cannabis substitute in Mexico. Sida rhombifolia, a pantropical weed, can also be used as a stimulant because of its ephedrine content. Additionally, fibre of this species is used locally in basketry. Extract of scarlet globe mallow (Sphaeralcea angustifolia) is used as ‘negrita extract’ in hair conditioners and other cosmetic products. Apart from Tilia, there are several species

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EUDICOTS

that produce high-quality wood. Heritiera species are valuable timber trees, often used in ship building, including mengkulang (H. cochinchinensis), chumprak (H. littoralis), crowsfoot elm (H. trifoliolata) and niangon (H. utilis). Blue mahoe (H. elatus) has fine wood used for cabinetry and bark useful for ropes and weaving hats. Mahoe (Thespesia populnea) and other Thespesia species have a hard wood suitable for turning, wheel frames and furniture. Balsa wood is light and harvested mostly from a common, rapidly growing, weedy species, Ochroma pyramidale. Some timber species are now highly endangered. Examples are St Helena ebony (Trochetiopsis ebenus) and St Helena redwood (T. erythroxylon, now extinct in the wild), which yielded a dark hardwood, now critically endangered. The dwarf ebony (T. melanoxylon) formerly on St Helena but Hoheria sexstylosa, East Bergholt Place Arboretum, Suffolk, UK [283]

Gossypium darwinii, fruiting, Galápgos Islands [283]

now extinct, as well as some Hibiscadelphus and Kokia species from Hawaii due to overexploitation and mismanagement. Many species are used as ornamentals, especially those in Abutilon (velvetleaf), Adansonia (baobab), Alcea (hollyhock), Althaea (marshmallow), Alyogyne (lilac hibiscus), Anisodontea (African mallow), Azanza (snot apple), Bombax (silk cotton tree), Brachychiton (bottletree or kurrajong), Callianthe megapotamica (trailing abutilon), Callirhoe (poppy mallow), Ceiba speciosa (silk floss tree), Chiranthodendron pentadactylon (devil’s hand tree), Clappertonia ficifolia (bolo bolo), Dombeya (pink-ball), Firmiana simplex (parasol tree or wutong), Fremontodendron californicum (f lannelbush), Gossypium sturtianum (Sturt’s desert rose), Grewia (crossberry), Hibiscus (e.g. H. malvaviscus; H. mutabilis, Confederate rose; H. rosa-sinensis, Althaea officinalis, Brooklyn Botanical Garden, New York, USA [283]

China rose; H. syriacus, rose of Sharon; H. tiliaceus, hau), Iliamna (wild hollyhock), Kitaibela vitifolia (cedar cup), Malacothamnus (bush mallow), Malope (mallowwort), Malva (mallow and lavatera), Malvastrum (false mallow), Pachira aquatic (Guiana chestnut), Plagianthus (ribbonwood), Rulingia (kerrawang), Sida hermaphroditica (Virginia fanpetals), Sidalcea (prairie mallow), Sparrmannia (African linden), Sphaeralcea (globe mallow), Thespesia (portia tree), Tilia (lime or linden) etc. Etymology: Malva is the classical Latin name for mallow. It is of Semitic origin (e.g. maluakh in Hebrew), meaning a saltbush, which came into Ancient Greek as μαλακε (malakhe; compare also Malacha and Malacothamnus) from which it entered Latin and numerous other European languages.

Hibiscus insularis, Adelaide Botanic Garden, South Australia [283]

Sida neomexicana, New Mexico, USA (DZ) [283]

Malva sylvestris, Ioannina, Greece [283]

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MALVALES

EUDICOTS

Rhopalocarpus similis, Andranomaitso, Madagascar (CD) [284]

284. SPHAEROSEPALACEAE Lombiry family

Rhopalocarpus triplinervis, fruit, Irodo, Antsiranana, Madagascar (CD) [284]

middle and at the base of the ovary (gynobasic) in Dialyceras. The fruit is an indehiscent, dry capsule, each locule with a single seed, the fruit surface often ornamented, which is useful for species identification. Distribution: This family is restricted to northern, central and eastern Madagascar.

These are trees and shrubs with alternate, petiolate leaves, deciduous in the dry season. Stipules are broad and fused across the petiole and enclose the buds, leaving a circular scar around the stem. Blades are simple with entire margins and pinnate or palmate venation with three main veins emerging from the leaf base. Inflorescences are bracteate, terminal or axillary panicles composed of cymes. The actinomorphic flowers are bisexual and pedicellate. The four sepals are free; the outer two are smaller and incurved, and hairy inside. The four petals are free and hairy, or not. Stamens are numerous, free or shortly basally fused, with thread-like filaments and dorsifixed anthers that open by lengthwise slits. The superior ovary is usually elevated on a short outgrowth of the receptacle (gynophore), surrounded by an annular disk. Usually two, sometimes one to five, carpels are fused to form a locule, the styles fused with a disc- or globe-shaped entire stigma. The style is apical in Rhopalocarpus in the 394

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Phylogeny and evolution: The family was previously known as Rhopalocarpaceae, but the name Sphaerosepalaceae has nomenclatural priority. They have previously been placed in a variety of families, including Clusiaceae, Elaeocarpaceae, Flacourtiaceae, Capparaceae, Tiliaceae, Bixaceae and Sarcolaenaceae. Placement in Malvales is supported by anatomical, pollen and molecular data, and they are perhaps closest to Thymelaeaceae and Bixaceae. Dialyceras with its gynobasic style, is similar to Diegodendron (Bixaceae), which resembles Sphaerosepalaceae in several morphological characters. Genera and species: Sphaerosepalaceae include two genera and 18 species: Dialyceras (3) and Rhopalocarpus (15). Uses: Even though many species of this family are highly endangered, the wood of some is still logged for timber, tool handles and charcoal, placing these already rare plants under further threat. Phloem fibre has been used locally to make rope.

Etymology: Sphaerosepalum is composed of the Greek words σφαίρα (sfaira), a globe or sphere, and Latin sepalum, a sepal (from Greek σκέπη, skepe, a covering). Sphaerosepalum is a later synonym of Rhopalocarpus, which is also Greek, meaning ‘club fruit’ in reference to the two-parted, ornamented capsules.

285. THYMELAEACEAE Mezereon family

A family of bisexual, sometimes unisexual, shrubs, perennial herbs and small trees, rarely vines, they are evergreen and deciduous and are usually poisonous. They have a tough and fibrous bark, which is usually covered with silky fibres inside. Leaves are usually alternate, sometimes opposite or in threes (ternate) and lacking stipules. A petiole is sometimes present or the leaf sessile and articulated to the stem. Leaf blades are simple, with entire margins and pinnate venation, with the secondary veins sometimes terminating in a marginal vein. Inflorescences are terminal, or less often axillary, sessile or stalked cymes, racemes, heads, spikes,

MALVALES

EUDICOTS

umbels or fascicles, sometimes produced on short shoots (brachyblasts), with inflorescence bracts sometimes colourful, often forming an involucre around the inflorescence or bracts absent. Flowers are usually actinomorphic and bisexual or unisexual. A floral tube is usually present and well developed, cup-shaped or tubular, often colourful and corolla-like, topped by usually four or five (rarely three to six) free sepals. Petals are either fully developed (clawed in Tepuianthus) or reduced to small scales, usually as many or twice as many as sepals, inserted on the rim of the floral tube and sometimes fused to form a ring. The one or two to many stamens (usually as many or twice as many as sepals) are inserted opposite the sepals in the floral tube or on

the receptacle. Filaments are free or reduced (and the anther sessile), rarely fused into a tube, and anthers are basifixed or dorsifixed, opening by lengthwise slits. A disk is usually present at base of ovary, scale-like, ring-like, cup-shaped or absent. The superior ovary is one- to 12-locular and sessile or shortly stalked (on a gynophore), with a style placed terminally or laterally on the ovary topped with a papillose, capitate, globular, pyramidal or club-shaped stigma or the stigma sessile on the ovary, globose, subglobose, subclavate, or pyramidal, sometimes papillose. Two or three free styles are found in Tepuianthus. Fruits are usually indehiscent, dry or fleshy capsules or berries, sometimes loculicidal with up to nine parts.

Distribution: Thymelaeaceae have a nearly worldwide distrubution in the Americas in eastern North America and California and from central Mexico and the Caribbean to Fuegia and the Falklands. They grow throughout Europe, Asia and Africa, apart from drier areas. They also occur throughout Australasia and the Pacific.

Pimelea imbricata var. piligera, Mt Benia, Western Australia [285]

Stellera chamaejasme, Yunnan, China [285]

Wikstroemia indica, New Caledonia [285]

Edgeworthia chrysantha, Royal Botanic Gardens, Kew, UK [285]

Daphne bholua ‘Jacqueline Postill’, private garden, Kingston upon Thames, Surrey, UK [285]

Phylogeny and evolution: Previously Thymelaeaceae were thought to be closely related to Euphorbiaceae, and indeed their pollen is similar to that of Euphorbiaceae subfamily Crotonoideae. Moreover, specimens of Euphorbiaceae/Peraceae and Thymelaeaceae were often reciprocally confused, especially genera like Gnidia versus Clutia,

Phaleria capitata, Singapore (WA) [285]

Gnidia latifolia in fruit, Taita Hills, Kenya [285]

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MALVALES with some having been labelled alternately as belonging to each family several times. Molecular analyses place Thymelaeaceae in Malvales, close to Bixaceae and far away from Euphorbiaceae (Malpighiales). Their age has been estimated to around 75 million years, but the first fossil pollen records of Thymelaeaceae subfamily Octolepidoideae are known from the Eocene. Tepuianthus, sometimes placed in its own family, Tepuianthaceae, is included based on molecular and morphological characters. Generic delimitation is still in need of study, although some realignments have already been made. Daphne and Wikstroemia are not easily delimited and may have to be redefined. Restella is placed in Wikstroemia, and the position of Stellera with regard to Wikstroemia is uncertain. Gnidia has been expanded to include Arthrosolen, Atemnosiphon, Craspedostoma, Englerodaphne, Pse udog nidia a nd Struthiolopsis, but Lasiosiphon is excluded from Gnidia. Thecanthes is now part of Pimelea, and Eriosolena belongs in Daphne. Genera and species: Thymelaeaceae consist of c. 46 genera and c. 913 species in three subfamilies; Tepuianthoideae – Tepuianthus (7); Octolepidoideae – Aëtoxylon (1), Amyxa (1), Arnhemia (1), Deltaria (1), Gonystylus (32), Lethedon (15), Octolepis (6) and Solmsia (2); Thymelaeoideae – Aquilaria (20), Craterosiphon (9), Dais (2), Daphne (98), Daphnopsis (72), Diarthron (19), Dicranolepis (16), Dirca (4), Drapetes (1), Edgeworthia (5), Enkleia (4), Funifera (4), Gnidia (c. 100), Goodallia (1), Gyrinops (9), Jedda (1), Kelleria (11), Lachnaea (40), Lagetta (3), Lasiadenia (2), Lasiosiphon (85), Linodendron (4), Linostoma (3), Lophostoma (4), Ovidia (3), Passerina (20), Peddiea (14), Phaleria (24), Pimelea (c. 110), Rhamnoneuron (2), Schoenobiblus (10), Stellera (1), Stephanodaphne (9), Struthiola (35), Synandrodaphne (1), Synaptolepis (5), Thymelaea (32) and Wikstroemia (90). Uses: The phloem of many species contains strong fibres, so strong that one can make a

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knot in a thin branch without breaking it. This makes the bark (which can be recognised by the silky hairs inside) suitable for manufacture of high-quality paper, such as that previously used for bank notes (now largely replaced by artificial fibres or plastics). Paperbush or mitsumata (Edgeworthia chrysantha) has been used to make high-quality traditional paper in Japan since ancient times. Himalayan E. gardneri has also been used locally for papermaking. Both make attractive garden plants. African Gnidia glauca also makes a high-quality paper. Wikstroemia sikokiana, known as gampi in Japan, and some species of Thymelaea in the Mediterranean, are also used to make paper. Bast fibre (surrounding the phloem) of many other Thymelaeaceae (e.g. Aquilaria, Dais, Daphne, Daphnopsis, Dirca, Gnidia, Gyrinops, Phaleria, Pimelea, Thymelaea and Wikstroemia) is good for making string, rope and high-quality fabric. Lacebark (Lagetta lagetto) makes a lace-like inner bark, used as a textile in Jamaica. Agarwood or aloewood is the decaying heartwood of species of Aquilaria, Aëtoxylum sympetalum, Gonystylus and Gyrinops. Agarwood is saturated with oleoresins, which can be used as incense and medicinally in commercial products against intestinal parasites. These species are threatened due to overexploitation. Ahaloth or calambac (Aquilaria malaccensis) was the aloe of the Bible and has been used and traded from Indomalaysia since ancient times. Ramin (Gonystylus bancanus) from western Malesia gives light wood for construction, furniture and walking sticks. Wallapatta (Gyrinops walla) is harvested in Sri Lanka for its light timber. In China Daphne genkwa is used clinically as a reputedly safe abortifacient. Mezereon (D. mezereum) was formerly used in Europe as a pepper substitute, often with fatal consequences. In Greek mythology the fountain nymph or naiad Daphne was so beautiful that she attracted the attention of Apollo. Her father, the river-god Ladon, then changed her into a laurel bush, hence the scientific name of the spurge laurel (Daphne laureola). Many species of Daphne and to a lesser extent Dais cotinifolia (pompom tree), Dirca palustris (moosewood), Edgeworthia

chrysantha (paperbush), Ovidia andina (trarovoqui), Pimelia spp. (riceflowers), Stellera chamaejasme (lang du), Struthiola myrsinites (gonnabos) and Wikstroemia are cultivated for their scented flowers in gardens. Etymology: Thymelaea is probably derived from Greek θυμός (thymos), anger or poison, and ελιά (elia), the olive, in its resemblance to an olive branch but with a poisonous fruit.

286. BIXACEAE Annatto family

Bixaceae are deciduous and evergreen trees, shrubs and perennial herbs with woody rootstocks. Leaves are alternate, with free stipules on both sides of the petioles (stipules are fused and enclose the bud in Diegodendron). Blades are simple to deeply palmately dissected or lobed, with entire to toothed margins and palmate venation (pinnate in Diegodendron). Inflorescences are terminal, rarely axillary, thyrses. The bisexual flowers are actinomorphic, slightly zygomorphic in Amoreuxia, and often large and showy. In Bixa and Diegodendron pedicels have five (peltate) glands below the sepals. The five free sepals and petals alternate and are yellow, white or pink. Stamens are numerous and free, with the filaments evenly clustered in two or five fascicles. Anthers are basifixed and open with two small pores or slits at the tip and sometimes also with two pores at the base. They split lengthwise in Diegodendron. The superior ovary is composed of two to five fused carpels topped with an elongate style and cleft stigma. Fruits are two- to fiveparted capsules, the seeds curved around and sometimes with a sarcotesta that turns soft when moistened (Bixa) or the capsule indehiscent (Diegodendron).

MALVALES

EUDICOTS

Amoreuxia palmatifida, Chontales, Nicaragua (CD) [286]

Cochlospermum vitifolium, near Guayaquil, Ecuador [286]

Distribution: Bixaceae have a pantropical distribution but are most diverse in the Neotropics where Amorouxia and Bixa are endemic and Cochlospermum is diverse from Mexico and the Caribbean to tropical Brazil and Bolivia. Cochlospermum is also found in the African tropics and India, Indochina and northern Australia. Diegodendron is restricted to eastern and northern Madagascar. Phylogeny and evolution: Bixaceae are sister to the Dipterocarpaceae-Cistaceae clade from which they separated 52.4 million years ago. Diegodendron has wood and many other characters that are similar to Sphaerosepalaceae, but molecular studies have shown that it is close to Bixa and embedded in Bixaceae if they also include former Cochlospermaceae. Even though the genera have many differences in morphology, they also share sufficient characters to warrant them to be united into a single family.

Bixa orellana in fruit, Singapore Botanical Garden [286]

and widely used in many processed foods such as cheese (especially Double Gloucester and Red Leicester) and candy, and in cosmetics. In pre-Columbian times in South America it was already cultivated and used for body paint in religious ceremonies or as a sunscreen. Seed hairs of Cochlospermum can be used as surgical wool. Many species are harvested locally for fibre, firewood, gum and dye (from the seeds). Cochlospermum vitifolium is sometimes grown as a garden ornamental in the tropics. Etymology: Bixa is derived from bija or biché, a Carib name for the plant.

287. CISTACEAE Rock-rose family

Genera and species: This is a family of four genera and c. 23 species: Amoreuxia (4), Bixa (5), Cochlospermum (13) and Diegodendron (1). Uses: The bright red seeds of the lipstick tree or annatto (Bixa orellana) produce a common food dye (E160b), consisting mainly of bixin. The colour is resistant to heat, light and frost

This family comprises annual and perennial herbs and shrubs (rarely canopy trees) that often have volatile oils. Hairs are simple or clustered so as to seem stellate, and they are

Bixa orellana, Singapore Botanical Garden [286]

often glandular, making the plant feel sticky. Leaves are simple and usually opposite, sometimes alternate or rarely whorled. They usually lack stipules, although they can be present and are then minute or present and large in Pakaraimaea. Leaf blades have an entire margin and pinnate or palmate venation or with three more or less parallel veins. Inflorescences are ancillary cymose panicles or racemes, or the flowers are solitary. Flowers are actinomorphic and bisexual but sometimes not opening (cleistogamous). The usually persistent sepals are three and free or five and then the outer two narrower and sometimes fused to the inner three. The five, sometimes three (Lechea), petals are crumpled in bud and fall off within hours of anthesis, sometimes contorted in bud (Pakaraimaea). Petals are sometimes absent in cleistogamous flowers. Stamens are numerous, inserted on a disklike receptacle, which is enlarged into a short androgynophore in Pakaraimaea, and anthers open with longitudinal slits, sometimes with broad connectives. The superior ovary is composed of three to five fused carpels forming a single locule (pentalocular in Pakaraimaea). It is topped with a simple style or one to three sessile stigmas. Fruits are many-seeded loculicidal capsules. Distribution: This family is most diverse in the Mediterranean region, but also found in Plants of the World

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MALVALES North, Central and southern South America, the Caribbean, throughout Europe and North Africa, Macaronesia, and from the Middle East to central Asia. Outside the maquis in the European and North African Mediterranean, species usually occur on exposed dry rocky, forested slopes. Phylogeny and evolution: In some morphology-based classifications, Cistaceae were placed in Violales or close to Bixaceae, but molecular analyses support placement near Dipterocarpaceae and Sarcolaenaceae in Malvales. To preserve the integrity of these families, moving the Neotropical genus Pakaraimaea from Dipterocarpaceae to Cistaceae creates clades that can be maintained as families. Some generic rearrangement has also occurred, for instance, Crocanthemum and Hudsonia have been merged with Helianthemum. Few fossils of Cistaceae are known, but there is a flower preserved in Baltic amber, and fossil wood was found in marine sands in Germany, both sites from the Late Eocene. The crown group is dated to c. 14 million

Cistus ladanifer, Royal Botanic Gardens, Kew, UK [287]

Cistus creticus, private garden, Kingston upon Thames, Surrey, UK [287]

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years, and there is evidence of frequent longdistance dispersal across the Atlantic. The Mediterranean basin has been the main centre of diversification for this family. Flowers of Cistaceae are ephemeral and open for only a few hours in full sunlight. Many have stamens that are sensitive to touch and actively dust the visiting insect with pollen.

Cistus, Halimium and Helianthemum are frequently grown as garden ornamentals, and many cultivars, selections and hybrids have been developed.

Genera and species: This is a family of seven genera and c. 170 species: Cistus (c. 17), Fumana (10), Halimium (9), Helianthemum (c. 110), Lechea (c. 18), Pakaraimaea (1) and Tuberaria (c. 10).

288. SARCOLAENACEAE

Uses: A resin called labdanum or ladan is harvested from several Cistus species, especially C. creticus and C. ladanifer. In ancient times (it is mentioned in Genesis in the Bible), it was collected from the hairs of goats and sheep that had been grazing among the shrubs. Ladan was used as a medicinal incense against colds, coughs etc. Today, it is mainly used in the perfume industry, where it is considered a good substitute for whale ambergris. Many species of especially

Helianthemum nummularium, Seglinge, Åland Islands, Finland [287]

Etymology: Cistus is the Latin name for the plant, derived from the Greek κιστος (kisthos), a basket.

Tunic-bells family

These are evergreen trees and shrubs that are covered with peltate and stellate hairs or tufts of hairs, often mixed with multicellular, glandular or simple hairs. They have alternate, petiolate simple leaves with free or fused, Pakaraimaea dipterocarpacea, collected by J. A. Steyermark (no 117745) in Venezuela in 1978 (Herbarium Kew) [287]

MALVALES

EUDICOTS

Leptolaena bojeriana, Isalo National Xyloolaena perrieri, Sahafary, Sarcolaena oblongifolia, fruit, Sarcolaena oblongifolia, Tsimbazaza Park, Madagascar (CD) [288] Andranomadiro, Madagascar (CD) [288] Tsimbazaza Botanical and Zoological Botanical and Zoological Park, Park, Antananarivo, Madagascar (CD) [288] Antananarivo, Madagascar (CD) [288]

small to large stipules that often fall off early. The simple blades are pinnately veined. and secondary venation is curved along the entire margins. Inflorescences are terminal panicles or few-flowered axillary clusters, singly or in pairs surrounded by bracts that develop into an enlarged involucre enclosing the fruit. Flowers are bisexual and actinomorphic. The three to five basally fused sepals are persistent in fruit. The five petals are contorted and free or slightly basally fused. A prominent, cupular or fivelobed disk surrounds ten to numerous stamens. Filaments are free or fused at base, and anthers are basifixed or dorsifixed and open by lengthwise slits. The superior ovary is composed of three (sometimes two to five) fused carpels topped with a persistent style and a broadly lobed stigma. Fruits are usually dehiscent. Distribution: Sarcolaenaceae are restricted to Madagascar.

Leptolaena (9), Mediusella (1), Pentachlaena (3), Perrierodendron (5), Rhodolaena (7), Sarcolaena (8), Schizolaena (20), Xerochlamys (8) and Xyloolaena (5). Uses: Because of their restricted distribution, they are of little economic importance. Some species have involucres that become fleshy in fruit and are edible. Xerochlamys bojerana has been used to flavour rum (rhum arrangé), and several species are locally used for timber and firewood, resulting in deforestation and threats to the native flora of Madagascar, which is full of endemic plants. Larger-flowered species have potential in the horticultural market. Etymology: Sarcolaena is composed of the Greek words σαρκός (sarkos), flesh, and χλαινα (chlaina), a cloak or blanket.

289. DIPTEROCARPACEAE Phylogeny and evolution: This family is closely related to Cistaceae and Dipterocar paceae. Fossil pollen of Sarcolaenaceae is known from Miocene sediments in South Africa, suggesting that the family may have been more widely distributed in the past. Genera and species: Sarcolaenaceae include ten genera and 68 species: Eremolaena (2),

Maranti family

Dipterocar paceae are evergreen and sometimes (partially) deciduous, bisexual trees. Their wood has canals with aromatic resins. Leaves are simple and alternate (spiral or in a plane) and subtended by persistent or deciduous stipules that leave (ring-like) scars. Leaf blades have an entire or sinuatecrenate margin and pinnate venation, often with nectaries on the upper surface when the leaves are young. Inflorescences are terminal or axillary racemes or panicles with minute and short-lived, rarely large and persistent, bracts, and inflorescence and flower parts are usually covered with stellate, scale-like or simple, usually fascicled hairs. Flowers are actinomorphic and bisexual. The five sepals are free or united at the base, and the five petals are fused, sometimes only basally. A stalk bearing the sex organs (androgynophore) is present in Monotoideae. The five to 15 or numerous (up to 110) stamens are in one to three rows and free or fused to the petals. Filaments usually are basally wider, and the connective is pointed at the tip (aristate), filiform or stout. Anthers are basifixed and basiversatile and open by lengthwise slits or sometimes by two apical pores. A superior or rarely partially inferior ovary is usually composed of three carpels, rarely bi- to penta-loculate with a style at the top that is broader at the base (stylopodium) with a trifid or entire tip. The stigma has three Plants of the World

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or one to six lobes. Fruits are usually nut-like, sometimes capsular and three-valved, usually with a single seed. The fruit is enclosed in the persistent, variously developed sepals, of which two or more lobes are usually developed into wings. Distribution: This family is mostly found in tropical Asia and the Seychelles (Dipterocarpoideae; with the highest diversity in northwestern Borneo), but Monotoideae are found in northern South America (Pseudomonotes tropenbosii) and tropical Africa (Marquesia, Monotes). Dipterocarpaceae are dominant rainforest elements, especially on peat lenses in Southeast Asia and Malesia. Phylogeny and evolution: Dipterocarpaceae are most closely related to Cistaceae and Sarcolaenaceae, with which they share many characters, and it has been suggested that the three families could be united. The family is divided into two subfamilies that appear to have evolved out of Africa (where 50 Monotes glaber by O. H. Coates Palgrave, from K. Coates Palgrave (1916) Trees of Central Africa [289]

million-year-old fossils of Dipterocarpoideae are known but where the subfamily does not currently occur), but diversified in tropical Asia. An association with ectomycorrhizae has been suggested to have occurred in this family on the Gondwana supercontinent, 135 million years ago, and all species are strongly ectomycorrhizal today. Some realignment of genera may be in order in light of molecular studies, e.g. Shorea and Hopea could perhaps be merged because Shorea is polyphyletic. Neobalanocarpus heimii may even be an intergeneric hybrid between these two genera. Genera and species: Dipterocarpaceae include 16 genera and c. 695 species, in two subfamilies: Monotoideae – Marquesia (4), Monotes (c. 30) and Pseudomonotes (1); Dipterocarpoideae – Anisoptera (10), Cotylelobium (5), Dipterocarpus (c. 70), Dryobalanops (7), Hopea (c. 104), Neobalanocarpus (1), Parashorea (14), Shorea (360), Stemonoporus (c. 20), Upuna (1), Vateria (3), Vateriopsis (1) and Vatica (c. 65).

Hopea wightiana, Singapore (WA) [289]

Uses: In Asia, dipterocarp species are dominant rainforest elements, and there they have been unsustainably logged during the last century to supply the international market with tropical hardwoods. It is now much less important as a source of timber due to diminished stocks, but plantations have emerged. The best-known woods are red and yellow maranti (Shorea spp.) for cabinetry and construction, balau (also Shorea), popular for flooring and decking, and keruing or mai yang (Dipterocarpus), which was popular for railway sleepers. Seeds of some species are high in fats and can be used as a substitute for cocoa butter. Gurjun oil, derived from Dipterocarpus, is used in paints and lubricants. Camphor wood (Dryobalanops aromatica) has been harvested for its scent compounds since ancient times and is used for perfume. Etymology: Dipterocarpus is composed of the Greek words δύο (dio), two, πτερων ( pteron), a wing or feather, and καρπός (karpos), a fruit. Neobalanocarpus heimii in fruit, Singapore Botanical Garden (CD) [289]

Dipterocarpus kerrii, habit, Malacca, Malaysia [289]

Dipterocarpus baurii, Singapore (WA) [289]

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BRASSICALES Families 290 to 306 comprise Brassicales, an order united by their shared production of glucosinolates, precursors of mustard oils that give Brussels sprouts and capers their characteristic ‘love them or hate them’ taste. Mustard oils (or isothiocyanates) are hydrolytic products of glucosinolates and have been shown to occur in some 15 families of eudicots. Conversion of glucosinolates to mustard oils is mediated by the enzyme myrosinase, which is normally sequestered in myrosin cells. This complicated shared chemical pathway led researchers before the advent of molecular studies to suggest that plants producing these must be closely related to each other, despite their morphological differences. Molecular analyses have since corroborated these relationships, and most are now placed in Brassicales, apart from Drypetes (Putranjivaceae, Malpighiales). Thus, this unique chemical pathway, the ‘‘mustard oil bomb’’, was invented at least twice. Family delimitation in Brassicales has also been a thorny issue, particularly for Brassicaceae, Capparaceae and Cleomaceae. Capparaceae in the traditional (broad) sense have been shown to be polyphyletic in many studies in which most taxa fall in a large clade in which Brassicaceae are embedded; Cleome is sister to Brassicaceae, and Capparaceae s.s. are sister to this. Possible nomenclatural solutions were to transfer the Cleome group to Brassicaceae, to recognise only one family, Brassicaceae s.l., including Capparaceae s.l. or to recognise three families Brassicaceae, Capparaceae and Cleomaceae. The consensus opinion has so far been in favour of the latter option, accepted in APG IV and followed here. The remaining taxa of Capparaceae s.l. sampled to date fall elsewhere in Brassicales or in other orders, resulting in the recognition of several small families.

290. AKANIACEAE Turnipwood family

versatile and open by lengthwise slits. The superior ovary is sessile and composed of three (to five) fused carpels, each forming a locule. The style is simple, longer than the stamens and topped by a capitate stigma. The fruit is a hairy, leathery loculicidal capsule splitting into threes (or fives). Akania hillii (included in A. bidwillii) by M. Smith from Curtis’s Botanical Magazine vol. 138, plate 8469, 1912 [290]

Akaniaceae are trees with myrosin cells in the bark and inflorescences, producing glucosinolates and giving off a mustardy smell. Leaves are alternate, unevenly pinnate with small or reduced stipules. Leaf lets are stalked and opposite or the lower ones alternate, with entire or toothed margins and pinnate venation. Inflorescences are erect terminal or axillary racemes or panicles with or without minute bracts. The bisexual f lowers are actinomorphic or slightly zygomorphic with a cup-shaped receptacle. The five sepals are fused to the receptacle, and the five free petals are longer than the sepals, clawed basally and asymmetrical. The eight to ten stamens are fused to the thin ring-shaped nectary disk at the base of the petals. Filaments are hairy at the base, and the dorsifixed or nearly basifixed anthers are

Distribution: Bretschneidera is found in southern China and adjacent Indochina and Taiwan, and Akania occurs only in eastern Australia. Phylogeny and evolution: Akania and Bretschneidera were originally described

Bretschneidera sinensis, Taiwan (KC) [290]

Bretschneidera sinensis, Taiwan (KC) [290]

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in Sapindaceae, but both were subsequently moved into their own families. The taxonomic placement of Bretschneidera has been problematic, with widely postulated alliances that include Capparaceae and Moringaceae, based on the cabbage-like smell of the wood, due to presence of glucosinolates. Morphologically, the two genera are distinct, but evidence from embryology and wood anatomy indicates a close relationship between them. Akaniaceae s.l. are strongly supported as the sister group of the herbaceous family Tropaeolaceae in molecular studies, and Palaeocene fossil leaves of Akania from Argentina suggests a South American origin of the group or at least a much wider distribution in the Southern Hemisphere in the past.

and κανιά (kania), a bract, referring to the absence of bracts at the base of the flower stalk, distinguishing it from Harpullia (Sapindaceae), to which it previously was believed to be related.

Genera and species: A family of two genera, each with a single species: Akania bidwillii and Bretschneidera sinensis. Etymology: No derivation of Akania is given in the original publication of the name, but it may be from the Greek α- (a-), without,

Tropaeolaceae are annual and perennial herbs and herbaceous vines with the scent of mustard. Perennial species often have tuberous roots, with which they survive adverse periods. They have climbing or prostrate stems, and leaves are alternate and long-petiolate, usually with stipules that are

Tropaeolum tricolor, Royal Botanic Gardens, Kew, UK [291]

Tropaeolum majus, private garden, Kingston upon Thames, Surrey, UK [291]

291. TROPAEOLACEAE Nasturtium family

small and scale-like and fall off early, rarely larger and leaf-like. Leaf blades are simple or palmately compound, the margins entire or palmately parted, lobed or sinuate, often peltate (with the petiole inserted in the middle of the blade). Venation is palmate in simple leaves and pinnate in leaflets of palmately compound leaves. Inflorescences are axillary, indeterminate racemes, sometimes condensed into umbels. Flowers are each subtended by a leaf-like bract, making the flowers appear solitary in the axils. The bisexual flowers are zygomorphic. The five sepals have one (or three) nectariferous spur(s). The five petals (or two due to reduction of lower ones) are free and clawed at the base, and margins are often lobed, dissected, fringed or ciliate. The eight stamens have free filaments that are in a single whorl, and basifixed anthers open by lengthwise slits. The superior ovary is composed of three carpels that form a trilobed ovary with three locules, topped with a single apical style and three linear stigmas. Fruits are schizocarps, splitting into three single-seeded fleshy or dry mericarps. Distribution: Tropaeolaceae are found in the mountains of southern Mexico, Central America and Andean and temperate South America. They have been introduced into Europe, Asia, Africa and Australia, where in some cases they can be aggressively invasive. Phylogeny and evolution: Tropaeolaceae have previously been included in Geraniaceae, from which they differ in having distinct stamens and fruits that split into single-seeded mericarps. Molecular studies have indicated

Tropaeolum speciosum, Royal Botanic Gardens, Kew, UK [291]

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Tropaeolum azureum, Royal Botanic Gardens, Kew, UK [291]

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EUDICOTS

Moringa drouhardii, Toliara, Madagascar

Moringa oleifera, Réunion [292]

Moringa hildebrandtii, The Living Desert, Palm Desert, California, USA [292]

placement in Brassicales, with which they share the production of glucosinolates. The nearest relatives of the Neotropical family Tropaeolaceae are Australian and East Asian Akaniaceae, which have fossils from South America.

Etymology: Tropaeolum is derived from the Greek τροπαίων (tropaion), a trophy, which in classical times was a wooden pole on which the shields of the defeated enemy were fixed, resembling the peltate leaves of some species of nasturtium. The common name should not be confused with the scientific name Nasturtium, a genus of Brassicaceae.

often replaced by glands. Leaf blades are one to three times unevenly pinnate with opposite leaflets, which have entire margins and pinnate venation. Inflorescences are axillary panicles forming thyrses, with bracts and bracteoles usually present but often replaced by glands. The bisexual flowers are zygomorphic to nearly actinomorphic with a cup-shaped or tubular receptacle and a nectary at its base. The five sepals are free and petal-like. The five, free petals are white, yellow or red, and equal or unequal in size, often forming a legume-like (papilionoid) flower. The five stamens are inserted on the rim of the disk, alternating with three to five staminodes (sterile filaments). Anthers are dorsifixed with a single theca opening by a lengthwise slit. The superior ovary is stalked (on a gynophore) and composed of three fused carpels, forming a unilocular cylinder topped by a single hollow style with a truncate stigma. Fruits are elongate, beaked capsules that open by three valves, with oily seeds that are sometimes winged.

(CD) [292]

Genera and species: The family consists of the single genus Tropaeolum with 94 species. Previously four genera were accepted, but three of these (Chymocarpus, Magallana and Tropheastrum) are embedded in Tropaeolum, and all of them have therefore been merged. Uses: Flowers and leaves of garden nasturtium (Tropaeolum majus) are good in salads, a pesto can be made from the leaves, and fruits can be pickled as a substitute for capers. Mashua (T. tuberosum) is cultivated by indigenous peoples in the Andes for its edible tubers, which are eaten like potatoes. They have a peppery flavour when raw, but men should take care in consuming this root vegetable, as it is said to reduce testosterone levels by as much as 45%. Tropaeolum includes a number of well-known garden ornamentals, especially garden nasturtium (T. majus), canary creeper (T. peregrinum) and perennial nasturtium (T. speciosum); other species are becoming better known in specialist collections.

292. MORINGACEAE Horseradish-tree family

Moringaceae are trees and shrubs, rarely herbs with woody rootstocks, sometimes with swollen, succulent trunks. They often smell and taste of mustard. Gum ducts are often present in the bark, exuding a strawcoloured or pink gum. The brittle wood is white or yellowish and fibrous. The alternate leaves are drought-deciduous, but the petiole and rachis are often persistent. Stipules are

Distribution: Moringaceae can be found in arid regions of Africa and southern Asia; most species are found in Madagascar, Namibia, the Horn of Africa, southern Arabia and the Indian subcontinent. Moringa oleifera is widely cultivated throughout the tropics.

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Phylogeny and evolution: Because the f lowers are strongly zygomorphic, they have a striking resemblance to those of Fabaceae, which caused Linnaeus to treat Moringa as a legume. Moringaceae are sister to Caricaceae, with which they share the glandular appendages at the apex of the petiole. Together they form a clade sister to Akaniaceae and Tropaeolaceae, and this larger clade is sister to the remainder of Brassicales. These families share wood and bark features, a complicated coat on the seeds and, of course, glucosinolates with the other Brassicales. An Upper Cretaceous fossil from eastern North America (Dressiantha) shows remarkable similarity to Moringa, but it has two carpels, which makes it similar to the CapparaceaeBrassicaceae clade of Brassicales.

than oranges and higher iron content than spinach. It is most likely also the most proteinrich plant on Earth. Seeds can be used to clean water, and the bark has strong fibres that can be used to make rope. Oil from the seeds is used in skin creams, soap and other cosmetic products. The flour-sack tree from Madagascar (Moringa drouhardii) produces an oil that is used as base for cosmetic products. Etymology: Moringa is a Latinised form of murungai, the Tamil word for the plant.

293. CARICACEAE Papaya family

Genera and species: This family has a single genus, Moringa, with 13 species. Uses: The miraculous, drumstick or horseradish tree (Moringa oleifera) is widely planted throughout the tropics. It has incredibly nutritious leaves, with a higher calcium content than milk, higher vitamin C content

Carica peltata, female in fruit, Yucatán, Mexico [293]

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The papaya family consists of trees and shrubs with swollen, often spiny stems and

(rarely) scrambling vines (Jarilla). Their sap is usually milky or yellow, and one species is covered with stinging hairs (Horovitzia cnidoscoloides). Leaves are alternate and spirally arranged near the tips of branches, and they are simple or palmately divided, the margins deeply lobed to entire. The unisexual (rarely bisexual) f lowers are actinomorphic, and male and female flowers are usually on different plants (dioecious), rarely on the same plant. The five free sepals form a cup. Male flowers have five petals that are fused into a thin tube with nectaries at the base. The ten stamens form two whorls, one fused with the corolla tube, the other with free filaments or fused at the base forming a stamen tube. The basifixed or dorsifíxed anthers open by lengthwise slits. A sterile pistillode is often present. Female flowers have an open corolla of five basally fused petals and lack stamens and staminodes. The superior ovary is composed of five carpels fused to form one or five locules, and this is topped with a fused style with branched stigmas. Fruits are succulent berries, and seeds form five lines inside, each seed covered in a mucilage sac.

Carica papaya, male, Copenhagen Botanical Garden, Denmark [293]

Jacaratia dolichaula, male flower, Selva Bananito, Limon, Costa Rica (CD) [293]

Carica papaya, female flower, Ubatuba, Brazil [293]

Carica peltata, female, Yucatán, Mexico [293]

Vasconcellea pubescens, Ecuador [293]

BRASSICALES

EUDICOTS

Limnanthes vinculans, University of California Botanical Garden, Berkeley, California, USA [294]

Limnanthes douglassii, private garden, Kingston-upon.Thames, Surrey, UK [294]

Distribution: Predominantly a Neotropical family (five genera), their greatest diversity is in Mexico and the Andes. The sixth genus (Cylicomorpha) has two species disjunctly distributed in Africa, one being native to Tanzania, Kenya and Malawi and the other to the Central African Republic, Nigeria, Congo and Cameroon. Phylogeny and evolution: As for nearly all families that produce glucosinolates (mustard oils), Caricaceae belong to Brassicales, in which they are most closely related to Moringaceae. Cylicomorpha from Africa is sister to the other genera, all of which are Neotropical (Mexico to Argentina); thus it and the outgroup exhibit an origin in Africa and diversification in the Americas. Vasconcellea was separated from Carica on morphological and molecular evidence; it includes all species previously included in Carica, except C. papaya and its wild ancestor, C. peltata. Genera: Caricaceae include six genera with about 35 species: Carica (2), Cylicomorpha (2), Horovitzia (1), Jacaratia (8), Jarilla (3) and Vasconcellea (c. 20). Uses: Fruits of all species of Caricaceae are eaten locally, but only the papaya or pawpaw, Carica papaya, is of economic importance, with a history as a crop plant dating to pre-Columbian times in Mesoamerica, where the wild relative

C. peltata also occurs. It was distributed by the Spaniards in early colonial times throughout the Old World Tropics via Luzon in the Philippines in the early 16th century. Because papaya has been known throughout the tropics for some 500 years, many cultures have adopted it as a “traditional” food plant. Fruits of the mountain papaya (Vasconcellea pubescens) are also cultivated and eaten in the Andes, especially where the climate is too cool to grow C. papaya. The wood of Jacaratia is succulent and eaten locally in the Guianas. Tissues and seeds of C. papaya and other species produce papain (papaya proteinase I), an enzyme that breaks down proteins and is used as a meat tenderiser and in medicine to reduce swelling associated with some types of hernia. Etymology: Carica is classical Latin for a kind of fig, hence Ficus carica (Moraceae) for the common domesticated fig.

294. LIMNANTHACEAE Meadowfoam family

Floerkea proserpinacoides, Elmore County, Idaho, USA (CD) [294]

These are annual herbs that produce mustard oils. They have alternate, simple or one to three times pinnately compound leaves without stipules. Leaf blades are entire, pinnately lobed, bipinnate or trifoliolate, and venation is pinnate. Inflorescences are axillary, foliose or bracteate racemes, often flowers solitary in the leaf axils with pedicels elongating after the flowers open. The usually bisexual flowers are actinomorphic and often showy. The three to five sepals are free or basally fused, persistent in fruit. Petals are usually the same number as sepals, free, equal in size and shape, with a basal nectary. The three, six, eight or ten stamens have free, glabrous filaments and dorsifixed anthers that open by lengthwise slits. The sessile superior ovary is composed of two to five carpels that are fused at the base only and have a central style emerging from the base of the ovaries (gynobasic), topped with two to five dry, papillate stigmatic branches. Fruits are schizocarps, dividing into one-seeded, indehiscent nutlets (mericarps). Distribution: Limnanthaceae include two North American genera. Limnanthes (meadowfoam) is restricted to western North America from California to southern Oregon, with one disjunct species on Vancouver Island and adjacent islands in Canada. Floerkea proserpinacoides (false mermaid) ranges from California to Washington in the west Plants of the World

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Setchellanthus caeruleus, Mexico (VS) [295]

and Missouri to Quebec in the east. They are often found in vernal pools and wet meadows. Phylogeny and evolution: Limnanthaceae were previously placed in Geraniaceae, but chemical and molecular data place them firmly in Brassicales, although within this order their position is not fully resolved. The genera diverged an estimated 9–17 million years ago. Genera and species: Limnanthaceae are a small family of two genera with ten species: Floerkea proserpinacoides and Limnanthes (9). Uses: Poached egg plant (Limnanthes douglasii) is a widely grown ornamental. The seeds are high in oil, and the plants have potential as seed crops, producing a wax similar to that of jojoba (Simmondsiaceae). The seeds are also reasonably high in protein. Etymology: Limnanthes is composed of the Greek words λίμνη (limne), a marsh or lagoon, and άνθος (anthos), a flower.

295. SETCHELLANTHACEAE Azulita family

EUDICOTS

Koeberlinia spinosa, New Mexico, USA (DZ) [296]

These are much-branched shrubs with distinct short and long shoots, all parts covered in T-shaped hairs. Succulent leaves are alternate and lack stipules and have short petioles or sessile leaves. Inflorescences consist of solitary, pedicellate, axillary flowers formed on long shoots. The bisexual flowers are actinomorphic. The five to seven sepals are fused along their entire length to form a cap that ruptures irregularly when the flower opens, usually into one or two flaps. The five to seven petals are lilac or blue and clawed at the base. The usually up to 60 stamens are aggregated into five to seven fascicles. Anthers are basifixed and open by two longitudinal slits. The superior ovary is placed on a short stalk (gynophore) and composed of three carpels, each forming a locule only fused on the inner side. A short style with three short stigmatic branches tops the ovary. Fruits are linear, furrowed capsules curved downwards with three to ten seeds per carpel. Distribution: This plant is only found in the arid zones of Mexico, in the states of Chihuahua, Coahuila and Durango and in the Tehuacán Valley on the border between Oaxaca and Puebla States. Phylogeny and evolution: Like many other members of Brassicales, this shrub exhibits marked xerophytic adaptations, notably in having much reduced leaves with slightly sunken stomata. Due to the presence of glucosinolates, it has a pungent, mustardy smell. The trimerous gynoecium, straight seeds with a spatulate embryo and the

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Koeberlinia spinosa in fruit, New Mexico, USA

(DZ) [296]

non-fibrous exotegmen are shared with some other brassicalean families, and it used to be included in Capparaceae. However, Setchellanthus does not fully agree with any one family, and its position in the order is not fully resolved by molecular analyses. It possibly diverged from the rest of Brassicales 54–73 million years ago. Genera and species: This family includes only a single species, Setchellanthus caeruleus. Etymology: Setchellanthus is named for New England botanist William Albert Setchell (1864–1943), and Greek άνθος (anthos), a flower. After botanising in various parts of the world, Setchell became Professor of Botany at the University of California at Berkeley where he built up the herbarium and botanical garden.

296. KOEBERLINIACEAE Allthorn family

This is a family of deciduous, thorny shrubs and small trees. Plants have myrosin cells but do not produce glucosinolates. Stems are erect and profusely branched, the branches green

BRASSICALES

EUDICOTS

with thorny tips. Minute, scale-like leaves are alternate and spirally arranged along the stem. They lack stipules and petioles and have entire leaf margins and pinnate venation. Inflorescences are short, axillary umbel-like racemes with tiny bracts. Flowers are bisexual and actinomorphic. The four persistent sepals are free, and the four petals are free and slightly clawed at the base. There are eight (rarely ten) stamens with free, flattened filaments that each bear a basal nectar-appendage. The dorsifixed anthers are slightly tipped and open with lengthwise slits. The superior overy is composed of two (rarely three) carpels that are fused into one or two locules and topped with a rounded style and a small bilobed stigma. Fruits are round berries with one or two seeds.

Etymology: Koeberlinia is named for German clergyman and amateur botanist Christoph Ludwig Köberlin (1794–1862). He was befriended by C. von Martius and J. Zuccarini, with whom he frequently exchanged plant specimens. He studied the plants of the Alps and Upper Swabia in southern Germany.

slits. A rudimentary pistillode is sometimes present in the middle of the male flower. Female flowers are naked without sepals, petals or staminodes. The ovary is composed of two carpels forming four (false) locules, topped with a bilobed subsessile stigma. Fruits are drupe-like syncarps, each with four hardcoated seeds.

297. BATACEAE

Distribution: Coastal warm-temperate and tropical America on both the Atlantic and Pacific sides, including the Galápagos (B. maritima) and in the seas between southern New Guinea and northern Australia (B. argillicola). Batis maritima is invasive in Hawaii.

Turtleweed family

Genera and species: This family includes the single genus Koeberlinia with two species; K. holacantha and K. spinosa.

These are low, scrambling, succulent shrubs without spines. Plants are unisexual (B. maritima) or bisexual (B. argillicola). Leaves are opposite and simple, with minute paired basal stipules that fall off early. Blades are sessile, succulent and linear, with entire margins and obscure venation. Inflorescences are unisexual, axillary or terminal cone-like spikes (catkins), sometimes lax and spikelike, or the flowers are solitary. Flowers are unisexual and actinomorphic, subtended by scale-like bracts. Sepals form a sheath that splits into two lobes. The four petals are free and clawed, and stamens are four, in between and longer than the petals with a free filament and dorsifixed anthers that open by lengthwise

Phylogeny and evolution: Because they grow in saline habitats, they have evolved morphological similarities to halophytic members of Amaranthaceae, and therefore early taxonomists placed Bataceae in Caryophyllales (alternatively known as Centrospermae). This placement was unlikely because they differ in their type of sieveelement plastids, have different chromosome numbers and lack betalains, typical for that order. Phylogenetic studies have now resulted in the more likely placement of Bataceae as sister to Salvadoraceae in Brassicales, which agree in the presence of glucosinolates and myrosinase and in morphological similarities. Bataceae and Salvadoraceae are similar in vegetative morphology and could well be merged. The biogeography of Bataceae is puzzling. Fruits are viable in seawater for up to three months, When seeds can tolerate

An undescribed new species of Bataceae or Salvadoraceae from Namibia [297]

Batis maritima, male flowers, Galápagos Islands [297]

Batis maritima, Galápagos Islands [297]

Distribution: This family is found in arid regions in southwestern North America (from Arizona to southern Texas), Mexico (Sonora to Tamaulipas south to Oaxaca) with a disjunct population in Bolivia. They are extremely xerophytic shrubs with photosynthetic stems and reduced, short-lived leaves. Phylogeny and evolution: Koeberlinia has previously been included in Capparaceae, but molecular studies support its removal from this family. A relationship with Bataceae and Salvadoraceae has been indicated. It used to consist of a single variable species, but a second species was recently described.

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BRASSICALES submergence in seawater and preferably grow on beaches and in mangrove forests, one would expect the species to be distributed throughout the tropics. However, the two species are found in isolated regions far away from each other and, even more strangely, in the Neotropics one species is found disjunctly along the Atlantic and Pacific coasts. Longdistance dispersal by means other than ocean currents must have played a role here. It is possible that the distribution is restricted by water temperatures. Genera and species: Bataceae include a single genus Batis with two species: Batis argillicola and B. maritima. An undescribed genus and species from the Namibian desert also seems to belong here (perhaps sister to Bataceae plus Salvadoraceae), but further studies of this taxon and a formal description are needed.

been conserved as Bataceae, retaining the incorrect spelling.

with an entire or bilobed stigma. Fruits are indehiscent berries or drupes.

298. SALVADORACEAE

Distribution: This family is distributed across the drylands of tropical and subtropical Africa, Madagascar, Arabia, India and Southeast Asia, east to the Philippines, often in somewhat saline soils.

Toothbrush-tree family

Etymology: Βάτης (batis) is Greek for walking, in reference to the scrambling nature of this shrub. When grammatically correctly conjugated, this family should be called Batidaceae, but the name has

Salvadoraceae are shrubs and small trees that are sometimes scrambing or scandent. Leaves are opposite and simple, thickly leathery or succulent, with minute basal stipules and short petioles. Sometimes plants are armed with axillary spines (Azima). Inf lorescences are axillary or terminal racemes, often compound in panicles or flowers fasciculate in leaf axils. Flowers are bisexual or unisexual and actinomorphic. The two to four (rarely five) sepals are fused to form a cup-shaped tube. The four (rarely five) petals are free or basally fused (Salvadora), usually with three or four teeth or lobes and often with a scale-like gland on the inner side. The four (rarely five) stamens are free, fused into a tube sometimes formed with the petals, with dorsifixed anthers that open via lengthwise slits. Connectives often have a point at the apex. The superior ovary is composed of two carpels that form one or two locules and a short style topped

Salvadora persica, near Malindi, Kenya [298]

Emblingia calceoliflora, Western Australia (KD) [299]

Uses: Even though concentrations of salt are high in Batis, the plants can be consumed by humans as a salad. Stems and leaves can also be pickled. Native Americans used to chew the roots, and a beverage was prepared from it. The seeds are a good source of nutrition, being high in protein, oil and starch, a useful plant when stranded on an uninhabited tropical island beach!

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EUDICOTS

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Phylogeny and evolution: In the past, Salvadoraceae were sometimes associated with Celastraceae, although this is unlikely on morphological grounds. Because of their tetramerous flowers and glucosinolates the family was placed in Brassicales, which was corroborated by molecular studies. Salvadoraceae are closely related to Bataceae, with which they share many characters. Genera and species: Salvadoraceae include three genera and ten species: Azima (4), Dobera (2) and Salvadora (4). Uses: The fibrous stems of Salvadora, and to a lesser extent of Dobera, are used as toothbrushes in Africa and Middle East, where they are called miswak or siwak. Twigs of Salvadora persica are antibacterial and commercially marketed as natural toothbrushes. Additional uses are the edible fruit, which is said to have aphrodisiac properties, the young shoots, which can be eaten as salad, and the leaves, which are a good source of food for camels. Dobera and Salvadora are also a source of essential oils used in the treatment of rheumatism, and leaves of both genera can be burned to extract salt.

Emblingia calceoliflora, Western Australia (KD) [299]

BRASSICALES

EUDICOTS

Etymology: Salvadora is named in honour of Catalan apothecary, botanist and plant collector Juan Salvador y Bosca (1598–1681).

299. EMBLINGIACEAE Slippercreeper family

when originally described, studies of palynology, morphology and anatomy resulted in controversy. Pollen resembles that of Polygalaceae, floral characters suggested affinities with Sapindaceae or Goodeniaceae and vegetative anatomy suggested affinities with Goodeniaceae or possibly Polygalaceae. Molecular studies have clarified the situation to some extent, placing Emblingia firmly in Brassicales, but relationships with other families in this order remain unresolved. Genera and species: This family encompasses a single genus and species: Emblingia calceoliflora.

This family includes prostrate, herbaceous shrubs with long and short shoots covered in stiff hairs. More-or-less opposite, simple leaves have minute stipules and short petioles. Blades have entire, cartilaginous margins and obscure pinnate venation. Inflorescences are reduced to axillary flowers that are bisexual, strongly zygomorphic and held upside-down (resupinate). The five sepals are densely hairy and fused for about half the length, apart from the upper side where the calyx is split to the base. The two petals are hooded, slipperlike and hairy. The receptacle is enlarged into a flattened, linear-curved stalk, bearing stamens and ovary (androgynophore), which emerges through the calyx slit, has flaring lobes at the top and is overtopped by the hooded petals. Eight or nine stamens (of which four are fertile on one side, the rest staminodial) are placed in a spreading ring at the top of the androgynophore. The superior ovary is situated in the staminal ring and composed of one or three carpels, each forming a winged locule, topped with a single hollow stigma. Fruits are one-seeded indehiscent, dry capsules. Distribution: Emblingiaceae are distributed along the western coast of Western Australia, where plants emerge following fires and disappear two or three years later. They are only infrequently encountered as a result. Phylogeny and evolution: Although Emblingia was referred to Capparaceae

Etymology: Emblingia is named for English physician Thomas Embling (1814–1893), who introduced many ‘useful’ animals and plants to the Colony of Victoria (often with disastrous results). He proposed for instance the introduction of camels for the exploration of the deserts of Australia. He had an interest in the humane treatment of prisoners and, after migrating to Melbourne, became a senator in the parliament of Victoria.

300. TOVARIACEAE Stinkbush family

Tovaria pendula, Copenhagen Botanical Garden, Denmark [300]

clawed. A nectary disk is present around the stamen filaments. The usually eight stamens have hairy filaments that are sometimes basally expanded and unequal in length. Anthers are basifixed, arrow-shaped with the apex sometimes pointed and the thecae opening by lengthwise slits. The superior ovary is sometimes stalked and unilocular, but partially separated into four, six or eight chambers, the placentas protruding as arms into the locule. A stout or nearly absent style bears a six- to eight-rayed, dark stigma. Fruits are berries with a persistent stigma. Distribution: Tovariaceae are found in tropical mountain forests in Mexico, Mesoamerica, Jamaica and northern South America following the Andes south to Bolivia.

These are upright and sprawling, perennial herbs and weak shrubs, sometimes somewhat tree-like, with a foetid odour (glucosinolates) and clear sap. Leaves are alternate and trifoliately compound with minute stipules. Leaf lets are nearly sessile with entire margins and pinnate venation. Inflorescences are lax, pendulous, terminal or axillary, bracteate racemes. Flowers are bisexual and actinomorphic. The usually eight (sometimes six to ten) caducous sepals are free, and the eight (sometimes six to nine) free petals are

Phylogeny and evolution: Tovaria was previously considered a member of Capparaceae but more frequently has been classified in its own family, which is corroborated by morphology and phylogenetic analyses based on DNA evidence. It is closely related to Resedaceae. Genera and species: Tovariaceae have a single genus Tovaria, with two similar species: Tovaria diffusa and T. pendula. Etymology: Tovaria is named for 18th century Spanish physician Simón Tovario.

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301. PENTADIPLANDRACEAE Oubli family

EUDICOTS

Distribution: Pentadiplandraceae are only found in tropical West Africa, from Nigeria to Angola.

302. GYROSTEMONACEAE Buttoncreeper family

Phylogeny and evolution: Pentadiplandra was originally placed in Tiliaceae and then in Capparaceae before being transferred to its own family. Its exact position in Brassicales is not fully resolved, although it may be most closely related to Tovariaceae, with which it shares many characters. Oubli or j’oublie is a hairless shrub that sometimes develops into a liana with a tubar underground. It has simple, alternate leaves with minute stipules and entire leaf blades with pinnate venation. Inflorescences are axillary, terminal, short racemes, in which each flower is subtended by leaf-like bracts. The bisexual or unisexual flowers are actinomorphic. The five sepals are nearly free and somewhat inflated at the base. The five free petals are basally thickened, and the bases are covered by woolly hairs. Stamens and ovary are elevated on a stout, stalk-like receptacle (androgynophore). The ten (rarely nine to 13) stamens are placed along the rim of the androgynophore and have filaments fused at the base. Anthers are basifixed and open by lengthwise slits. Stamens are sterile (staminodes) in female flowers. The superior ovary is placed amid the sterile stamens (staminodes) and composed of four or five carpels, each forming a locule. The entire style is topped with a four- or five-lobed stigma. Fruits are sweet, round berries.

Genera and species: Pentadiplandra brazzeana is the sole species of Pentadiplandraceae. Uses: The berries contain a sweet pulp, making them a popular snack with local people. The sweetness is due to a proteinbased compound, brazzein, which has been patented as a sweetener many times sweeter than sugar. This compound can now be produced in transgenic maize and bacteria, opening up possibilities for use in foodstuffs. By using lactic acid, bacteria that have been altered to produce brazzein can be used to produce sugar-free, sweet yoghurts. Other local uses of Pentadiplandra are as a medicine, fish poison, aphrodisiac and condiment. Etymology: Pentadiplandra is composed of the Greek words πέντε ( pente), five, διπλό (diplo), double and άνδρα (andra), a man or stamen, in reference to the usually ten stamens.

Pentadiplandra brazzeana, Equateur Tersoonia cyathiflora, female flower, Province, DR Congo (AK) [301] Mt Benia, Western Australia [302]

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These are short-lived annual shrubs and small trees, often with succulent, rubbery stems with soft wood and usually orange, red or brown branchlets. Leaves are alternate, simple, sessile or petiolate and have small stipules. Blades are entire, succulent and pinnately veined or have only a midvein visible. Inflorescences are axillary or terminal racemes or spikes, or the flowers are solitary. The unisexual flowers are (nearly) actinomorphic and usually on separate plants (male and female plants of the same species can be so distinct that they have previously been thought to be different taxa). Sepals are fused to form a four- or five-lobed broad cup surrounding a fleshy disk. Petals are absent. Male flowers have seven to 100 stamens in one or several series, the anthers almost sessile and filaments short, basifixed and opening by lengthwise slits. Female flowers have a superior ovary composed of one to numerous carpels that are either fused around a central column or entirely fused, each carpel with a stigmatic style. Fruits are

Gyrostemon ramulosus, male flowers, near Geraldton, Western Australia [302]

Codonocarpus cotinifolius, Pilbara, Western Australia [302]

BRASSICALES

EUDICOTS

Stixis ovata, Laos (ZZ) [303]

Reseda luteola, Wavrons, France [303]

Stixis sp., in fruit, Laos (ZZ) [303]

Reseda phyteuma, near Ronda, Spain [303]

succulent, later dry schizocarps, breaking radially into separate, one-seeded nutlets (mericarps) or hard achenes. Seeds are arillate and usually germinate in response to fire. Distribution: Gyrostemonaceae are restricted to Australia, where they occur in drier regions of most of Australia, except the monsoonal north and temperate southeast, but they do occur in Tasmania. Phylogeny and evolution: Gyrostemonaceae have been variously placed, sometimes in Phytolaccaceae or even in Sapindales, but molecular analyses indicated that they belong in Brassicales, which agrees with the presence of glucosinolates. In Brassicales they

Forchhammeria watsonii, Sonora, Mexico (SC) [303]

are most similar and sister to Resedaceae, although characters have yet to be identified that uniquely link these two families. Similarities between their fruit and that of Forchhammeria (Resedaceae) are obvious. Fossils of Gyrostemonaceae have been found from the Miocene of New Zealand. Tersonia is possibly embedded in Gyrostemon. Genera and species: Gyrostemonaceae include five genera with 18 species: Codonocarpus (3), Cypselocarpus (1), Gyrostemon (12), Tersonia (1) and Walteranthus (1). Etymology: Gyrostemon is composed of the Greek words γύρος (gyros), round, and στημων (stemon), a stamen or thread.

303. RESEDACEAE Mignonette family

This is a family of annual, biennial and perennial herbs, shrubs, vines and trees that produce glucosinolates. Leaves are usually alternate, sometimes in a basal rosette, fascicles or rarely opposite (Borthwickia), Plants of the World

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BRASSICALES with rudimentary stipules modified into glands or spines or absent. Blades are simple or trifoliate with pinnate venation and entire, toothed, crenate or pinnately lobed margins. Inflorescences are terminal, sometimes axillary, bracteate, racemes or spikes, or flowers are solitary in the leaf axils (Neothorelia). Flowers are usually bisexual, sometimes unisexual and strongly zygomorphic to almost actinomorphic. The two to eight free or fused sepals are equal or highly unequal, rarely fused and forming a tube that ruptures on one side when the flower opens (Borthwickia). The two to eight petals are free or fused, usually unequal in shape and size (the superior petal usually larger, clawed and often with incised margins) or equal and sometimes with a ciliate margin. Petals are rarely absent. A nectar disk is often asymmetrical and formed between the stamens or absent. The gynoecium/androecium is usually stalked (androgynophore), often hidden by the nectary disk. Three to 70 stamens have free or basally fused filaments and basifixed anthers that open by lengthwise slits. The superior ovary is composed of two to eight free or basally fused carpels, each with an apical stigma. Fruits are capsules, unfused follicles that open apically or fleshy drupes. Distribution: A widespread but patchily distributed family in North America, Mesoamerica, the Antilles, throughout central, western and southern Europe, east to the Baltic states and the Caucasus, northern, eastern and southern Africa, Anatolia, Arabia, southwestern and Central Asia, South Asia, southwestern China, Southeast Asia and western Malesia (Sumatra, Borneo, Philippines). Reseda has been introduced in South America and Australia. Phylogeny and evolution: Resedaceae i n clu d e Borthwickia (s o m e t i m e s Borthwickiaceae), Neothorelia, Stixis, Tirania; (sometimes Stixidaceae), Forchhammeria (formerly Capparaceae) and maybe also Gyrostemonaceae with which some Resedaceae share some characters. Separation of these families was based on morphological

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EUDICOTS

grounds, but in molecular studies no resolution was obtained between Stixidaceae, Resedaceae s.s. and Forchhammeria, and therefore APG IV has been conservative. Borthwickia, Forchhammeria, Neothorelia, Stixis and Tirania are included in Resedaceae to prevent unnecessary inflation of families in Brassicales. Nevertheless, there are many shared characters between some taxa of Resedaceae and Gyrostemonaceae, and the two may not warrant recognition in the long run. Forchhammeria, for instance, has an irregular number of sepals and lacks petals, as do Gyrostemonaceae. Oligomeris is embedded in Reseda, and the two may have to be merged despite their many morphological differences. Further studies with more complete sampling of the taxa will be needed to ascertain proper generic limits. It is possible that only seven or eight genera should be recognised in total. Oligomeris linifolia has a disjunct distribution, found in arid regions of the Old World and southwestern USA and Mexico; it has also been reported from southern China. This disjunction has been suggested to result either from ancient dispersal or from more recent human introduction into the New World. However, genetic variation in this species indicates that its occurrence in the New World was due to relatively recent but natural long-distance dispersal, despite the lack of obvious seed dispersal mechanisms in this species. Genera and species: Resedaceae are a family of eight to 12 genera and c. 90 species: Borthwickia (1), Caylusea (3), Forchhammeria (10), Homalodiscus (2), Neothorelia (1), Ochradenus (4), Oligomeris (3), Randonia (1), Reseda (c. 55), Sesamoides (1), Stixis (7) and Tirania (1). Uses: Mignonette (Reseda odorata) provides oil used in perfume and is an ornamental. Dyer’s rocket or weld (R. luteola) is the source of an orange-yellow dye used since antiquity. Etymology: Reseda is the Latin name from the plant derived from resedo, I heal, in reference to its medicinal properties.

304. CAPPARACEAE Caper family

These are shrubs and trees usually without spines, rarely thorny, producing glucosinolates. Leaves are alternate and arranged spirally or in a plane (distichous). They are simple or trifoliate to palmately compound, with scalelike or absent stipules and petioles often with nectaries and sometimes with petiolar spines. Blades have an entire margin and pinnate venation. Inflorescences are terminal or axillary racemes, sometimes corymbs or solitary flowers, without bracts. Flowers are usually bisexual and actinomorphic or (slightly) zygomorphic. The four, free sepals are usually persistent, and the four petals are free and equal in size. Nectary-disks or glands are often present, and the receptacle sometimes forms a rim on the inside (corona). Stamens and pistils are usually on a stalk (androgynophore). The six to numerous (up to 250) stamens have free filaments and basifixed or dorsifixed anthers that open by longitudinal slits. The ovary is sometimes stalked (gynophore) and composed of a single carpel that forms two locules. The style is relatively short and thick and topped by a single, capitate stigma. Fruits are capsules or berries that often open by two lateral valves. Distribution: Capparaceae are a pantropical family extending into the temperate zones in North America, Mediterranean Europe and North Africa, western and Central Asia and Australia. They are prominent in seasonally dry forests. Phylogeny and evolution: Capparaceae and their close relatives Cleomaceae and Brassicaceae, emerged during the Miocene. In their original circumscription,

BRASSICALES

EUDICOTS

Euadenia eminens, Royal Botanic Gardens, Kew, UK [304]

Steriphoma cleomoides, Ruissalo Botanical Garden, Turku, Finland [304]

Capparaceae were found to be polyphyletic. Tribe Stixideae and Forchhammeria have many characters that distinguish them from this concept and were found to be closer to Resedaceae in molecular studies. Also Cleome (Cleomaceae), Koeberlinia (Koeberliniaceae), Pentadiplandra (Pentadiplandraceae), Setchellanthus (Setchellanthaceae) and Tovaria (Tovariaceae) were previously included in Capparaceae, but these were found to belong elsewhere in Brassicales. Capparaceae in a broader sense have been previously considered to include two subfamilies, Capparoideae and Cleomoideae, based on flower and fruit structure. Molecular and morphological analyses of the family revealed that this circumscription of Capparaceae is paraphyletic, with the larger Brassicaceae embedded within it, but monophyly of each of the three clades was strongly supported. Therefore, three families, Brassicaceae, Capparaceae and Cleomaceae, are accepted rather than merging the three families into one as done in earlier APG systems, even though they share many morphological characters. Placement and circumscription of some genera are still uncertain, and the genera accepted below are tentative, pending further taxonomic study. Keithia brasiliensis is probably not part of Capparaceae because it has five fused petals that have glands at their base. Where it does belong is unclear. Similarly Poilanedora unijuga has five petals, five sepals and an intrastaminal disc, making this a poor fit in this family.

Capparis flexuosa, Copenhagen Botanical Garden, Denmark [304]

Genera and species: Capparaceae include c. 30 genera and about 324 species: Anisocapparis (1), Apophyllum (1), Atamisquea (1), Bachmannia (1), Beautempsia (1), Belencita (1), Boscia (c. 30), Buchholzia (2), Cadaba (c. 30), Calanthea (10), Caphexandra (1), Capparicordis (3), Capparis (c. 250), Cladostemon (1), Colicodendron (14), Crateva (8), Cynophalla (21), Dhofaria (1), Euadenia (4), Hispaniolanthus (1), Maerua (c. 100), Mesocapparis (1), Monilicarpa (2), Morisonia (5), Neocalyptrocalyx (7), Preslianthus (3), Ritchiea (c. 15), Sarcotoxicum (1), Steriphoma (8) and Thilachium (c. 13). Unplaced: Keithia (1) and Poilanedora (1). Uses: Flower buds of Capparis species, either pickled or salted, are the capers of commerce. Pickled fruits and stems of some species are also eaten, notably in southeastern Spain. Much of the material harvested for commerce is gathered from plants in the wild, although there are several areas where plants are cultivated. Taxonomy of the taxa yielding capers is complex, and the main source, Capparis spinosa, may be of hybrid origin. It is seldom truly found in the wild, usually associated with archaeological sites or ancient agricultural landscapes, and may be a cultigen derived from C. sicula with some introgression from C. orientalis. Fruits of the bumble tree (C. mitchellii) from Australia are also edible. Some larger tree-like species of Capparis (e.g. C. mitchellii, C. nobilis) have a hard wood similar to that of box (Buxus,

Capparis spinosa, Syracuse, Sicily, Italy [304]

Buxaceae). Species of Steriphoma and Capparis are occasionally grown as garden ornamentals in the tropics. Etymology: Capparis is derived from the Greek κάππαρης (kapparis), caper, the classical name of the plant, its preserved buds and fruit. The genitive singular of this in Latin is ‘capparis’, and thus the correct family name is Capparaceae, not Capparidaceae as sometimes spelled.

305. CLEOMACEAE Spiderflower family

Cleomaceae are annual and perennial herbs and shrubs that produce glucosinolates and are glabrous or with glandular hairs. Leaves are alternate and spirally arranged, the blades simple or (usually) palmately compound with entire, serrate, or serrulate margins and pinnate venation. Stipules are usually present and variable in shape, often three to eight-palmatifid, linear, thread-like, scalelike, spine-like or (rarely) absent. Petioles usually have a thickened part (pulvinus) and

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BRASSICALES

Cleome graveolens, Ruissalo Botanical Garden, Turku, Finland [305]

Cleome serrulata, New Mexico, USA (DZ) [305]

Isomeris arborea, Palm Springs, California, USA

sometimes petiolar spines. Inflorescences are terminal or axillary corymbose racemes, or flowers are solitary, often with unifoliate or trifoliate bracts. Flowers are usually bisexual (sometimes functionally unisexual in part of the inflorescence), actinomorphic or zygomorphic. The four free or basally fused sepals are sometimes persistent. The four free petals are equal or unequal. A rim or corona formed from the receptacle is sometimes present, and nectary disks, scales or glands also may be present. The androecium/gynoecium is usually stalked (androgynophore). Usually six to 27 (rarely fewer or up to 35) stamens have filaments that are free or fused at the base to the ovary stalk (gynophore). Anthers are basifixed or dorsifixed and open by longitudinal slits. The superior ovary is stalked (gynophore) or not and composed of a single carpel forming two locules tipped with a single straight, thick style with a single, capitate, unlobed stigma. Fruits are capsules opening by two valves or nutlets, rarely inflated schizocarps (Isomeris), or samaras (Dipterygium).

extending into temperate North and South America, Europe, Central and East Asia and Australia. They are most diverse in the Americas.

productivity. The relatively close relationship to Brassicaceae (which includes the genetic model species, Arabidopsis thaliana) provides an opportunity for Cleome to be used in clarifying the genetic control of C4 photosynthesis.

Distribution: Cleomaceae are widespread in the tropics and warm temperate regions 414

EUDICOTS

Christenhusz, Fay & Chase

Phylogeny and evolution: Delimitation of genera in this family has been in great turmoil because Cleome in the traditional sense has been found to be polyphyletic with regard to other segregates, such as Arivela, Carsonia, Cleoserrata, Dactylaena, Dipterygium, Gynandropsis, Hemiscola, Melidiscus, Physostemon, Podandrogyne, Polanisia and Tarenaya, to name a few. Many segregates have been proposed, but not all combinations have been made, causing nomenclatural instability. It is therefore advisable to maintain Cleome in the broad sense and include the segregates mentioned above in it. Inclusion of Dipterygium is problematic because it was considered intermediate between Brassicaceae and Cleomaceae. In the latter it was usually maintained as a separate subfamily because of its winged, indehiscent fruits. Cleome spans a developmental progression from species with C3 photosynthesis through others carrying out C4 photosynthesis, which allows greater water- and nitrogen-use efficiency and higher

[305]

Genera and species: Cleomaceae include nine or fewer genera, most are now merged with Cleome and probably most, if not all, should be united into a single genus. There are a total of c. 346 species: Cleome (c. 320), Cleomella (c. 10), Cristatella (2), Haptocarpum (1), Isomeris (1), Oxystylis (1), Peritoma (6), Puccionia (1) and Wislizenia (3). Uses: African cabbage or cat whiskers (Cleome gynandra, synonym: Gynandropsis gynandra) is used as a leaf vegetable in South Africa. Some species are popular bedding plants, such as spider flower (C. sesquiorygalis, synonym: Tarenaya hassleriana), which is also used as a cut flower. Cleome viscosa seeds are sometimes added to curries in Sri Lanka as a cumin substitute. Etymology: Cleome is probably derived from Greek κλέος (kleos), glory.

BRASSICALES

EUDICOTS

306. BRASSICACEAE Cabbage family

Brassicaceae are annual, biennial and perennial herbs, shrubs and rarely small trees and vines (e.g. Cremolobus, Heliophila) that produce glucosinolates (mustard oils). They grow usually terrestrially, but are occasionally (partially) submerged aquatics. Leaves are alternate or rarely opposite or whorled, sometimes all in a basal rosette. They lack stipules but often have tiny, stipule-like glands at the base of the petioles and pedicels, and blades are usually simple, sometimes pinnatifid or trifoliolate or pinnately, palmately or bipinnately compound. Leaves are usually petiolate, sometimes sessile or subsessile

Matthiola crassifolia on the ruins of Byblos, Lebanon [306]

and then sometimes amplexicaul or clasping the stem with auricles. Margins are entire, dentate, crenate, sinuate, repand or dissected, and venation is pinnate, rarely palmate. Inflorescences are terminal racemes that are often corymbose or paniculate, or flowers are solitary on pedicels from axils of rosette leaves, usually without bracts. Flowers are usually bisexual and mostly actinomorphic, occasionally (somewhat) zygomorphic. The four free sepals are in two opposite pairs and saccate at the base or not. Petals are free in two pairs of two (cruciform), alternating with the sepals, and a petal claw is usually differentiated with the petal blade sometimes reduced and much smaller than the claw. Petals rarely have a basal appendage, and margins can be entire or emarginate to bifid, rarely fimbriate or filiform. Petals are rarely rudimentary or absent. Usually six (rarely two to 24) stamens are placed in two whorls, often the outer pair shorter than the inner two pairs, rarely equal in length or in three pairs of unequal lengths. Free, rarely fused filaments are simple or (rarely) winged, toothed or with appendages and nectar glands at the base. Anthers are basifixed or

Bunias orientalis, Turku, Finland [306]

dorsifixed and open towards the inside by lengthwise slits. The superior, usually sessile ovary is composed of two fused carpels, each forming a locule with a false septum connecting the two (rarely the septum absent and forming a single locule). The single style has a capitate or conical, entire or bilobed stigma, rarely elongated into horns or spines. Fruits are typically a bivalved capsule that opens from below and are called a silique or silicle (depending on length-width ratio). Fruits are sometimes indehiscent nutlets, schizocarps, samaras or lomenta, sometimes with a fruit stalk (carpophore). Distribution: A nearly worldwide family (but absent from parts of tropical America, Africa, Asia, Madagascar and Australia), and most diverse in temperate, montane and Arctic areas, with the highest diversity in the Irano-Turanian region, the Mediterranean Basin and western North America. Phylogeny and evolution: Brassicaceae have always been regarded as a natural group, but in the 1990s based on a morphological and an early molecular study, it was

Hornungia alpina, private garden, Kingston upon Thames, Surrey, UK [306]

Plants of the World

415

BRASSICALES

Brassica nigra, naturalised along the Californian coast at Point Bonita, USA [306]

recommended that Brassicaceae and Capparaceae (including Cleomaceae) be united in a single family, Brassicaceae, a suggestion that was followed by APG. Subsequent studies proposed that three closely related families should intead be recognised, with Brassicaceae sister to Cleomaceae, and both sister to Capparaceae, which is now the standard followed. Tribal classification of Brassicaceae has been subject to controversy, and the early system, based on fruit morphology and sepal orientation, was highly artificial. Convergence is common throughout the family, and most morphological characters evolved independently in separate groups. This complicates generic delimitation in Brassicaceae because most genera are distinguished primarily by fruit characters. Because of the incomplete molecular knowledge of many genera, delimitations have fluctuated greatly in recent times. Several molecular studies have demonstrated that there is a major split in Brassicaceae, corresponding to the Mediterranean and southwestern Asian tribe Aethionemeae and the rest of the family. The tendency in Brassicaceae has been to 416

Christenhusz, Fay & Chase

EUDICOTS

Descurainia pinnata, Manatee County, Florida, USA [306]

split rather than to lump genera when nonmonophyly has been encountered, although in desperation one previous author (Krause, 1902) concluded that all might be better merged into the single genus Crucifera (a later synonym that he published). This relatively unstable taxonomy, combined with wide convergence of characters and unworkable original circumscriptions of genera created an unwieldy number of genera; one solution to these problems would be treat current tribes at the genus level. However, not all currently recognised genera are placed, and more work will be needed to establish all relationships. Many realignments following molecular studies have taken place. The genera of tribe Brassiceae, which includes most of the commercial crops, are over-split, and some merging at the generic level seems to be indicated, especially because genera such as Brassica, Diplotaxis and Erucastrum are polyphyletic. For the list below, we followed Al-Shehbaz (2012), with species numbers from BrassiBase (Kiefer et al., 2014), in the hope that this will suffice for the moment.

Noccaea caerulescens, Helsinki Botanical Garden, Finland [306]

Genera and species: Brassicaceae are by far the largest family of Brassicales with c. 343 genera and about 3,630 species: Abdra (2), Acuston (1), Aethionema (45), Alliaria (1), Alyssoides (1), Alyssopsis (2), Alyssum (c. 100), Ammosperma (2), Anastatica (1), Anchonium (2), Andrzeiowskya (1), Anelsonia (1), Anzhengxia (1), Aphragmus (11), Aplanodes (2), Arabidella (7), Arabidopsis (10), Arabis (60), Arcyosperma (1), Armoracia (3), Aschersoniodoxa (4), Asperuginoides (1), Asta (1), Atacama (1), Atelanthera (1), Athysanus (2), Atropatenia (2), Aubrieta (12), Aurinia (7), Baimashania (2), Ballantinia (1), Barbarea (22), Berteroa (5), Biscutella (45), Bivonaea (1), Blennodia (2), Boechera (110), Bornmuellera (9), Borodinia (8), Borodiniopsis (1), Botschantzevia (1), Brachypus (1), Brassica (38), Braya (25), Brayopsis (7), Bunias (2), Cakile (7), Calepina (1), Callothlaspi (5), Calymmatium (2), Camelina (8), Camelinopsis (2), Capsella (5), Cardamine (c. 200), Carinavalva (1), Carrichtera (1), Catenulina (1), Catolobus (1), Ceratocnemum (1), Chalcanthus (1), Chamira (1), Chartoloma (1), Chaunanthus (3),

BRASSICALES

EUDICOTS

Chilocardamum (4), Chlorocrambe (1), Chorispora (11), Christolea (2), Chrysochamela (3), Cithareloma (2), Clastopus (2), Clausia (4), Clypeola (9), Cochlearia (20), Coincya (6), Coluteocarpus (1), Conringia (6), Cordylocarpus (1), Crambe (35), Crambella (1), Cremolobus (7), Crucihimalaya (11), Cryptospora (4), Cuphonotus (2), Cuprella (2) Cusickiella (2), Cymatocarpus (3), Cyphocardamum (1), Dactylocardamum (1), Degenia (1), Delpinophytum (1), Dendroarabis (1), Descurainia (35), Diceratella (11), Dichasianthus (1), Dictyophragmus (3), Didesmus (2), Didymophysa (2), Dielsiocharis (2), Dilopha (2), Dimorphocarpa (4), Diplotaxis (25), Dipoma (1), Diptychocarpus (1), Dithyrea (2), Dontostemon (10), Douepea (3), Draba (390), Drabastrum (1), Drabella (1), Dryopetalon (8), Eigia (1), Elburzia (1), Enarthrocarpus (5), Englerocharis (4), Eremobium (1), Eremoblastus (1), Eremophyton (1), Eruca (1), Erucaria (10), Erucastrum (25), Erysimum

Armoracia rusticana Helsinki Botanical Garden, Finland [306]

(c. 180), Euclidium (1), Eudema (4), Eutrema (26), Exhalimolobos (9), Farsetia (27), Fezia (1), Fibigia (3), Foleyola (1), Fortuynia (2), Friedrichkarlmeyeria (1), Galitzkya (3), Geococcus (1), Glastaria (1), Goldbachia (7), Graellsia (7), Guiraoa (1), Halimolobos (8), Harmsiodoxa (3), Heldreichia (1), Heliophila (90), Hemicrambe (3), Hemilophia (5), Henophyton (1), Hesperidanthus (5), Hesperis (34), Hirschfeldia (1), Hollermayera (1), Hormathophylla (10), Hornungia (3), Horwoodia (1), Ianhedgea (1), Iberis (30), Idahoa (1), Ihsanalshehbazia (1), Iodanthus (1), Ionopsidium (9), Irania (5), Irenepharsus (3), Isatis (86), Iskandera (2), Ivania (2), Kernera (1), Kotschyella (2), Kremeriella (1), Lachnocapsa (1), Lachnoloma (1), Ladakiella (1), Leavenworthia (8), Leiocarpaea (1), Leiospora (6), Lepidium (c. 250), Lepidostemon (6), Lepidotrichum (1), Leptaleum (1), Leptoplax (1), Lithodraba (1), Litwinowia (1), Lobularia (4), Lunaria (3), Lutzia (1), Lyrocarpa (3),

Eutrema wasabi, Royal Botanic Gardens, Kew, UK [306]

Macropodium (2), Malcolmia (41), Mancoa (8), Masmenia (2), Mathewsia (5), Matthiola (50), Megacarpaea (9), Megadenia (1), Meniocus (7), Menkea (6), Menonvillea (25), Metashangrilaia (1), Micrantha (1), Microlepidium (2), Microstigma (3), Microthlaspi (7), Morettia (4), Moricandia (7), Morisia (1), Mostacillastrum (30), Murbeckiella (5), Muricaria (1), Myagrum (1), Nasturtiopsis (2), Nasturtium (5), Neotorularia (6), Nerisyrenia (8), Neslia (2), Neuontobotrys (13), Neurotropis (3), Nevada (1), Noccaea (85), Noccidium (2), Notoceras (1), Notothlaspi (2), Ochthodium (1), Octoceras (1), Odontarrhena (86), Olimarabidopsis (3), Onuris (6), Oreophyton (1), Ornithocarpa (2), Orychophragmus (2), Otocarpus (1), Pachycladon (12), Pachymitus (1), Pachyneurum (1), Pachyphragma (1), Parlatoria (3), Parodiodoxa (1), Parolinia (5), Parrya (50), Parryodes (1), Paysonia (8), Pegaeophyton (7), Peltaria (3), Peltariopsis (2),

Eruca sativa, Gordes, France [306]

Plants of the World

417

BRASSICALES

EUDICOTS

Cochlearia officinalis, Lizard Peninsula, Cornwall, UK [306]

Pennellia (10), Petrocallis (1), Petroravenia (3), Phlebolobium (1), Phlegmatospermum (4), Phoenicaulis (1), Phravenia (1), Phyllolepidium (2), Physaria (106), Physocardamum (1), Physoptychis (2), Physorhynchus (2), Planodes (2), Polyctenium (1), Polypsecadium (15), Pringlea (1), Pseuderucaria (2), Pseudoarabidopsis (1), Pseudocamelina (3), Pseudodraba (1), Pseudofortuynia (1), Pseudosempervivum (6), Pseudoturritis (1), Pseudovesicaria (1), Psychine (1), Pterygostemon (1), Pugionium (2), Pycnoplinthopsis (1), Pycnoplinthus (1), Quezeliantha (1), Raffenaldia (2), Raparia (2), Raphanorhyncha (1), Raphanus (5), Rapistrum (2), Resetnikia (1), Rhammatophyllum (8), Rhizobotrya (1), Ricotia (9), Robeschia (1), Romanschulzia (14), Rorippa (86), Rudolf-kamelinia (1), Rytidocarpus (1), Sandbergia (2), Sarcodraba (5), Savignya (1), Scambopus (1), Scaphiarabis (4), Schimpera (1), Schizopetalon (10), Schouwia (1), Scoliaxon (1), Selenia (5), Shangrilaia (1), Shehbazia (1), Sibara (13), Sinapidendron (5), Sinapis (5), Sinoarabis (1), Sisymbrella (1), Sisymbriopsis (5), Sisymbrium (41), Smelowskia (25), Sobolewskia (4), Solms-laubachia (33), Sphaerocardamum (4), Spryginia (7), Stanleya (7), Stenopetalum (10), Sterigmostemum (11), Stevenia (8), Straussiella (1), Streptanthus (54), Streptoloma (2), Succowia

418

Christenhusz, Fay & Chase

Vella pseudocytisus, Royal Botanic Gardens, Kew, UK [306]

(1), Synstemon (2), Synthlipsis (2), Takhtajaniella (1), Tchihatchewia (1), Teesdalia (3), Tetracme (9), Thelypodiopsis (7), Thelypodium (16), Thlaspi (2), Thlaspiceras (11), Thysanocarpus (7), Tomostima (6), Trachystoma (3), Transberingia (1), Trichotolinum (1), Tropidocarpum (4), Turritis (2), Vania (4), Vella (7), Veselskya (1), Warea (4), Weberbauera (22), Xerodraba (8), Yinshania (13), Zerdana (1), Zilla (2) and Zuluagocardamum (1). Uses: Many members of Brassicaceae have great economic importance as food, fodder, industrial crops and ornamentals. A good number of species and cultivars are important vegetables, notably Brassica oleracea, a native of coastal Europe and now domesticated with many forms, such as kale (‘Acephala Group’), Chinese kale (‘Alboglabra Group’), broccoli and cauliflower (‘Botrytis Group’), cabbages (‘Capitata Group’), Brussels sprouts (‘Gemmifera Group’), kohlrabi (‘Gongylodes Group’), sprouting broccoli and calabrese (‘Italica Group’) and Portugese cabbage (‘Tronchuda Group’), of which the leaves, compact inflorescences, axillary buds and stems are frequently eaten. Swede or rutabaga (B. napus ‘Napobrassica Group’) has an edible root-like structure that is mostly derived from the hypocotyl (stem

below the seed leaves and above the true root); it originated from a hybrid between cabbage and turnip (B. rapa). European sarson (B. rapa) is an important oilseed in India and best known for producing turnips (‘Rapifera Group’), but also is the origin of a number of Oriental vegetables such as pak-choi and chard (‘Chinensis Group’) and Chinese cabbage (‘Pekinensis Group’), all commonly used in stir-fries. Mizuna and mibuna (‘Japonica Group’) are used as salad greens in Japanese cuisine and komatsuma (‘Perviridis Group’) is rich in vitamins and can be cooked like spinach. Abyssinian mustard or Texsel greens (Brassica carinata) are eaten in northeastern Africa and have high protein content. Japanese greens or senposai (B. juncea var. crispifolia) are often grown for spring greens, especially in South and East Asia. Seakale (Crambe maritima) is sometimes eaten as a vegetable, especially if the stems and leaves have been blanched by heaping stones around the plants to remove the bitterness. Many species are eaten fresh in salads, such as salad rocket (Eruca sativa), popular in Italian cuisine, white rocket (Diplotaxis erucoides), watercress (Nasturtium officinale, synonym: Rorippa nasturtium-aquaticum) and the peppery roots of radish (Raphanus sativus) and its larger

BRASSICALES

EUDICOTS

cultivar from East Asia (daikon or muli, R. sativus ‘Longipinnatus’). Young sprouts of several species are also popular, especially cress (Lepidium sativum), wintercress (Barbarea verna) and mustard cress (Sinapis alba). Young sprouts of Brassica rapa subsp. campestris are commonly eaten with mashed potatoes in spring in the Netherlands where they are called raapstelen. Dittander (Lepidium latifolium) was a salad plant of the Ancient Greeks, but it is largely unknown in cultivation now. Maca (L. meyenii) is a salad vegetable in the Andes, allegedly with aphrodisiac properties, and its dried roots are now exported to China where it has become popular. Turkish rocket (Bunias orientalis) was previously used as a salad and animal fodder. Garlic mustard (Alliaria petiolata) has an onion-like flavour and was formerly used as a garlic substitute. The plant is undervalued, as it is often weedy and invasive, but it has more vitamin A than spinach and more vitamin C than oranges. Also high in vitamin C is scurvy grass (Cochlearia officinalis), which is often prepared as a juice but can also be eaten in salads and was used by the UK Royal Navy to treat scurvy on long sea voyages. Seeds of hoary cress (Lepidium draba) were formerly used as a pepper substitute. Because of their mustard oils, many species are used as condiments, especially mustards, such as white mustard (Sinapis alba, synonym: Brassica hirta) and Chinese mustard (Brassica juncea var. crispifolia), the latter most commonly used for Dijon mustard. Black mustard (Brassica nigra) is also sometimes used, and this was probably the mustard seed of the Bible. Flixweed (Descurainia sophia) was also grown for its mustard-like seeds. A well-known food flavouring is horseradish (Armoracia rusticana), a sterile cultigen originating somewhere in Central Asia or Assyria and propagated through root cuttings of a single clone for over 2,000 years. A similar but much stronger flavour is wasabi (Eutrema wasabi, synonym: Wasabia japonica), a condiment commonly eaten with Japanese raw fish dishes (sushi, sashimi etc.). Rapeseed (Brassica napus subsp.

oleifera) is used as fodder in addition to being an important source of cooking oil, especially canola or Canadian rapeseed (B. napus ‘Canola’). Rapeseed or canola oil is commonly used in salads and cooking, for deep-fried and processed foods, mayonnaise and margarine and as a biofuel. It is also used industrially in soaps, plastics and antifreeze, lubricants etc. False flax (Camelina sativa) was originally domesticated for fibre but was and sometimes is still grown as a substitute for rapeseed oil, called camelina oil. Other minor oil seeds are tansy mustard (Descurainia pinnata) and yellowtop or lesquerella (Physaria fendleri), the latter being a good substitute for castor oil. For a long time, a blue dye from the fermented leaves of woad (Isatis tinctoria) was produced, which was cheaper and easier to make than imported indigo and thus commonly used to dye fabric, linen and wool for tapestries. It was the blue of many national flags in Europe. Some species, such as alpine pennycress (Noccaea caerulescens, formerly placed in Thlaspi) and Pyrenean scurvygrass (Cochlearia pyrenaica), are hyperaccumulators of metals including cadmium and zinc and have potential use in the development of plants to be used in bioremediation of polluted soils. Among the many ornamentals grown in temperate gardens are Aethionema saxatile (burnt candytuft), Alyssum alyssoides (small alison), A. spinosum (spiny madwort), Arabis caucasica (mountain rockcress), Aubrieta deltoidea (purple rockcress), A. ×cultorum (garden aubrieta), Aurinia saxatilis (gold-dust), Cardamine (spinks), Cochlearia acaulis (violet cress), Draba aizoides (yellow whitlow-grass), Erysimum cheiri (wallf lower), Fibigia lunarioides (roman shields), Heliophila longifolia (blue diamonds), Hesperis matronalis (dame’s violet), Hornungia alpina (hutchinsia), Iberis sempervirens (candytuft), I. umbellata (florist’s cress), Lobularia maritima (sweet alyssum), Lunaria annua (honesty), Malcolmia maritima (Virginia stock), Matthiola incana (Brompton stock), M. longipetala subsp. bicornis (night-scented stock), Moricandia ramburii (violet cabbage),

Parrya nudicaulis (nakedstem wallflower) and Schizopetalon walkeri (snowflake stock), to list a few. Genetic model: Thale cress (Arabidopsis thaliana) has developed notoriety as a genetic model species used to study many aspects of plant biology. It has a small genome, and this allows it to be a successful weed, going through several generations in a single growing season. Its small genome and relationships to important crop plants were among the reasons for its selection as the first plant for which the genome was sequenced, although it has not yet been fully completed (even though claims have been made in this respect). Availability of the sequences for at least the vast majority of the genome is an extremely valuable resource for plant geneticists to improve understanding of plant and genomic function. Carnivory: Carnivorous seeds are perhaps a strange concept, but tests have been carried out on the seeds of shepherd’s purse (Capsella bursa-pastoris). In these studies, it was shown that the mucilage layer of seeds upon imbibition contains chemicals that attract soil nematodes, protozoa and soil bacteria and a toxin that kills these organisms. The mucilage also produces proteinases, and labelled amino acids were found to be absorbed by the seeds. It is easy to imagine that seedlings would benefit nutritionally from the breakdown of these organisms. Record: At 7,756 m in the Himalayas, Solms-laubachia himalayensis is one of the three seed plants that occurs at the highest elevation. The other high-elevation species are members of Ranunculaceae (Ranunculus lobatus) and Caryophyllaceae (Arenaria bryophylla). Etymology: Brassica is the classical Latin name for cabbage. The family is also sometimes called Cruciferae, which is not based on the genus Crucifera (a much later name) and is therefore better avoided. Cruciferae means cross-bearers, in reference to the four-petalled flowers.

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419

BERBERIDOPSIDALES

EUDICOTS

BERBERIDOPSIDALES Families 307 and 308 make up the small order Berberidopsidales. This is putatively the first branching clade in the larger clade including the asterids. An age of c. 113 million years is estimated. The spirally arranged perianth with variable numbers of parts in Berberidopsis is reminiscent of some flowers of Ranunculales.

307. AEXTOXICACEAE Olivillo family

These are unisexual trees with the branches, lower leaf surfaces and inf lorescences covered in rust-coloured scales. Leaves are nearly opposite and have a simple, entire blade with a pinnate venation are often slightly peltate at base; they lack stipules. Inflorescences are racemes and are clustered in groups of three or more in leaf axils. Inflorescence bracts are small and round. Flowers are unisexual, actinomorphic and enveloped in bud by a bract. The four to six spirally arranged sepals are round, free and thin, the usually five (sometime four to

six) free spirally arranged petals are broadly clawed and have a thick midrib. Male flowers have five stamens and kidney-shaped nectar glands. Anthers are dorsifixed and open by short apical slits. Female flowers have five f leshy sterile stamens (staminodia) that alternate with nectar glands, the superior ovary is composed of a single carpel and covered with large scales. The short apical style is bent to one side and appressed to the ovary. Fruits are dry one-seeded indehiscent capsules.

Christenhusz, Fay & Chase

308. BERBERIDOPSIDACEAE Coral-vine family

Distribution: Aextoxicaceae are found around the Andean Lake District and coastal region of central and southern Chile and adjacent Argentina. Phylogeny and evolution: The position of this family has long been disputed, but molecular analyses found them sister to Berberidopsidaceae. Genera and species: Aextoxicon punctatum is the only species in this family.

Aextoxicon punctatum, Royal Botanic Gardens, Kew, UK [307]

420

Etymology: Aextoxicon is composed of the Greek words αίξ (aix), a buck, and τοξικών (toxicon), poisonous, because this species is poisonous to goats.

This is a family of woody vines with spirally arranged leaves without stipules. Leaf blades are simple, three- to five-palmately veined at base with the margins entire or coarsely toothed. Petioles often have a swollen pulvinus. Inflorescences are axillary racemes, or flowers are solitary in the leaf axils. The bisexual, actinomorphic flowers are pendent.

Aextoxicon punctatum, male flowers, Chile (IV) [307]

SANTALALES

EUDICOTS

shaped stigma. The fruits are berry-like.

The perianth is spirally arranged and either has five distinct sepals and five distinct petals (Streptothamnus) or the sepal-like tepals gradually integrating into the petal-like ones (Berberidopsis). Stamens are placed either in a single whorl of six to 13 or numerous and densely packed. Filaments are free, and the anthers are (nearly) basifixed, opening by longitudinal, lateral slits. The superior ovary is unilocular and topped with a club-shaped style and an unremarkable or mushroom-

Phylogeny and evolution: Originally placed in Flacourtiaceae, members of this family were placed as sister to Aextoxicaceae in early molecular analyses. The family is probably c. 100 million years old. Wood anatomy supports inclusion of this family with Aextoxicaceae.

Berberidopsis beckleri, Royal Botanic Gardens, Kew, UK [308]

Berberidopsis beckleri in fruit, Royal Botanic Gardens, Kew, UK [308]

Distribution: They are disjunct between southern Chile and eastern Australia.

Genera and species: A family of two genera with three species: Berberidopsis beckleri, B. corallina and Streptothamnus moorei. Uses: The coral vine, Berberidopsis corallina, is sometimes planted as an ornamental evergeen vine in mild temperate gardens. Etymology: Berberidopsis refers to the similarity of the leaves of B. corallina to some Berberis (Berberidaceae) species. Berberidopsis corallina, Royal Botanic Gardens, Kew, UK [308]

SANTALALES Families 309 to 313 compose the order Santalales, which are one of the largest groups of parasitic flowering plants (the other being Orobanchaceae), but relationships between the families and even which families should be recognised are problems still not fully resolved. Some more recent molecular analyses have included different subsets of Santalales, and due to these differences in sampling, it is still not possible to assess all relationships. It seems that Olacaceae form a grade leading up to the rest, with Balanophoraceae embedded in Santalaceae. Here we follow the families provided in APG IV (2016), recognising that some of these may be combined or split further as our knowledge of these plants develops. The crown group of Santalales has been dated to c. 101–108 million years old.

309. OLACACEAE Tallow-wood family

Olacaceae are terrestrial, evergreen shrubs, trees and vines. They are green and autotrophic in many cases, but several members are also parasitic, making haustorial connections with their roots underground. Leaves are simple, alternate and spirally arranged or in a plane (distichous). Petioles are well developed, slender or thickened towards the end, and there are no stipules. Leaf blades have pinnate venation, sometimes with several prominent basal veins where the blades are

sometimes peltate or cordate. Leaf margins are entire or toothed, with the teeth sometimes glandular. Inflorescences are dichotomous cymes, racemes, axillary fascicles, short spikes or simple to compound umbels, sometimes cauliflorous or appearing capitate, with or without bracts. Flowers are bisexual or partially unisexual and actinomorphic. The usually five (rarely three to seven) sepals are fused into a cup-shaped, five-lobed calyx that is persistent in fruit, sometimes absent

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SANTALALES (in some Heisteria). The four or five (rarely three to seven) petals are free or slightly fused at the base, cup-shaped or long-tubular and sometimes hairy on the inside. A nectar disc is absent or present between the stamens and style. The four or five (or eight to ten, rarely up to 20) stamens are in one or two whorls, sometimes partly sterile (all staminodial in female flowers of Octoknema), and filaments are fused to the base of the petals. Anthers are basifixed to (rarely) dorsifixed and open with inward (introrse) or lateral slits, apical pores or valves (trivalvate in Hondurodendron), and the connective is apical or not; anthers can be connate and surround the style or reflexed (Aptandra). The superior or half-inferior ovary is composed of three to five fused carpels forming one to five locules. A style is absent or conical and short, and the stigma is entire or

slightly three- to five-lobed. Fruits are drupes, usually with a cupule formed by the enlarged persistent calyx.

Phylogeny and evolution: We have followed the traditional circumscription for this family (excluding Schoepfiaceae, which are not directly related), even though this set of genera is not monophyletic. Some

authors recognise the seven clades that make up this family as independent families (Aptandraceae, Coulaceae, Erythropalaceae, Octoknemaceae, Olacaceae, Strombosiaceae and Ximeniaceae). Also, relationships among these families are poorly supported, and therefore it is premature to recognise all these clades at the family level. For some families there is also no evidence that they form a clade, and it is likely that in a broad circumscription Olacaceae are polyphyletic. Erythropalaceae and Strombosiaceae may need to be recognised in future classifications. All families of Santalales could instead be placed in a single family. Because of the lack of resolution in this group, age estimates (c. 96 million years has been suggested) are preliminary. A fossil similar to Anacolosa is known from the Upper Cretaceous of Europe,

Erythropalum scandens, Singapore (WA) [309]

Strombosia ceylanica, Singapore (WA) [309]

Aptandra zenkeri, Lope Reserve, Gabon (CD) [309]

Octoknema affinis, Mt Cameroon, Cameroon

Ochanostachys amentacea flowers, Singapore

Ochanostachys amentacea, Singapore (WA)

(CD) [309]

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Distribution: This is a pantropical family found also in subtropical North and South America. A number of species are parasitic in heathlands, where this life-history strategy enables plants to survive under conditions of low soil nutrients and water. Parasitic Olacaceae are capable of attaching haustoria to root systems of widely different hosts during dry periods, mostly to obtain water.

(WA) [309]

[309]

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Harmandia mekongensis, Laos (YN) [309]

Olax latifolia, Gabon (AP) [309]

Ximenia americana, Singapore (WA) [309]

Ximenia americana in fruit, Singapore (WA) [309]

Olax benthamii, Western Australia (KD) [309]

and this fossil taxon (Anacolosidites) became more widespread during the Eocene.

varnishes, paints etc. It is thick, adhesive, heat resistant and slowly drying. Malania oleifera oil is commercially valuable because it is high in acetylenic fatty acids. Nuts of Coula edulis are commonly eaten by local people in Central Africa. Fruits of pantropical Ximenia americana are eaten in Africa. Leaves and fruits of Erythropalum scandens have a strong garlic scent and are eaten in Southeast Asia. Wood of some species is valued for its scent.

Opiliaceae are evergreen trees, shrubs and (rarely) lianas that are all root parasites forming haustorial connections to neighbouring plants. They have simple alternate leaves, without stipules, arranged in two ranks (distichous). Blades have an entire margin, pinnate venation and cystoliths that look like tubercles when leaves are dried. Inf lorescences are caulif lorous or axillary spikes, catkins, umbels, racemes or panicles with scale-like bracts. Flowers are actinomorphic and bisexual or unisexual and plants then dioecious (Agonandra, Gjellerupia) or gynodioecious (Champereia). Sepals (if present) fused into a tiny cupule with an entire or four- to five-toothed margin, but sepals are often absent and then the perianth is replaced by valvate, fused tepals. The four or five petals are present in bisexual and male flowers and valvate and free or fused at the base. Nectaries alternate with stamens in between the petal lobes and are free or fused and circular or cupular. Stamens are as many as and opposite the petals, free or the filaments fused with the petal bases.

Genera and species: This family consists of 29 genera and 150 species in six putative groups that we treat as subfamilies here: ‘Erythropaloideae’ – Brachynema (2), Erythropalum (1), Heisteria (34) and Maburea (1); ‘Strombosioideae’ – Diogoa (1), Engomegoma (1), Scorodocarpus (1), Strombosia (10), Strombosiopsis (3) and Tetrastylidium (2); ‘Couloideae’ – Coula (1), Minquartia (1), Ochanostachys (1) and Octoknema (14); ‘Ximenioideae’ – Curupira (1), Douradoa (1), Malania (1) and Ximenia (10); ‘Aptandroideae’ – Anacolosa (16), Aptandra (4), Cathedra (5), Chaunochiton (3), Harmandia (1), Hondurodendron (1), Ongokea (1) and Phanerodiscus (3); ‘Olacoideae’ – Dulacia (13), Olax (42) and Ptychopetalum (5). Uses: In West Africa, Ongokea gore is a valuable oil-seed crop called boleco or isano oil, which is used as an additive in making

Etymology: Olax is Latin for odour, smell, in reference to the fragrant wood.

310. OPILIACEAE Bally-coma family

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Champereia manillana, Singapore (KH) [310]

Champereia manillana in fruit, Singapore

Lepionurus sylvestris, Singapore (WA) [310]

Opilia amentacea, Fogg Dam, Northern Territory, Australia (JC) [310]

The superior or half-inferior ovary (sunken in a disk) is composed of two to five carpels forming a single locule. It is topped by a simple style and capitate stigma, or the stigma is sessile and shallowly lobed. Fruits are drupes with three or four linear cotyledons per seed.

Uses: Fruits of Cansjera leptostachya and Opilia amentacea are eaten locally. In Southeast Asia, young leaves and fruits of Champereia manillana and Melientha suavis are sometimes eaten.

Rhizomes grow from a large tuber and are generally branched, usually with scales, warts and lenticels, forming haustoria where they encounter host roots. Shoots are subtended by scale-like or peltate, reduced bracts, the flowering stalks with or without leaves and unbranched. Leaves are scale-like, lack chlorophyll and stomata and are alternate and spirally arranged. Inflorescences are unisexual or contain both male and female flowers in terminal racemes that are reduced to spadices or spadix-like structures covered with minute branches, each frequently subtended by variously modified bracts. Flowers are small (some of the smallest flowers in the angiosperms), unisexual and actinomorphic. Male flowers are larger than female flowers with three or four (sometimes eight) free or (basally) fused sepals, or sepals absent. Petals are missing. When the perianth is absent, there are one or two stamens or as many stamens as sepal lobes when present. Filaments are free or fused into a synandrium, and anthers are free or fused with two or more locules that open by transverse slits. Female flowers are congested. Sepals and petals are absent or are reduced to two tepals that are free or fused into a cup,

(KH) [310]

Distribution: Opiliaceae are a pantropical family, extending into temperate Australasia. The largest genus, Agonandra, is found from Mexico to Argentina, but the other nine genera are all from the Old World tropics.

Etymology: No derivation was given in the original description of Opilia. It has been suggested that it is a cognate with Opilio, a genus of harvestmen, but this seems unlikely.

311. BALANOPHORACEAE Snake-mushroom family

Phylogeny and evolution: Opiliaceae were often associated with Olacaceae in previous classifications, but recent molecular results indicate a closer relationship with Santalaceae, with which they may have to be combined in the future. Genera and species: Opiliaceae include 11 genera and 36 species: Agonandra (10), Anthobolus (3), Cansjera (3), Champereia (1), Gjellerupia (1), Lepionurus (1), Melientha (1), Opilia (2), Pentarhopalopilia (4), Rhopalopilia (3) and Urobotrya (7). 424

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Balanophoraceae are fleshy, unisexual and bisexual, herbaceous parasites without chlorophyll that grow on roots or rhizomes of various hosts, usually trees or shrubs.

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Phylogeny and evolution: Balanophoraceae are strange plants, forming an underground tuber that consists of either parasite or mixed host-parasite tissue and lacking a proper root system. These tubers rupture to leave a circular structure at ground level from which the inflorescence emerges. Inflorescences are racemose, often peculiarly club-shaped, and densely covered in many minute unisexual flowers, in Balanophora so small that they

consist of only 50 cells and are perhaps the smallest flowers of any angiosperm. The leaves when present are arranged spirally or whorled. A relationship between Balanophoraceae and Santalaceae was suggested because of similarities in haustorium structure, and molecular evidence has confirmed this, placing Balanophoraceae inside Santalaceae, close to Nanodea. Because these plants are so strange, there is some aversion to including them in Santalaceae, which would result in splitting of the latter. For the time being, the family is accepted here until further studies show the position of Balanophoraceae more clearly. The floral morphology is often unclear, for instance whether the ovaries are inferior or superior. Only the South African Mystropetalon has a clearly inferior ovary, and this genus may form a clade with Dactylanthus (New Zealand) and Hachettia (New Caledonia), together sister to the rest of the family (and is sometimes treated as a separate family, Mystropetalaceae). Balanophora, Langsdorffia and Thonningia produce a wax-like substance, balanophorin,

Balanophora fungosa, male, China (YN) [311]

Langsdorffia hypogaea, Cordillera del Condor, Ecuador (CD) [311]

which is adnate to the ovary (Mystropetalon). The ovary is inferior (or possibly superior, but this is difficult to see) and composed of two or three fused carpels, each capped by a free style or a single style with a capitate stigma that is sometimes sessile on the ovary. Fruits are achenes, sometimes surrounded by fleshy, fruit-like tissue of the inflorescence, bracts or perianth. Distribution: Balanophoraceae are an almost pantropical family, extending into subtropical and warm temperate South Africa, the Himalayas, China, Japan and New Zealand.

Balanophora fungosa, female, China (YN) [311]

rather than starch as their main food reserve. Cynomorium, previously placed here, has been shown to be unrelated, and is now placed in Cynomoriaceae of uncertain affinity (either Saxifragales or Rosales), Genera and species: Balanophoraceae include 17 genera and c. 44 species: Balanophora (15), Chlamydophytum (1), Corynaea (2), Dactylanthus (1), Ditepalanthus (1), Exorhopala (1), Hachettea (1), Helosis (1), Langsdorffia (3), Lathrophytum (1), Lophophytum (3), Mystropetalom (2), Ombrophytum (4), Rhopalocnemis (1), Sarcophyte (2), Scybalium (4) and Thonningia (1). Uses: Inflorescences of Hachettea, Langsdorffia, Lophophytum and Ombrophytum are locally eaten (South America, New Caledonia). Etymology: Balanophora is composed of the Greek words βάλανος (balanos), an acorn or glans, and φόρος (phoros), to bear, referring to the shape of the male inflorescence. Helosis cayennensis, Mt Matoury, French Guiana (CD) [311]

Hachettea austrocaledonica, Col d’Amieu, New Caledonia (CD) [311]

Scybalium jamaicense, Jamaica [311]

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Cervantesia tomentosa, Pichincha, Ecuador (CD) [312]

Thesium refractum, Yunnan, China

Osyris lanceolata, Yale Rock, Taita Hills, Kenya [312]

Exocarpos sparteus, Flinders Ranges, South Australia [312]

[312]

312. SANTALACEAE Sandalwood family

Santalaceae are a diverse family of green stem-parasites (mistletoes), root-parasitic trees, shrubs and perennial herbs and stem-parasitic vines. Roots are haustorial on branches of the host or roots make haustoria when in contact with host roots, the haustorium bell-shaped and tightly attached to the host. Leaves can be alternate or opposite, rarely subverticillate. They can be persistent or not and well-developed or reduced and scale-like (photosynthesis then carried out by 426

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Comandra umbellata, New Mexico, USA (DZ) [312]

Santalum lanceolatum, near William Creek, South Australia [312]

green stems), lacking cystoliths and stipules. Blades are entire and have pinnate or obscure, parallel venation. Inflorescences are axillary or terminal spikes, cymes, panicles, corymbs, fascicles, umbels or clusters, sometimes the flowers solitary in the leaf axils, without bracts or bracts present and foliaceous or scalelike. Flowers are unisexual or bisexual and actinomorphic. Sepals are generally absent, rarely present as a small rim (calyculus). The three to seven petals are free or fused into a campanulate or urn-shaped tube. A nectary is lobed or absent. Stamens are as many as petal lobes and opposite these with short filaments or the anthers sessile. Anthers are basifixed or dorsifixed and open by longitudinal slits. Sterile stamens and ovaries (staminodia and pistillodia) are sometimes present in functionally unisexual flowers, absent in bisexual or fully unisexual flowers. The halfinferior to inferior ovary is composed of two to six fused carpels forming one locule that is sometimes partially divided or lobed. The

Phoradendron henslowii, Galápagos Islands [312]

Viscum triflorum, Réunion [312]

apical style is simple and topped by a lobed stigma, or the stigma is sessile. Fruits are drupes or false-drupes with a stony seed (in terrestrial taxa), dry nutlets or sticky berries with one or two seeds (in branch parasites). Distribution: This is a widespread family, centred in the tropics. They extend to southern Alaska and Quebec in North America, Terra del Fuego and the Falklands in South America, southern Scandinavia, Siberia and the Russian Far East in temperate Eurasia and south to South Africa and New Zealand. Phylogeny and evolution: Santalaceae here include the formerly separate Viscaceae. To maintain that family, all seven clades would have to be treated as families, adding another five families (Amphorogynaceae, Cervantesiaceae, Comandraceae, Nanodeaceae and Thesiaceae), as done by some authors. Balanophoraceae (probably closely related to Nanodea) will also

SANTALALES

EUDICOTS

render Santalaceae paraphyletic, so if this family is to be maintained Santalaceae will need to be split. Relationships between the various clades are, however, not yet well understood, so we prefer to follow APG IV and keep this larger, easily diagnosable single family. Branch parasites have occurred in this family for at least 72 million years. Herbaceous Thesium probably originated in South Africa c. 60 million years ago, and Santalum, now widespread in the Old World, has a centre of origin in Australia, from where it dispersed across the Pacific and Indian Oceans several times, most likely carried by birds.

– Antidaphne (9), Colpoon (2), Eubrachion (2), Exocarpos (26), Lepidoceras (2), Myoschilos (1), Nestronia (1), Omphacomeria (1), Osyris (4), Rhoiacarpus (1) and Santalum (16); ‘Nanodeae’ – Mida (1) and Nanodea (1).

Genera and species: Santalaceae include 44 genera and c. 1,000 species, here treated as tribes (mostly unpublished): ‘Cervantesieae’ – Acanthosyris (6), Cervantesia (2), Jodina (1), Okoubaka (2), Pilgerina (1), Pyrularia (2), Scleropyrum (4) and Staufferia (1); ‘Thesieae’ – Buckleya (5), Lacomucinaea (1), Osyridicarpos (1), Thesidium (9) and Thesium (340); ‘Comandreae’ – Comandra (1) and Geocaulon (1); ‘Amphorogyneae’ – Amphorogyne (3), Choretrum (7), Daenikera (1), Dendromyza (c. 20), Dendrotrophe (7), Dufrenoya (13), Leptomeria (17), Phacellaria (6) and Spirogardnera (1); ‘Visceae’ – Arceuthobium (29), Dendrophthora (125), Ginaloa (9), Korthalsella (30), Notothixos (8), Phoradendron (c. 250) and Viscum (130); ‘Santaleae’

Uses: Fruits of several species (Buckleya, Geocaulon, Pyrularia) are locally eaten. Quandong (Santalum acuminatum) has a delicious fruit often made into preserves in Australia. Economically, the most important are the fragrant woods of Santalum and Osyris species, particularly Indian sandalwood (S. album), Australian sandalwood (S. spicatum) and African sandalwood (O. lanceolata), which are used for the production of incense and essential oils in the perfume industry mostly from wild but sometimes also from cultivated sources. Poaching of wood from the wild can have a serious impact on populations of these species, and trade is now restricted in East Africa and Australia. Santalum is often cultivated, but host trees (usually Fabaceae) have to be provided in successful plantations. Common mistletoe (Viscum album) has minor economic importance as a Christmas decoration. Kissing under the mistletoe is a remnant of a past in which it was one of the most revered plants of early herbalists and a surviving remnant of northern European plant magic and druidism. It has been used for numerous medicinal purposes and of course also made into a human fertility potion and aphrodisiac. Sticky berries of Viscum were

Misodendrum sp., in fruit, Argentina (SK) [313]

Misodendrum punctulatum on Nothofagus antarctica (Nothofagaceae), Chile (JC) [313]

(and maybe still are) boiled into a glue used to catch songbirds for food in the Mediterranean, a practice that has resulted in the decline of many migratory songbirds in Europe and is now outlawed. Trees infested with mistletoes can be weakened by the parasites, causing damage and reduced yields in fruit orchards and timber plantations. Etymology: Santalum is Latin for sandalwood, possibly originating from Sanskrit candanam, incense wood.

313. MISODENDRACEAE Feathery-mistletoe family

These evergreen, unisexual mistletoes are shrubs that parasitise Nothofagus (Nothofagaceae) branches using a cup-like haustorium. Their leaves are alternate and simple, sometimes reduced to scales, with an entire margin and parallel venation. Inf lorescences are catkins organised in racemes or spikes. The unisexual flowers are actinomorphic, and male flowers lack a

Misodendrum oblongifolium, Argentina (SK) [313]

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SANTALALES perianth but have a small lobed nectar disk. The two or three stamens have basifixed anthers with a single theca that opens by a terminal slit. Female flowers have three tepals that are fused to the ovary. Two or three staminodes are present and enlarge to become feathery in fruit. The half-inferior ovary is composed of three carpels that are fused to form a single locule. The ovary is topped with a short, stout style and a trilobed stigma. Fruits are achenes or nutlets crowned by the feathery staminodes that help in dispersal by wind. Distribution: Misodendraceae are known only from southern beech forests in Chile and Argentina, where they parasitise branches of Nothofagus (Nothofagaceae). Phylogeny and evolution: Misodendrum was often placed in Olacaceae in previous classifications, but molecular studies indicated a relationship to Loranthaceae and Schoepfiaceae. The genus is now treated in its own family. Stem parasitism evolved in this family c. 40 million years ago, before this trait evolved in Loranthaceae. Genera and species: The sole genus of this family is Misodendrum with eight species.

EUDICOTS

Etymology: Misodendrum is composed of the Greek words μίσος (misos), hatred, and δένδρων (dendron), tree.

314. SCHOEPFIACEAE Whitewood family

This is a family of green, parasitic trees, shrubs (Schoepfia, some Arjona) and perennial herbs (some Arjona, Quinchamalium) with haustorial roots. Leaves are simple, alternate and petiolate or sessile, without stipules. Blades are elliptic to linear with pinnate venation and entire margins. Inflorescences are terminal or axillary spikes, axillary cymes or umbels, with bracts that often are fused below the flower to form a lobed, cup-shaped epicalyx. Flowers are bisexual and actinomorphic, usually sessile. Sepals are absent or represented by a short rim fused to the ovary. The four or five petals

are fused to form a urn-shaped or salverform tube with a tuft of hairs on each lobe inside, directly behind each anther. The four or five stamens have filaments fused to the petal tube, sometimes nearly absent in Schoepfia and then the anthers sessile on the tube. Anthers are dorsifixed or basifixed (in Quinchamalium) and open with inward or outward slits. A nectar disk is a ringshaped mound around the base of the style. The inferior ovary is usually composed of three fused carpels forming a single locule (but can be bicarpellate in Schoepfia species with four petals and stamens). A thin and filiform style is placed in the middle of the nectary and topped with a tri- (or bi-)lobed stigma. Heterostyly (with pin and thrum flowers) is found in all genera. Fruits are nut-like achenes. Distribution: This family occurs in tropical America, temperate South America and East Asia from South Korea and Japan south to Sumatra. Schoepfia occurs in Southeast Asia and tropical America, where it extends north to southern Florida (where it is known as Gulf greytwig or whitewood). Arjona and Quinchamalium are both found in temperate South America. Phylogeny and evolution: Schoepfia was previously included in Olacaceae, but it was found to be closer to Arjona and Quinchamalium, formerly in Santalaceae, with which it shares many characters. Molecular analyses place these genera together as sister to Misodendraceae with strong support. Genera and species: This is a family of three genera and 58 species: Arjona (8), Quinchamalium (25) and Schoepfia (25). Uses: Arjona makes edible tubers.

Schoepfia schreberi, Yucatán, Mexico [314]

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Etymology: Schoepfia is named for German physician, zoologist and botanist Johann David Schöpf (1752–1800). He travelled in New England, Florida and the Bahamas during the late 1770s where he collected many specimens, which he described in his Travels in the Confederation (1783).

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315. LORANTHACEAE Showy-mistletoe family

Species of Loranthaceae are evergreen branch parasites (mistletoes) and parasitic vines, and a few are root parasitic trees and shrubs (Atkinsonia, Gaiadendron, Nuytsia). Roots grow along the host epidermis and make intermittent haustoria that are large and may cause host-tissue proliferation. Leaves are opposite or in whorls of three (ternate), simple and well developed or reduced to scales. Margins are entire, and venation is parallel-pinnate. Inflorescences are axillary or terminal racemes, spikes, umbels or heads, all derived from compound dichasia. Flowers are bisexual or rarely unisexual and usually actinomorphic, but some are slightly zygomorphic and may open explosively. Sepals are usually reduced to an entire, toothed or lobed rim or shallow cup around the tip of the ovary (calyculus). The five or six (rarely three to nine) petals are free or slightly fused at the base forming a long tube that is lobed equally or cleft unequally. A nectary is often present. As many stamens as petals are placed opposite them, often with the filament fused to the petal base. In some African taxa, filaments are coiled and spring outwards when touched. Anthers are basifixed and open by longitudinal slits. The inferior ovary is composed of three or four carpels, fused to form a single, or rarely four (Lysiana), locules. Fruits are berries or drupes with milky sap and a single seed. The calyculus is sometimes enlarged around the top of the fruit. Nuytsia has a winged samarra. The two cotyledons often become fused. Distribution: Loranthaceae are widespread, in the Americas from Mexico to central Chile

A system of natural plant families — Adolf Engler (1844–1930) and Karl Prantl (1820–1888) Since Linnaeus, there has been only one nearly complete detailed work attempting to describe and classify all plants of the world, the 23-volume Die Natürlichen Pflanzenfamilien (The Natural Plant Families, 1887–1915) written and edited by German botanists Adolf Engler and Karl Prantl. Placing algae, mosses, ferns, gymnosperms and angiosperms in a natural rather than an artificial classification, it was a monumental work involving numerous collaborators and experts. Even today, this system is still used, although it is now superseded by the APG system used in this volume. There is no complete revised classification including all plant lineages. Adolf Engler was a pioneer in interdisciplinary sciences, emphasising the importance of, for instance, geology in the study of biodiversity and ecology. Although Engler worked more on phanerogams, Prantl was a specialist on cryptogamic plants, which proved to be an excellent combination of talents. The genus Englerina (Loranthaceae) was named for Adolf Engler because he described the type species, Loranthus holstii, in an article describing plants from Africa.

Photograph of Heinrich Gustav Adolf Engler in 1884. Unknown photographer (public domain)

and Argentina, in eastern Europe, Anatolia, Sub-Saharan Africa, Yemen, Madagascar, tropical and subtropical Asia, north to Japan, and throughout the Pacific south to New Zealand. They are more diverse in the Southern Hemisphere, especially Australia, although flower morphology is also diverse in South America. Phylogeny and evolution: Loranthaceae were generally associated with Viscaceae (now tribe Visceae in Santalaceae) because they share the mistletoe habit. There are, however, many differences, and molecular studies place Loranthaceae closer to Misodendraceae and Schoepfiaceae, whereas Santalaceae are closer to Opiliaceae. The root parasite Nuytsia is sister to the rest of

Photograph of Karl Anton Eugen Prantl c. 1890. Unknown photographer (public domain)

the family, and other root parasitic genera (Atkinsonia, Gaiadendron) may also belong there, but this is not yet clear. In total, five clades are found in phylogenetic studies, which were formally recognised at the tribal level. The family started diversifying c. 28–40 million years ago. Gaiadendron is a root parasite that sometimes infects epiphytes in the canopy but does not infect the supporting tree. Like branch parasites of Santalaceae, fruits of Loranthaceae are sticky, and when birds eat the fruits, this sticky layer adheres to their bills and is then wiped off on a branch, thus dispersing the seed and infecting another branch. Australian mistletoes resemble their hosts, especially Amyema infecting Eucalyptus, which often have similar leaves.

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Amyema pendula, Burrinjuck, New South Wales, Australia [315]

Lysiana murrayi in fruit, near Coober Pedy, South Australia [315]

Lysiana murrayi, near Coober Pedy, South Australia [315]

Diplatia grandibracteata, west of Oodnadatta, South Australia [315]

Genera and species: Loranthaceae include 74 genera with c. 1,050 species: Actinanthella (2), Aetanthus (11), Agelanthus (62), Alepis (1), Amyema (107), Amylotheca (4), Atkinsonia (1), Bakerella (16), Baratranthus (3), Benthamina (1), Berhautia (1), Cecarria (1), Cladocolea (26), Cyne (6), Dactyliophora (3), Decaisnina (25), Dendropemon (31), Dendrophthoe (30–60), Desmaria (1), Diplatia (3), Distrianthes (1), Elytranthe (10), Emelianthe (2), Englerina (26), Erianthemum (16), Gaiadendron (1), Globimetula (13), Helicanthes (1), Helixanthera (55), Ileostylus (1), Lampas (1), Lepeostegeres (9), Lepidaria (14), Ligaria (2), Loranthus (9), Loxanthera (1), Lysiana 430

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(8), Macrosolen (25–50), Moquiniella (1), Muellerina (4), Notanthera (1), Nuytsia (1), Oedina (4), Oliverella (3), Oncella (4), Oncocalyx (12), Oryctanthus (15), Oryctina (10), Panamanthus (1), Papuanthes (1), Passovia (21), Pedistylis (1), Peraxilla (2), Peristethium (15), Phragmanthera (35), Phthirusa (c. 7), Plicosepalus (12), Psittacanthus (c. 119), Scurrula (43), Septulina (2), Socratina (2), Sogerianthe (4), Spragueanella (2), Struthanthus (c. 50), Tapinanthus (30), Taxillus (60–85), Thaumasianthes (1), Tolypanthus (7), Trilepidea (1), Tripodanthus (3), Tristerix (11), Trithecanthera (5), Tupeia (1) and Vanwykia (1).

Uses: Nuytsia floribunda is occasionally cultivated as an ornamental in Australia, but its haustoria can cut underground electrical and phone wires and even cause leaks in drainage pipes. Its common name is the Western Australian Christmas tree as it flowers around Christmas, during the dry Australian summer when other plants are dormant. Etymology: Loranthus is composed of the Latin lorum, a strap, and Greek άνθος (anthos), a flower.

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EUDICOTS

CARYOPHYLLALES Families 316 to 353 make up the large and diverse order Caryophyllales. The family circumscriptions in this order have long been difficult, especially in the MolluginaceaePortulacaceae assemblage, which was found to be widely polyphyletic, resulting in the recognition of a number of small, poorly defined families and leaving larger well-defined families such as Cactaceae, Didiereaceae and Nyctaginaceae untouched. More broadly defined families could be circumscribed, but this is not supported by the botanical community and thus the smaller entities are recognised in APG IV, which is here followed.

316. FRANKENIACEAE Sea-heath family

These are salt-loving shrubs and perennial and annual herbs, often cushion plants, with opposite, simple leaves that bear salt glands sunken in the lamina. Leaves lack stipules, but leaf pairs are often united at the base to form a sheath, or are petiolate. Leaf blades are flattened or nearly round, with an entire margin that is strongly recurved with obscure venation, Inflorescences are axillary dichasia that form leafy cymes, or the flowers are solitary in the leaf margins. The usually bisexual f lowers are actinomorphic and subtended by two prophylls and a pair of leaflike bracts. The four to seven (usually five) sepals are fused into a persistent, ribbed tube Frankenia serpyllifolia, near Oodnadatta, South Australia [316]

with short lobes. The five (sometimes four to seven) petals are free, basally clawed and usually with a scale-like appendage or ligule on the upper side of the claw. The three to six (rarely up to 25) stamens are usually in two whorls of three with filaments that are free or shortly basally fused. Anthers are versatile and open lengthwise to the outside (extrorse). Rarely the inner stamen whorl is staminodial (sterile). The superior ovary is composed of usually three (sometimes one to four) fused carpels forming a single locule. A slender apical style is topped with as many stigmatic branches as carpels (usually three). The fruit is a loculicidal or apicidal capsule surrounded by the persistent calyx.

Phylogeny and evolution: Frankeniaceae are closely related to Tamaricaceae, with which they share many characters such as the small leaves with salt-excreting glands, petal appendages, secondary chemistry and pollen morphology. Because of their parietal placentation, they were previously placed in Violales, but molecular studies placed these two families together as sister to Polygonaceae+Plumbaginaceae in Caryophyllales. The lineage diversified 30–43 million years ago.

Distribution: This family occurs throughout the warmer and drier regions of the world, especially coastal habitats and other sites with saline soils. It is found in western North America (southwestern USA and northern Mexico), dry western and southern South America, Atlantic islands, coastal western and Mediterranean Europe, North Africa, South Africa, western and Central Asia and Australia (the last where the greatest diversity occurs).

Uses: Frankenia laevis (a native of Europe including Britain) is occasionally grown as a rock garden ornamental.

Frankenia muscosa, Dalhousie Springs, South Australia [316]

Genera and species: The sole genus in this family is Frankenia with c. 90 species, probably more.

Etymology: Frankenia is named to commemorate Johann Francke (Johannus Frankenius, (1590–1661), Professor of Medicine at Uppsala University, who was the first to describe the plants of Sweden.

Frankenia laevis, Sicily, Italy [316]

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Reaumuria kuznetzovii, Rustavi, Republic of Georgia (CD) [317]

Tamarix ramosissima, Royal Botanic Gardens, Kew, UK [317]

Myricaria germanica, Ruissalo Botanical Garden, Turku, Finland [317]

317. TAMARICACEAE

sessile on the ovary. Fruits are loculicidal, abaxially dehiscent, conical capsules with numerous seeds that are hairy or with an awn at the tip.

Uses: Tamarix (tamarisk or salt cedar) is often planted in dry or coastal habitats because of its erosion-control properties, due to strong roots and tolerance to salt, flooding and drought. Galls are sometimes formed, which are a good source of tannins, and twigs are flexible and can be used for weaving and basketry. Myricaria germanica is sometimes used to produce a black dye. Tamarix is sometimes planted as an ornamental.

Salt-cedar family

This is a family of shrubs and trees with photosynthetic twigs. Their leaves are alternate, small and scale-like and lack stipules. They are usually sessile and have salt-excreting glands. Inf lorescences are racemes or panicles, often aggregated in bunches, rarely the flowers solitary in leaf axils. Flowers are usually bisexual and actinomorphic. The four or five sepals are free or rarely fused at the base and persistent in fruit. The four or five petals are free, sometimes (Reaumuria) with a small scale on the petal. The four, five or numerous stamens are usually equal to or several times the number of stamens. Filaments are usually free, rarely united into fascicles, usually placed on the nectar disk, sometimes merged with the petal base. Anthers are dorsifixed and open by lengthwise slits. A thick nectar disk surrounds the ovary, which consists of two to five carpels fused into a single or several locules. It is topped by two to five free styles, rarely fused into a single style, or the stigmas 432

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Distribution: Tamaricaceae are found in steppe and desert regions of the Old World, from Macaronesia across southern and eastern Europe and northern Africa, through Arabia, Anatolia, southern and Central Asia to southwestern China and the Russian Far East. The centre of diversity lies in Central Asia. They are also found in East Africa, southwestern Africa and as an isolated population in western Scandinavia. They usually grow in dry and/or salty habitats, often river beds, and have been introduced by humans to many areas outside their native range (North America, South America, Western Europe, Australia etc.). Phylogeny and evolution: Reaumuria, which has petal scales, is sister to the rest of the family, which lack these scales. Hololachna is included in Reaumuria, from which it differs little. Myrtama has been placed alternatively in Myricaria and Tamarix, but it is morphologically distinct and is thus treated as a separate genus. Pliocene and Pleistocene fossils of Tamarix are known from northern and eastern Africa and Central Asia. Genera and species: Tamaricaceae includes four genera and c. 78 species: Myricaria (10), Myrtama (1), Reaumuria (12), Tamarix and (c. 55).

Etymology: Tamarix is derived from the classical Latin tamariscus, a tamarisk bush.

318. PLUMBAGINACEAE Thrift family

This is a family of perennial herbs, shrubs and vines. Leaves are simple and spirally arranged, but sometimes with a basal rosette, without stipules. Leaf blades have entire margins, sometimes with basal auricles, pinnate or reduced venation and usually chalk-glands that exude water and calcium salts. Inflorescences are axillary, bracteate racemes, spikes, heads or thyrses. Bracts are

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usually sheathing and papery, and flowers are subtended by two (rarely one) bracteoles. Flowers are bisexual and actinomorphic. The five sepals are fused to compose a five- or ten-ribbed tube that is often petal-like and persistent. The five petals are nearly free to completely fused. A nectar disk is usually present, often as five separate glands between the stamens. The five stamens are opposite the petals, and filaments are free or fused with the petal tube. The basifixed anthers open by lengthwise slits. The superior ovary is composed of five fused carpels forming a single locule and topped with five free styles with dry, papillate stigmas. Styles of different lengths (heterostyly) are common in this family. Fruits are dry, one-seeded achenes or a circumscissile or apically opening capsule, all subtended by persistent sepals and sometimes petals. Distribution: This is a family with a worldwide distribution, but mostly coastal in North America and Europe. They occur throughout the tropics in wet and dry, fresh

and saline habitats and in mangroves in Bengal and northern Australia. They are most diverse in the Mediterranean and IranoTuranian regions. Phylogeny and evolution: Plumbaginaceae are closely related to Polygonaceae with which they share secondary chemistry, broadened leaf bases and an accrescent calyx. It is estimated that Plumbaginaceae evolved 111–118 million years ago, but radiation of the crown clade was more recent, c. 37–52 million years ago. There are two major clades in Plumbaginaceae, with the mangrove plant Aegialitis sister to both. The classification below is currently the one accepted, with Aegialitis usually segregated in its own subfamily. There are many genera with few species segregated from Limonium, the circumscription of which may need to be revisited. Taxonomy of Limonium is complex due to the widespread occurrence of polyploidy and apomixis. Genera and species: Plumbaginaceae include c. 30 genera and c. 725 species,

Plumbago scandens, Galápagos Islands [318]

Armeria maritima, The Burren, Ireland [318]

in three subfamilies: Aegialitidoideae – Aegialitis (2); Staticoideae – Acantholimon (c. 165), Afrolimon (7), Armeria (c. 100), Bakerolimon (2), Bamiania (1), Bukiniczia (1), Cephalorrhizum (2), Chaetolimon (3), Ceratolimon (4), Dictyolimon (4), Eremolimon (7), Ghaznianthus (1), Gladiolimon (1), Goniolimon (c. 20), Ikonnikovia (1), Limoniastrum (9), Limoniopsis (2), Limonium (c. 350), Muellerolimon (1), Myriolimon (2), Neogontscharovia (3), Popoviolimon (1), Psylliostachys (10), Saharanthus (1) and Vassilczenkoa (1); Plumbaginoideae – Ceratostigma (8), Dyerophytum (3), Plumbagella (1) and Plumbago (c. 24). Uses: Some species are grown as ornamentals, especially of Armeria (thrift), Ceratostigma, Limonium (sea lavender) and Plumbago (leadwort). Statice (Limonium sinuatum and other species) comes in many colour forms and is used as a cut flower, usually dried. Etymology: Plumbago is derived from Latin plumbum, lead, and the suffix -ago, to like.

Limonium sinuatum, Sicily, Italy [318]

Acantholimon venustum, Royal Horticultural Society Garden, Wisley, UK [318]

Ceratostigma plumbaginoides, Royal Botanic Gardens, Kew, UK [318]

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Symmeria paniculata, Samiria, Coccoloba unifera, Guadeloupe [319] Loreto, Peru (CD) [319]

319. POLYGONACEAE Knotweed family

Polygonaceae are a family of annual and perennial herbs, shrubs, trees and vines. Their stems are often swollen at the nodes and frequently hollow, sometimes prickly. Leaves are simple, alternate, rarely opposite or whorled, petiolate or blades nearly sessile; stipules are usually united to a usually papery tubular sheath around the stem (ochrea) that is often fringed or toothed, rarely reduced or absent. Blades usually have entire margins with pinnate venation. Inflorescences are terminal or axillary spikes, racemes, panicles or heads, portions of which are subtended by bracts, and each flower is subtended by a tubular sheath (ochreola) consisting of fused bracteoles. Flowers are small, actinomorphic and usually bisexual (rarely unisexual and then the plants dioecious). The two to six 434

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Chorizanthe fimbriata, Santa Barbara, California, USA [319]

tepals are in one or two whorls that are basally fused into a tube (sometimes short), persistent and often enlarged in fruit, producing wings, tubercules or spines. The usually six to nine stamens (rarely more, as few as two) have filaments that are free or fused at the base, often merged with the tepal tube forming a ring. Anthers are versatile and open on the inside (introrse) with lengthwise slits. The superior ovary is composed of usually three (sometimes two or four) fused carpels forming a single locule. The ovary is topped with one to three, rarely four, free or basally fused styles, and variously ornate stigmas. Fruits are usually trigonal or flat nutlets that are often subtended by persistent tepals. Distribution: Polygonaceae have a near-global distribution, but they are predominantly north temperate with only a few representatives in the tropics (absent from the Asian tropics) and the Southern Hemisphere. They are remarkably diverse in western North America (California). Phylogeny and evolution: The family is divided into three subfamilies, the New World (especially Neotropical) Eriogonoideae and the nearly cosmopolitan Polygonoideae, with tropical African Afrobrunnichia sister to both

Eriogonum kennedyi var. alpigenum, Royal Horticultural Society Garden, Wisley, UK [319]

or just Eriogonoideae, and South American and West African Symmerioideae sister to all. Even though long-distance dispersal is evident in all clades, the family seems to have an African-Gondwanan origin, and some clades are old enough to implicate continental drift as the cause. The diverse and possibly paraphyletic Eriogonum probably represents a recent radiation. Diversification of Rheum occurred contemporaneously with uplifting of the Tibetan plateau, 9.2–12.0 million years ago. Woodiness seems to have evolved several times in Polygonaceae from an herbaceous habit. A number of genera have recently been reorganised, but it seems best to apply a broad concept to Persicaria, including Aconogonon, Bistorta, Koenigia and Knorringia, although not all necessary combinations for these have thus far been made in Persicaria. A woody species, Fagopyrum tibeticum, has potential as a perennial source of buckwheat. Genera and species: This family has c. 50 genera, and includes some 1,200 species in three subfamilies: Symmerioideae – Symmeria paniculata; Eriogonoideae (28) – Acanthoscyphus (1), Afrobrunnichia (2), Aristocapsa (1), Centrostegia (1), Chorizanthe (50), Coccoloba (c. 120), Dedeckera (1), Dodecahema (1), Eriogonum (c. 250), Gilmania (1), Goodmania

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(1), Hollisteria (1), Johanneshowellia (2), Lastarriaea (3), Mucronea (2), Nemacaulis (1), Oxytheca (3), Pterostegia (1), Sidotheca (3), Stenogonum (2) and Systenotheca (1); Polygonoideae – Antigonon (9), Atraphaxis (25), Brunnichia (1), Calligonum (80), Duma (3), Emex (2), Fagopyrum (17), Fallopia (12), Gymnopodium (3), Harfordia (1), Homalocladium (1), Leptogonum (1), Magoniella (1), Muehlenbeckia (19), Neomillspaughia (2), Oxygonum (22), Oxyria (1), Persicaria (c. 235), Podopterus (3), Polygonella (11), Polygonum (c. 15), Pteropyrum (6), Rheum (c. 60), Rumex (c. 200), Ruprechtia (c. 14), Salta (1) and Triplaris (c. 17). Uses: Possibly originating in Yunnan, China, buckwheat (Fagopyrum esculentum) has been grown for food in Eurasia since ancient times and has been found in the stomachs of naturally mummified bodies in peat bogs (the

so-called bog men) in northwestern Europe. The flour is popular for noodles in Japan and most commonly used for pancakes in Europe (including Breton crepes and Dutch pannenkoeken) and North America. Pillows filled with the seeds can be heated (in an oven or microwave) and will retain the heat for a long time, being used to treat muscle pains. Tatar buckwheat (F. tataricum), probably also from China, is used locally for similar purposes. Black bindweed seeds (Fallopia convolvulus) were eaten in prehistoric Europe. Most Coccoloba species have edible, grapelike fruits that are actually a fleshy calyx surrounding the nutlet. The best known is seaside grape (Coccoloba uvifera), which is common on Neotropical beaches and planted elsewhere in the tropics. The fleshy calyx is made into jellies and wine and used to produce a red dye. Similarly those of climbing lignum (Muehlenbeckia adpressa) are used for pies

in Australia. Tubers of coral vine (Antigonon leptopus) are edible and have a nut-like flavour. This Mexican vine is commonly cultivated and frequently naturalised throughout the tropics. Petioles (leaf stalks) of rhubarb (mostly Rheum rhabarbarum and R. rhaponticum and their cultivars) are commonly picked in spring and early summer. They are sweet and sour with lots of malic acid and can be used for making jam, pies and other baked goods and wine. Similarly, Babylonian rhubarb (R. ribes) from southwestern Asia has stalks that can be eaten raw and taste like currants or are used in local Turkish or Iranian cuisine. Sorrel (Rumex acetosa) has edible leaves, and it is often cultivated in Eurasian allotments for use in salads, sauces and soups. Tanner’s dock (R. hymenosepalus) is used similarly in southwestern North America, and its leaves can be used as rhubarb. Patience dock (R. patientia), monk’s rhubarb (R. pseudoalpinus)

Antigonon pteropus, cultivated in Mombasa, Kenya [319]

Oxyria digyna, Lake District, England, UK [319]

Fallopia japonica, French Pyrenees [319]

Persicaria affinis, Royal Botanic Gardens, Kew, UK [319]

Polygonum aviculare, Canbury Gardens, Kingston upon Thames, Surrey, UK [319]

Rheum alexandreae, Yunnan, China [319]

Rheum rhabarbarum, showing the ochrea on inflorescence, Hengelo, the Netherlands [319]

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CARYOPHYLLALES and French sorrel (R. scutatus) are used like R. acetosa. Rumex species also produce tubers that are rich in tannins, formerly used to tan leather. Widespread mountain sorrel (Oxyria digyna) is locally grown as a rock garden ornamental and has edible leaves. Flowers of Calligonum polygonoides are used in baked goods and Indian cuisine. Vietnamese coriander (Persicaria odorata) is a potherb, commonly used in Southeast Asian cuisine, especially in salads and soups including laksa. An essential oil from the plant (kesom oil) is being investigated as a natural food flavouring and additive. In Vietnam it has a reputation as an antiaphrodisiac. It has a strong corianderlike smell with a lemony note and is not as pungent as water pepper (P. hydropiper), which was formerly used in Eurasia as a peppery spice. Water pepper should be used in moderation. Special cultivars selected for a milder taste are grown in Japan as a vegetable, and a red form is used as sashimi garnish. Seeds of water pepper are sometimes used to adulterate wasabi. Young shoots of soft knotweed (P. mollis) are eaten as a vegetable in Sikkim. An indigo dye is made from P. tinctoria in East Asia. The rhizomes of bistort (P. bistorta) are used in northern England in Easter-ledge pudding, probably an association with with its legendary properties of aiding conception. Knotgrass (Polygonum aviculare) is now known as a bad weed, but it was formerly brewed into a tea, effective in the treatment of asthma. Triplaris weigeltiana has hollow stems, but it makes valuable timber for Dionaea muscipula, Royal Botanic Gardens, Kew, UK [320]

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indoor carpentry. Shrubs of Central American Gymnopodium yield good charcoal. A number of species are cultivated as ornamentals, especially in Antigonon, Atraphaxis, Coccoloba, Eriogonum, Fallopia, Homalocladium, Muehlenbeckia, Persicaria and Rheum. Japanese knotweed (Fallopia japonica) is a difficult-to-eradicate invasive in many parts of the temperate zones, forming vast stands crowding out native vegetation, and removal of these has high costs. A female clone originally escaped from Von Siebold’s plant nursery in Leiden, the Netherlands. Male plants have since been introduced to Europe and North America causing widespread seedset and further naturalisation. Plants are still offered as an ornamental, and it is sometimes cultivated for its young shoots that can be eaten like rhubarb, its mature stems yielding a reasonable fibre for papermaking (good for making seed/cutting pots, hangingbasket liners and egg cartons). The long rhizomes can be collected to produce a yellow dye, so maybe eradication efforts can be put to some use. In the UK it is now illegal to allow Japanese knotweed to spread onto neighbouring properties and to dispose of plant material that still has the potential to regrow. This ban includes the movement of contaminated soil. Etymology: Polygonum is composed of the Greek words πολύς (polys), many, and γόνυ (gony), a knee, in reference to the swollen nodes of the stems. Dionaea muscipula, Green Swamp, North Carolina, USA [320]

320. DROSERACEAE Sundew family

These annual and perennial, carnivorous herbs usually grow terrestrially, but sometimes they occur as free-floating submerged aquatics (Aldrovanda). The stem has adventitious roots (much reduced leaves that function as rhizoids underground) and sometimes bears rhizomes, corms or tubers. Above-ground stems are reduced or elongate and then self-supporting or climbing. Leaves are alternate or rarely whorled and in a basal rosette or placed along the stem. They are usually petiolate and have stipules or not, and the blades are circinnate (like ferns) or infolded when they emerge. Leaf blades are motile, have sticky, glandular, slime-tipped tentacles (hairs) that trap and digest small animals (the fly-paper trap of Drosera), or leaves are modified into snaptraps with sensitive hairs inside that trigger the closing of the blade to trap small animals (Aldrovanda and Dionaea). Inflorescences are axillary cincinnae organised in thyrsoid panicles or (rarely) racemes, usually held well above the traps, or flowers are solitary Aldrovanda vesiculosa (Siberian form), Royal Botanic Gardens, Kew, UK [320]

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Drosera rotundifolia, with prey, Ireland [320]

Drosera macrantha, near Geraldton, Western Australia [320]

Drosera regia, emerging leaf (circinnate), Royal Botanic Gardens, Kew, UK [320]

Drosera glanduligera, Mt Benia, Western Australia [320]

(Aldrovanda). Flowers are actinomorphic and bisexual. The usually five, sometimes four, six or eight, sepals are persistent in fruit and more or less fused at the base. Petals are free and usually five, but normally as many as sepals or more. The four or five (or up to 20) stamens are usually free (fused at the base in Dionaea) and alternate with the petals. The usually basifixed anthers open on the outside with two longitudinal slits. The superior ovary is usually composed of three (rarely two to five) fused carpels, forming a single (rarely up to three) locules. The ovary is topped with free and often deeply branched styles (Aldrovanda, Drosera) or the styles fused into a simple common style (Dionaea). Fruit a loculicidal or (rarely) indehiscent capsule.

a broad distribution, extending from Central Europe south to southeastern Africa and Australia, but its distribution is now patchy due to local extinction resulting from water pollution. Dionaea is endemic to a restricted swampy area in coastal southeastern North America (North and South Carolina), but it has been introduced and has naturalised in California, New Jersey, Florida and Jamaica.

the Eocene of Europe. Drosera pollen has been found in Miocene formations of Europe and New Zealand. Diversification in southwestern Australia is probably linked to the development of the winter-rain Mediterranean style climate in that region. The Australian pygmy sundew clade includes a species from Guyana, which is a striking case of longdistance dispersal.

Phylogeny and evolution: Originally Droseraceae were associated with other car nivorous plant families, notably Nepenthaceae, Ancistrocladaceae and Dioncophyllaceae. The last two are indeed their closest relatives on the basis of molecular studies. They are not related to other families with flypaper traps such as Byblidaceae (Lamiales) or Roridulaceae (Ericales), with which, apart from the trapping mechanism, they share almost no characters. Dionaea and Aldrovanda share snap-traps and are sister genera. Diversification in Drosera may have started c. 42 million years ago, although older ages have been suggested. Aldrovanda fossils (pollen and macrofossils) are known from

Genera and species: Droseraceae are a family of three genera and 130 species: Aldrovanda (1), Dionaea (1) and Drosera (128).

Distribution: A cosmopolitan family, Droseraceae are usually confined to nutrient poor, sandy or peaty environments that are often seasonally dry. They are widespread across the cool north temperate zone, locally extending into the subtropics, and throughout the tropics/subtropics with the greatest diversity in Western Australia. Aldrovanda has

Uses: Several species are grown as curiosity ornamentals, especially Venus’ f ly trap (Dionaea muscipula), which, with its rapidly closing traps, captivates the imagination and fascinates children. The plant is probably one of the origins of stories about carnivorous plants actively grabbing its victim. The popularity of Venus’ fly traps in cultivation and the limited area of distribution in the Carolina swamps has resulted in a high level of threat to the native populations. Currently wild plants are protected, wild collections Plants of the World

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CARYOPHYLLALES are illegal and plants in the trade have been propagated by tissue culture. Nevertheless, the native sites are under threat from urban development, and the species is therefore classified as ‘Vulnerable’ by IUCN. Several species of sundew (Drosera) and waterwheel (Aldrovanda vesiculosa) are grown in specialist collections. Carnivory: Droseraceae demonstrate remarkable diversity in morphology, specifically in their mode of carnivory, with two major forms of traps: flypaper traps in Drosera and snap traps in Aldrovanda and Dionaea. Both trap types are active, although those of Drosera are slower in their response. The sticky tentacles of Drosera catch a small animal (often insects or other arthropods) in the mucilage drops formed on top of its glandular hairs, and the more the insect struggles, the more the hairs are stimulated (often beginning in as little as ten seconds). The stimulation of one tentacle causes movement in the surrounding tentacles. Organic substances cause more movement than inorganic substances, and movement occurs after the hairs have been irritated at least three times. The leaves are capable of true digestion, and the glands can absorb digested matter. The digestive enzymes are similar to the proteolytic enzymes of animals. Although the trapping mechanism of all Drosera species is basically the same, the vegetative morphology of the plants varies greatly between species. Many are rosette forming, but extreme variation is found in southwestern Australia, e.g. D. gigantea, which is about a metre tall and produces annual growths with many flypaper traps on a highly branched, erect, self-supporting stem. Other species in the same region of Australia have elongate scrambling stems that twine around supports using their trap leaves, making additional trap leaves on short axillary shoots to catch insects. Recently it has been found that the tentacles of the Australian pimpernel sundew (Drosera glanduligera) have a hinge and upon contact catapult their prey into the centre of the sticky leaf. The Venus’ fly trap (Dionaea muscipula) is one of the best-known carnivorous plants because of its quick movement. The plant has a

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leaf with two opposite terminal pinnae that are cup-shaped and hinged at the base. They have a series of teeth along the margin, of which the opposite sides interlock when closed. Inside are trigger hairs that, when stimulated, generate a current by the movement of calcium ions. The change in acidity makes the midrib cells change shape as a result of osmosis, and the trap closes, catching the prey. The trap closes tightly and releases digestive enzymes, and the dissolved proteins are absorbed by the leaf tissue. After several days the prey is reduced to its chitin skeleton, and the trap reopens. Despite their aquatic habit and general morphological dissimilarity, the traps of the waterwheel (Aldrovanda vesiculosa) bear clear similarity to those of Dionaea, albeit somewhat miniaturised. They function in a similar fashion, trapping small aquatic animals in oligotrophic still water. Etymology: Drosera is derived from the Greek δρόσος (drosos), dew, referring to the glistening excretion of the leaf glands.

321. NEPENTHACEAE Asian-pitcherplant family

Nepenthaceae are unisexual, carnivorous, terrestrial vines, shrubs and epiphytes. Seedlings form rosettes, and, in the majority of species, plants then form an erect, scrambling or climbing stem. Stems are woody, covered in glandular hairs or glabrous. Leaves are alternate, petiolate or sessile and then the leaf base clasping, without stipules. The midvein is prolonged into a tendrillate tip, and the apex of the tendril usually develops into an urn-shaped to cylindrical pitcher with a lid usually projecting over the pitcher mouth and a spur inserted at the base of the lid. The pitcher has a slippery

zone and digestive glands further inside. Inflorescences are terminal (but appearing leaf-opposed because of continued axillary growth) racemes, thyrses or botryoids, usually sparsely bracteate. Flowers are unisexual and actinomorphic, with a single whorl of (three or) four, free (rarely basally fused) tepals and often with nectar glands within. Male flowers have four to 24 stamens, with the filaments fused into a column and highly reduced pistillodes. Anthers open by longitudinal slits. Female flowers have a whorl of highly reduced staminodes, and a superior, sometimes stalked (gynophore) ovary composed of usually four (rarely three to six) fused carpels each forming a locule. A disc-like stigma tops the ovary, sessile or with a short style. Fruits are four-valved, loculicidally dehiscent capsules with many hair-like seeds. Distribution: This family is predominantly distributed in tropical East and Southeast Asia, extending west to Madagascar, the Seychelles and Sri Lanka, Assam and in tropical Southeast Asia to the Caroline Islands, New Caledonia and northern Australia. It has its greatest diversity in Borneo, Sumatra, Sulawesi and the Philippines (particularly Palawan). Nepenthes species generally grow in thin mats of leached organic matter that can overlay alkaline, neutral or acid substrates. Phylogeny and evolution: The species from the Seychelles (Nepenthes pervillei) and Madagascar (N. madagascariensis) were found to be sister to the rest of the genus, suggesting that the origin may have been west of Malesia, where the greatest diversity currently occurs. The family may have diverged c. 55–67 million years ago. Genera and species: There is only one genus in Nepenthaceae: Nepenthes (c. 150 species). Uses: Nepenthes is frequently grown as an ornamental, and complex man-made hybrids are often offered. Unusual species can be found in specialist collections. Carnivory and other uses of the trap: The traps of Nepenthes are pitchers formed from

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Nepenthes ampullaria, Malacca, Malaysia [321] Nepenthes spectabilis, male flowers, Royal Botanic Gardens, Kew, UK [321]

Nepenthes pervillei, Mahé, Seychelles [321]

Nepenthes lowii, Borneo (WA) [321]

Nepenthes vieillardii, female flowers, New Caledonia [321]

tendrils with a mouth into which prey can fall. The pitchers have nectar glands near the top, which attract insects and other invertebrates, and these in turn may attract predators of these creatures. The rim and upper part of the pitcher are slippery, smooth and waxy, and the pitcher is filled with water, so when an animal loses its grip, it lands in the water and drowns. The pitchers are lined with secretory glands that produce a digestive liquid, and the digestive process is thought to be partly due to the enzymes secreted into the liquid by the pitchers and partly because of microorganisms living in the liquid. In newly opened pitchers, the liquid can be highly acidic (pH 2.5), which also may help digestion. Nepenthes species

are thus active carnivores because they trap prey, digest them and absorb nutrients. Prey items are predominantly invertebrates, although occasional cases have been reported of the remains of amphibians, lizards and even small mammals or birds in pitchers of the largest species (N. rafflesiana, N. rajah). Specialisation on particular types of prey has also been reported, such as termites (N. albomarginata), but some species appear to have forgone carnivory altogether. Several species are detritivores, gaining nutrients from fallen leaf litter (N. ampullaria) and reports of a shrew (Tupaia montana) defecating in a pitcher while eating nectar from the lid of N. lowii are remarkable cases of symbiosis. Several other species (e.g.

Nepenthes bicalcarata, Royal Botanic Gardens, Kew, UK [321]

N. ephippiata, N. macrophylla, N. rajah) are also likely to have similar symbioses. Pitchers of Nepenthes can provide a home for various microorganisms, invertebrates and even vertebrates, which may take advantage of the trapping of other animals by the plants. Etymology: Nepenthes is combined of the Greek νε (ne), not, and πενθές ( penthes), mourning: “nepenthe” thus being a plant that would drive away sadness, an antidepressant or a drug of forgetfulness, a name already used in Homer’s Odyssey. Pliny used the name for a plant that, mingled with wine, had an exhilarating effect, but it was later applied by Linnaeus to these pitcher plants.

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322. DROSOPHYLLACEAE Dew-pine family

EUDICOTS

capsules that open along the locules in the upper half, containing numerous round, black seeds. Distribution: This species is found in the southwestern Iberian peninsula (Atlantic coast of southern Portugal, the southernmost tip of Spain and Gibraltar) and northernmost Morocco. It grows on dry, basic, nutrientdeficient soils.

These are carnivorous herbs with a woody rootstock and a tap root. Stems and leaves are covered with rows of stalked slimeexcreting tentacles and irregularly distributed sessile digestive glands. Leaves are alternate, arranged spirally in a crowded rosette at the stem ends without stipules or petioles. The leaves are reversely circinnate (curling downward) in bud. Inf lorescences are terminal, few-flowered thyrsoid panicles. The bisexual flowers are actinomorphic and have five basally fused sepals that are glandular outside. The five free petals are yellow. The ten stamens are arranged in two whorls of five, with free filaments and dorsifixed anthers that open outwardly by longitudinal slits. The superior ovary is composed of five fused carpels topped with five, free, reflexed styles, each with a capitate stigma. Fruits are

Drosophyllum lusitanicum, emerging leaf (reverse circinnate), Royal Botanic Gardens, Kew, UK [322]

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Phylogeny and evolution: Drosophyllum lusitanicum was originally described as a species of Drosera, and it does have similarities with that genus, except for its reversely circinnate leaves that do not move when capturing prey. Molecular phylogenetic studies have demonstrated that Drosophyllum is more closely related to Dioncophyllaceae (with which it shares the trapping mechanism and gland types of Triphyophyllum) and the non-carnivorous Ancistrocladaceae.

rosette of leaves covered with sticky, mucilage-secreting, long glands that trap prey (flypaper traps). Once trapped, insects are dissolved by enzymes released by short glands on the leaves, and the released nutrients are absorbed by the plant. It is an active carnivore because it has all the cellular mechanisms to trap, digest and absorb. Etymology: Drosophyllum is composed of the Greek words δρόσος (drosos), dew, and φύλλων ( fyllon), a leaf, in reference to the glistening excretions of the leaf glands, and their similarity to Drosera (Droseraceae).

323. DIONCOPHYLLACEAE Hookleaf-vine family

Genera and species: This family consists of one genus with the single species: Drosophyllum lusitanicum. Uses: It is occasionally grown as a curiosity in specialist collections. Carnivory: Drosophyllum forms a stalked

Drosophyllum lusitanicum, Royal Botanic Gardens, Kew, UK [322]

This is a family of climbing shrubs and woody lianas with alternate sessile leaves that lack stipules. There are two or three types of leaves on the same plant, all with reverse circinnation

Drosophyllum lusitanicum, Santarem, Portugal (CD) [322]

CARYOPHYLLALES

EUDICOTS

(as in the related Drosophyllaceae): normal leaves with a simple blade, entire margin and closely parallel secondary venation and a simple acute apex; leaves with the apex bearing a pair of recurved hooks; and in Triphyophyllum, leaves that are seasonally formed, filiform and covered in stalked sticky glands and sessile digestive glands. Inf lorescences are axillary cymes with actinomorphic, bisexual flowers. The five sepals are free or basally fused into a short tube. The five free petals and ten or 25–30 stamens are in several whorls, with filaments that are slightly fused or free. Anthers are basifixed and open inwards with longitudinal slits. The superior ovary is composed of two or five fused carpels topped with two or five free styles that are simple or plumed at the tip. Fruits open in two or five locules when immature, exposing the disc-like, broadly winged seeds that hang on a funicle from the open valves. Distribution: This family is restricted to the rainforests of West Africa and the Congo. Phylogeny and evolution: The family was originally placed in the now defunct Flacourtiaceae, but relationships with Nepenthaceae and later with Ancistrocladaceae and Drosophyllum were soon realised. Molecular studies have shown this family to be most closely related to Ancistrocladaceae. This pair of families is sister to Drosophyllaceae and then more distantly related to Droseraceae, Nepenthaceae and other Caryophyllales, as was suggested earlier on the basis of morphology. Genera and species: This family consists of three genera, each with a single species: Dioncophyllum thollonii, Habropetalum dawei and Triphyophyllum peltatum. Uses: Triphyophyllum peltatum has proven to have antimalarial compounds, and it is locally used to treat this and other diseases. It may also have antitumour properties. Carnivory: In this family only Triphyophyllum peltatum is carnivorous, at least part of the

Habropetalum dawei, type specimen collected by G. Meama in Sierra Leone in 1926, ex coll. M. T. Dawe (Herbarium Kew) [323]

Triphyophyllum peltatum, Cote d’Ivoire (SP) [323]

time; the other two genera lack carnivorous leaves. Dioncophyllum appears to lack glands altogether, and the glands in Habropetalum are simple and present only on young stems. This absence of carnivory probably represents losses in these genera, given the relationship to carnivorous Drosophyllaceae, Droseraceae and Nepenthaceae. All species of this family are heterophyllous, but only Triphyophyllum produces a third type of leaves just before the height of the rainy season. These have a filiform midrib that is produced above a short blade and is covered with two types of glands, one long-stalked making a sticky, sweet sap that attracts and traps insects and a second type of sessile glands that produce digestive enzymes, similar to those on the leaves of Drosophyllum. The leaves have reverse circinnation and are short-lived, surviving for only a few weeks. Their prey is mostly insects, predominately beetles, and large numbers are trapped. The flytrap leaves only appear part of the time, probably due to decreasing potassium levels triggering the formation of trap-leaves. Triphyophyllum can therefore rightfully be called a ‘part-time carnivorous plant’.

Etymology: Dioncophyllum is composed of the Greek words δύο (dio), two, όγκος (onkos), hook, and φύλλων ( fyllon), a leaf, in reference to some of the leaf types found in members of this family with two hooks on the tip of the leaves, which they use to climb in the vegetation.

324. ANCISTROCLADACEAE Kardal family

Lianas and scandent shrubs climbing by hooked stem apices make up this family. Their alternate leaves are crowded in terminal rosettes or distributed spirally along stems. Leaves lack petioles, and stipules are usually absent or tiny and soon falling off,

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Ancistrocladus tectorius, China (SK) [324]

but leaves are articulate to the branch and leave a saddle-shaped scar. Leaf blades are simple and often appear petiolate because of the long-attenuate leaf base, and the margin is entire and venation pinnate. Blade surfaces are covered with small pits, each of which contains a hair secreting a waxy substance. Inflorescences are in axillary or terminal dichotomously branched panicles, sometimes composed of racemes or spikes. Bracts are tiny, subtending inflorescence axes and pedicels. Flowers are bisexual and actinomorphic, except for the five unequal sepals that are emergent from the upper or middle part of the ovary. Petals are five, free or basally fused and slightly fleshy. The five, ten or 15 stamens are all equal or different in length and have filaments that are slightly fused at the base. Anthers are basifixed and open by four longitudinal slits. The halfinferior ovary is composed of three carpels that are fused to form a single locule. The three styles are free and articulate to the tip of the ovary, each with a capitate, moonshaped or pinnatifid stigma. Fruits are nuts surrounded by the corky hypanthium and crowned by the unequal sepals and a single hard seed.

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EUDICOTS

Ancistrocladus tectorius in fruit, Vietnam (CD) [324]

Distribution: This family is widespread in the Old World tropics, occurring in tropical Africa, India, Sri Lanka and Southeast Asia.

Rhabdodendron amazonicum, female, Les Nouragues, French Guianas (CD) [325]

325. RHABDODENDRACEAE Clubfruit-tree family

Phylogeny and evolution: Ancistrocladaceae were previously often placed in Violales or Theales, within which their position had been unclear. Molecular and morphological evidence place them close to Dioncophyllaceae, with which they show striking resemblance in habit. Genera and species: This is a family of a single genus, Ancistrocladus, with 21 species. Uses: Several species of Ancistrocladus are used medicinally, and some species have been proven to have antimalarial, antiviral, anti-HIV and fungicidal properties. They are not yet commercially employed by the pharmaceutical industry but are frequently used by local communities. Etymology: Ancistrocladus is composed of Greek αγκίστρων (ankistron), a fish-hook, and κλάδος (klados), a branch, in reference to the hooked branches.

This is a family of shrubs and small trees. Leaves are simple, alternate and without stipules, but with the leaf bases sometimes resembling stipules. Blades have an entire margin, pinnate venation with secondary venation parallel, a gland-dotted surface and peltate hairs with fringed margins on the lower surface. Inflorescences are axillary simple or compound racemes, with scale-like bracts. Flowers are bisexual and actinomorphic with a broad cup-shaped hypanthium. The five sepals are fused around the hypanthium and sometimes form five short lobes or a simple ring. The (four or) five petals are sepal-like, free, and slightly punctate. A nectar disk is absent, and the numerous (c. 30–50) stamens

CARYOPHYLLALES

EUDICOTS

have short flattened filaments. The basifixed anthers open by longitudinal slits but fall off early. The half-inferior, sessile ovary has a single locule, a gynobasic style and an elongate stigma. The fruit is a small drupe borne on a short stalk emerging from the receptacle. Distribution: This family is confined to the Guianas and northeastern Brazil. Phylogeny and evolution: Rhabdodendon has been placed in many families in the past, including Chrysobalanaceae (as Lecostemon), Rutaceae and Phytolaccaceae. Molecular studies placed Rhabdodendron firmly in Caryophyllales, but without a close relationship to any other family. Rhabdodendron may have diverged from other Caryophyllales c. 83–90 million years ago. Genera and species: The sole genus in this family is Rhabdodendron with three species. One (R. gardnerianum) is probably extinct, R. macrophyllum is confined to threatened white-sand forests in Amazonia and the third (R. amazonicum) is a more widespread but uncommon rainforest tree. Etymology: Rhabdodendron is derived from the Greek ράβδος (rabdos), a stick or rod, and δένδρων (dendron), a tree, referring to the stalked fruit.

small bract (male) or solitary in the leaf axils subtended by two large bracts (female). Flowers are actinomorphic and unisexual with five (sometimes four or six) sepals. Petals are absent. Male flowers have eight or 16 free stamens with stout filaments and basifixed anthers that open outwards with lengthwise slits. There are no pistillodes, staminodes or nectaries. Female flowers are pendent on a recurved peduncle and much larger than male flowers. The superior ovary is composed of three (rarely four) fused carpels, each forming a locule. This is topped with three, free, reflexed styles. Fruits are capsules enclosed by the enlarged, fleshy sepals and open loculicidally in three valves, each having a single (rarely more) seed. Distribution: This shrub is only found in the deserts of Baja California, Mojave and Sonora in western North America (Arizona, California and northwestern Mexico). It is found on dry slopes with well-drained soils and occurs only in areas with less than 240 mm annual precipitation.

Monimiaceae. They especially differ from Buxaceae in lacking nectaries and seed morphology, embryology, wood anatomy, chemistry etc. Molecular studies have placed Simmondsiaceae in an expanded Caryophyllales, agreeing with many morphological characters such as the free styles, persistent sepals and, of course, wood anatomy. The family may have diverged c. 70–88 million years ago. Genera and species: The family is composed of a single species, Simmondsia chinensis. Uses: Jojoba is grown commercially to produce a liquid wax, commonly called jojoba oil, that is extracted from the seed. The oil is unique in that it is more similar to human oil than any other produced by the plant kingdom. It is therefore used predominantly in skin care products and other pharmaceuticals. It is also used as a lubricant and biofuel. Its cultivation is also used to curb desertification in some parts of the world.

Phylogeny and evolution: Even though the anomalous secondary growth of the wood should have given a hint of a relationship with Caryophyllales, the family was frequently placed close to Buxaceae, Euphorbiaceae (on the basis of pollen morphology) or even

Etymology: Simmondsia is named in honour of English botanist Thomas William Simmonds (1767–1804). The epithet chinensis refers to China, where the species does not occur naturally, but it was erroneously applied because an abbreviation ‘Calif’ (for California) on the specimen label was misread as ‘China’.

Simmondsia chinensis, male flowers, Palm Desert, California, USA [326]

Simmondsia chinensis, in fruit, Palm Desert, California, USA [326]

326. SIMMONDSIACEAE Jojoba family

Simmondsiaceae are evergreen, unisexual shrubs with nearly sessile, leathery, greyish leaves without stipules. Inflorescences are axillary capitate clusters subtended by a

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CARYOPHYLLALES

Physena madagascariensis, male, Lokobe Forest, Antsiranana, Madagascar (CD) [327]

EUDICOTS

Physena madagascariensis, female, Lokobe Forest, Antsiranana, Madagascar (CD) [327]

327. PHYSENACEAE Balloonfruit family

Physena sessiliflora, male, Orangea Forest Reserve near Antsiranana, Madagascar (CD) [327]

length. Fruits are inflated, stalked, indehiscent capsules with a dry wall and a single seed with a vascularised seedcoat.

328. ASTEROPEIACEAE Manoko family

Distribution: This family is restricted to the eastern parts of Madagascar.

These are shrubs and trees with alternate, simple leaves and short, articulated petioles and no stipules. Blades have pinnate venation and entire margins. Inflorescences are axillary racemes with unisexual actinomorphic flowers. Sepals are five to nine, fused at the base only, persistent in fruit and covered in hairs. Petals and nectaries are absent. Male flowers usually have ten to 14 (rarely eight or up to 25) stamens, which are either free or some filaments basally fused into a short tube. The elongate anthers (making up most of the stamen length) are basifixed and open along the sides with longitudinal slits. Stamens surround a rudimentary pistillode. Female flowers lack staminodes, and the superior ovary is composed of two fused carpels. From a dimple in the top of the ovary, two elongate styles (to 3 mm long) are covered with fibrous appendages along their entire 444

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Phylogeny and evolution: Relationships of Physena have been problematic. Because of its wind-pollination syndrome, it has been associated with Urticaceae in the past, but its parietal placentation made botanists think that it may be related to Capparaceae or the now defunct Flacourtiaceae, although there are no morphological or anatomical characters that would support these placements. The family does share wood-anatomical characters with another Madagascarian endemic family Asteropeiaceae, a relationship that is supported by molecular studies. Their morphological differences in flower structure are because Physena evolved to become pollinated by wind, whereas Asteropeia relies on insects. Genera and species: The single genus Physena consists of two species, P. madagascariensis and P. sessiliflora. Etymology: Physena is derived from the Greek φυσάω ( fysao), inflated, referring to the swollen fruit.

Trees and scrambling shrubs make up this family. They have alternate, simple, shortpetiolate leaves without stipules. Blades are thick and leathery with entire margins and pinnate venation that may be obscured by succulence. Inflorescences are axillary or terminal thyrsoid panicles, without or with two to eight spirally arranged, minute bracts. Flowers are bisexual and actinomorphic. The five sepals are fused at the base, persistent and enlarging in fruit, and there are five free petals. The ten (to 15) stamens are fused at the base into a broad ring, the filaments persistent in fruit and dorsifixed anthers opening inwards by lengthwise slits. The superior ovary is composed of (two or) three carpels that are fused to each form an incomplete locule. The ovary is topped by a short style with a trilobed stigma, or the three stigmas are free and sessile. Fruits are thick-walled,

CARYOPHYLLALES

EUDICOTS

dry capsules that are surrounded by the calyx and indehiscent or rupture irregularly, each with a single seed.

329. MACARTHURIACEAE Macarthuria family

Distribution: The genus is restricted to coastal regions of Australia, with its greatest diversity (six species) in Western Australia.

Distribution: Asteropeiaceae are restricted to Madagascar, where they grow in the eastern, central and northwestern parts of the island in coastal and humid forests up to 900 m and on rocky outcrops. Phylogeny and evolution: Originally published as an anomalous genus, it was later moved to Theaceae, where it remained until the 1990s when wood anatomy and phylogenetic studies showed its affinity to Caryophyllales and close relationship to Physenaceae, another Madagascan endemic. Genera and species: The only genus in this family, Asteropeia, consists of eight species. Uses: Asteropeia rhopaloides produces a locally valued timber, used for construction. Etymology: Asteropeia is derived from the Greek αστραπή (astrapi), lightning. Asteropeia sp., Mandena Reserve, Toliara, Madagascar (CD) [328]

unlobed stigma. Fruits are loculicidal capsules with up to ten seeds.

Phylogeny and evolution: Macarthuria was originally included in Aizoaceae but later moved together with several other genera to Molluginaceae. It was then found, however, that this family was grossly polyphyletic and probably had been recognised on the basis of a few characters plesiomorphic within the larger assemblage. On the basis of its morphology, Macarthuria was placed with Limeum in Limeaceae, but it was found on molecular grounds that these also were not related. Finally, Macarthuria has been found to be sister to all other core Caryophyllales, which diversified c. 60 million years ago.

These are rigid and wiry, rush-like, perennial herbs with green stems that are sometimes woody at the base. Leaves are alternate, petiolate or not, and lack stipules. Blades are simple, linear or progressively reduced to scales on the stem. Leaf blades have entire margins and obscure venation. Inflorescences are many-f lowered, axillary or terminal compound cymes, or flowers are solitary in the leaf axils. Flowers are actinomorphic and bisexual. The five or ten sepals (if the latter, then in two whorls) are fused at the base. Petals are five and free or absent. The eight stamens alternate with the sepals, and their filaments are fused at the base. Anthers are basifixed and open by lengthwise slits. The superior ovary is composed of three carpels that are fused into a single (rarely three) locule. The three styles are fused at the base, each with an

Etymology: Macarthuria is named for Australian horticulturist, botanist and agriculturist Sir William Macarthur (1800–1882), who was important for the promotion of grape cultivation and wine-making in Australia.

Asteropeia rhopaloides, Anjozorobe, Madagascar [328]

Macarthuria australis, Perth, Western Australia [329]

Genera and species: Macarthuriaceae include just one enigmatic genus, Macarthuria, with about ten species.

Asteropeia amblyocarpa, Irodo, Antsiranana, Madagascar (CD) [328]

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330. MICROTEACEAE Jumby-pepper family

EUDICOTS

of three to five fused carpels forming a single locule topped with two to five (nearly) sessile papillose stigmas. The fruit is a muricate or spiny achene. Distribution: This family is restricted to tropical America.

These are annual herbs that may become slightly woody at their base. They have alternate, sessile leaves without stipules, entire leaf margins and simple venation. Inflorescences are axillary spikes or spikelike racemes. Flowers are bisexual and actinomorphic, subtended by a single bract and two bracteoles, or bracteoles absent. The (four or) five sepals are free and linear-elliptic to elliptic, and petals are absent. The (two to) five to eight stamens have free filaments and dorsifixed, nearly globose anthers that open by lengthwise slits. The ovary is composed

Microtea debilis, Bahía, Brazil [330]

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Phylogeny and evolution: Like Gisekia, Microtea was previously included in Phytolaccaceae, within which it was usually associated with Lophiocarpus (now Lophiocarpaceae). Molecular evidence places it as sister to the rest of core Caryophyllales (minus Macarthuriaceae), in a rather isolated position. It has morphological characters like those of Amaranthaceae. The genus is not well known and needs more study.

markets in tropical America, where they are used for kidney problems (especially proteinuria). The f lavonoid cirsimarin triggers lipid mobilisation, and it can be used by athletes to enhance their performance. Etymology: Microtea is probably derived from Greek μικρότητα (mikrotita), smallness, in reference to the small stature of the plant and its flowers, but a derivation is not provided in the original publication of the name, so we cannot be certain.

331. CARYOPHYLLACEAE Carnation family

Genera and species: The sole genus in this family is Microtea, with nine species. Uses: As jumby pepper, Microtea species are offered in traditional, medicinal plant Illecebrum verticillatum, Bergius Botanical Garden, Stockholm, Sweden [331]

Silene uniflora, Lizard Peninsula, Cornwall, UK [331]

CARYOPHYLLALES

EUDICOTS

Cerastium tomentosum, private garden, Hengelo, the Netherlands [331]

Arenaria dyris, Royal Botanic Gardens, Kew, UK [331]

Dianthus longicalyx, Royal Botanic Gardens, Kew, UK [331]

This family consists mostly of annual and perennial herbs, sometimes small shrubs and rarely treelets. Plants are usually bisexual, sometimes unisexual and grow terrestrially or on rocks, and stems are often swollen at the nodes. Roots sometimes form tubers or taproots, and perennial plants usually survive adverse seasons with their woody roots; underground rhizomes are rare. Leaves are mostly opposite or appearing whorled, rarely alternate, and they are often connate at the base across the node, usually without stipules, which when present are scarious. Blades are simple and entire and sessile or petiolate. Inflorescences are usually terminal, bracteate dichasia, sometimes a panicle or reduced to a monochasium, or flowers are solitary. The actinomorphic (rarely weakly zygomorphic) flowers are usually bisexual (sometimes unisexual). The usually five (rarely four) sepals are free or fused, veined or not. Petals are usually five, sometimes four or absent, free, entire, toothed, emarginate or bifid, sometimes dissected, often clawed, sometimes with corona scales on top of the claw. Stamens are usually as many or twice as many as sepals, or absent in female flowers, free or filaments fused. Anthers are versatile and open with longitudinal slits. A nectary is often found as a disc at the base of the stamens or in the tube of the filaments. The superior

(rarely partly inferior) ovary is composed of two to five fused carpels, topped with free or more or less basally fused styles, and stigmas are as many as the styles. The ovary is sometimes borne on a stalk (gynophore) or the entire petal-stamen-ovary unit borne on a stalk (anthophore). The fruit is usually a capsule that opens from the tip into as many or twice as many teeth as there are carpels, exposing numerous seeds. The fruit is an indehiscent berry, nutlet or achene with one or two seeds.

Phylogeny and evolution: Fossil records are rare, but the oldest palynological fossil is known from the Late Cretaceous (c. 73 million years old), which has been matched with macrofossils of Caryophyllif lora from Tasmanian Eocene sediments. The diversification rate of European Dianthus is one of the highest known for any group of organisms. The traditional subfamilies of Caryophyllaceae are not upheld by molecular systematics, Paronychioideae forming a grade. A subfamilial classification of this family has therefore been abandoned. On the basis of molecular results, several genera have been recently recircumscribed: Minuartia, which was polyphyletic in the traditional sense, has been split into 11 genera.

Distribution: Caryophyllaceae are one of the two families with the widest distributions, but with their greatest diversity in temperate climates. They are only absent from the permanent ice sheets in Antarctica and Greenland and are less diverse in the tropics, being absent from parts of tropical Asia and the Brazilian Highlands. Caryophyllaceae can be found in extreme habitats. Arenaria bryophylla is one of the three vascular plants that grows at high elevations; it can be found at 6,180 m on the upper slopes of Mount Everest. Antarctic pearlwort, Colobanthus quitensis, is one of two angiosperms, with the Antarctic hair grass (Deschampsia antarctica, Poaceae), that grows south of the Antarctic Circle. Genera such as Acanthophyllum diversified in xeric habitats.

Genera and species: Caryophyllaceae include 91 genera, with about 2,625 species: Acanthophyllum (57), Achyronychia (1), Agrostemma (2), Allochrusa (7), Ankyropetalum (4), Arenaria (c. 300), Augustea (3), Bolanthus (8), Brachystemma (1), Bufonia (20), Calycotropis (1), Cardionema (6), Cerastium (c. 100), Cerdia (4), Chaetonychia (1), Cherleria (19), Colobanthus (20), Cometes (2), Corrigiola (10), Cyathophylla (1), Dianthus (c. 320), Diaphanoptera (6), Dicheranthus (1), Drymaria (48), Drypis (1),

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CARYOPHYLLALES Eremogone (c. 90), Facchinia (5), Geocarpon (1), Gymnocarpos (10), Gypsophila (150), Habrosia (1), Haya (1), Herniaria (48), Holosteum (4), Honckenya (1), Illecebrum (1), Kabulia (1), Krauseola (1), Lepyrodiclis (3), Loeflingia (7), Mcneillia (5), Microphyes (3), Minuartia (c. 55), Minuartiella (4), Moehringia (c. 25), Moenchia (3), Mononeuria (9), Ochotonophila (1), Ortegia (1), Paronychia (110), Pentastemonodiscus (1), Petrorhagia (33), Philippiella (1), Phrynella (1), Pirinia (1), Pleioneura (1), Plettkea (4), Pollichia (1), Polycarpaea (50), Polycarpon (c. 10), Polytepalum (1), Pseudocherleria (c. 12), Pseudostellaria (21), Pteranthus (1), Pycnophyllopsis (2), Pycnophyllum (17), Rhodalsine (1), Robbairea (2), Sagina (c. 20), Sanctambrosia (1), Saponaria (40), Schiedea (28), Scleranthopsis (1), Scleranthus (c. 10), Scopulophila (2), Silene (c. 700), Spergula (5), Spergularia (c. 60), Sphaerocoma (2), Stellaria (c. 120), Stipulicida (1), Subulina (65), Telephium (5), Thurya (1), Thylacospermum (1), Triplateia (1), Uebelinia (3), Vaccaria (1), Velezia (6), Wilhelmsia (1) and Xerotia (1).

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EUDICOTS

also grown as a cut flower and traditional garden plant and has a similar scent. Soapwort, Saponaria officinalis, was formerly used as a soap substitute. Double-flowered forms of ‘baby’s breath’, Gypsophila paniculata, are popular additions to bridal bouquets, and it is also a popular garden plant. Gypsophila rokejeka is used to flavour halva in the eastern Mediterranean. Sea purslane, Honckenya peploides, is the only plant traditionally eaten by the Alaskan Inuit. Chickweed, Stellaria media, is a troublesome garden weed but can be eaten cooked or fresh when young. Sagina procumbens is also a weed, commonly growing in cracks in paving etc. Many species are grown as ornamental plants in cottage gardens and rockeries.

Record: Seeds of Silene stenophylla from a subfossil squirrel stash, estimated to be c. 31,800 years old were germinated and grew into plants that produced viable seeds. This is the longest seed dormancy recorded. Etymology: Caryophyllus is composed of the Greek κάρυο (karyo), a nut, and φύλλων ( fyllon), a leaf, in reference to the walnut-like scent of carnations. Caryophyllus is a later synonym of Dianthus.

332. ACHATOCARPACEAE Snake-eyes family

Uses: Carnation or clove pink (Dianthus caryophyllus) has been cultivated since classical times for its scent and as a source of oil to make soap. Many Dianthus hybrids (often D. caryophyllus × D. plumarius) are grown for the cut-flower industry or as garden ornamentals. Sweet William (D. barbatus) is

Carnivory: Various species of Silene have sticky hairs and are known by the common name of ‘catchfly’. Despite this, they are usually not considered carnivorous. Silene viscaria, for example, is one of the plants that has developed sticky leaves as a defence against feeding insects, but the insects thus trapped are not digested because these plants have no digestive enzymes. However, it has been demonstrated that S. viscaria and others (e.g. Cerastium arvense, Stellaria americana and S. jamesiana) have protease activity, but uptake of the products of digestion has not yet been investigated. Further research would be necessary to demonstrate full carnivory in this family.

This family of shrubs and small trees has branchlets that are often spinescent, sometimes with scattered thorns. Leaves are alternate, sometimes fasciculate, simple and without stipules. Blades are spatulate or oblanceolate to elliptic, margins are entire and venation is pinnate. Inflorescences are axillary racemes or panicles, or flowers are solitary; bracts are thin and linear-subulate, and bracteoles are present and minute or

Herniaria glabra, Turku harbour, Finland [331]

Drymaria cordata, Ecuador [331]

Gypsophila paniculata, Old Mission Peninsula, Michigan, USA [331]

Christenhusz, Fay & Chase

CARYOPHYLLALES

EUDICOTS

Achatocarpus pubescens, male, Nicaragua (OO) [332]

Achatocarpus pubescens, female, Nicaragua (OO) [332]

absent. The unisexual flowers of both genders occur on the same plant. The perianth consists of one whorl, usually of five, sometimes of four (Phaulothamnus), imbricate, distinct petals that are persistent in fruit. Male flowers have ten to 20 exserted stamens with filiform filaments that are free or fused at the base and have basifixed, oblong anthers. Female flowers are composed of two carpels forming a unilocular superior ovary with two or three sessile, filiform styles. Fruits are whitish or blackish, often translucent, sour or bitter berries with a single seed.

Genera and species: This family includes two genera and about 11 species, Achatocarpus (c. 10) and Phaulothamnus spinescens. Etymology: Achatocarpus is composed of the Greek words αχατος (achatos), an agate, and καρπός (karpos), a fruit, in reference to the translucent white berries of some species.

333. AMARANTHACEAE Spinach family

Distribution: This family occurs from southern North America (Mexico, Texas and California) through Central America to northern South America, with a disjunct population in southern South America (Peru, Bolivia, Paraguay and Argentina). Phylogeny and evolution: Achatocarpaceae have been included in Phytolaccaceae by numerous authors, probably due to the similarity in having racemes with berries. However, they are distinguished from Phytolaccaceae by a number of characters such as unisexual f lowers, pollen with obscure pores, unilocular ovaries with single ovules and different wood-anatomical structures. There are chemical characters that further separate Achatocarpaceae from Phytolaccaceae.

This family comprises unisexual and bisexual, annual and perennial herbs and shrubs, rarely vines and trees. The simple leaves are alternate or opposite, lack stipules, are often succulent, well-developed or reduced and then the stems succulent and green. Blades are flat or cylindrical, usually entire, sometimes lobed or pinnatifid. Inflorescences are cymose, often aggregated into loose thyrses, racemes or panicles or in dense heads or spikes. The often papery bracts can be green, white or brightly coloured and subtend one or several

Phaulothamnus spinescens (HP) [332]

flowers. The unisexual or bisexual flowers are actinomorphic and subtended by two bracts. Sterile flowers in the inflorescence are sometimes present and can be modified into scales, spines, bristles or hairs. The three to five (rarely up to eight) free or (basally) fused tepals are usually present and persistent, sometimes absent, and they can be green, white and sepal-like or brightly coloured and petal-like. There are as many stamens as there are tepals, rarely fewer or more, with the filaments all free, fused completely or fused only basally into a cup. Anthers are basifixed and open by one or two longitudinal slits. The superior (rarely partly inferior) ovary is unilocular and tipped with a short or long style and a capitate or branched stigma. Fruits are irregularly rupturing capsules, nuts or achenes, often enclosed in the persistent tepals and sometimes opening with a lid. Distribution: This is a more or less cosmopolitan family, but they are especially common and diverse in arid warm-temperate and subtropical regions and saline habitats. Phylogeny and evolution: This family evolved c. 47–87 million years ago. Traditionally two closely related families, Amaranthaceae and Chenopodiaceae, were recognised, albeit considered closely related in every classification. However, the latter was found to be polyphyletic, with

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450

EUDICOTS

Genera and species: Amaranthaceae have an estimated 170 genera with c. 2,040 species, which are placed in three subfamilies: Polycnemoideae – Hemichroa (1), Nitrophila (5), Polycnemum (8) and Surreya (2); Amaranthoideae – Achyranthes (c. 9), Achyropsis (6), Aerva (9), Allmania (2), Allmaniopsis (1), Alternanthera (c. 80), Amaranthus (70), Arthraerva (1), Blutaparon (4), Bosea (3), Brachylepis (8), Calicorema (3), Celosia (65), Centema (2), Centemopsis (3), Centrostachys (1), Chamissoa (2), Charpentiera (6), Chionothrix (3), Cyathula (25), Dasysphaera (4), Deeringia (12), Digera (1), Eriostylos (1), Froelichia (16), Froelichiella (1), Gomphrena (c. 125), Guilleminea (5),

Amaranthus retroflexus, Bucharest, Romania [333]

Ptilotus obovatus, near Coober Pedy, South Australia [333]

Achyranthes arborescens, Royal Botanic Gardens, Kew, UK [333]

Gomphrena haageana, Ruissalo Botanical Garden, Turku, Finland [333]

Froehlichia juncea, Galápagos Islands [333]

Christenhusz, Fay & Chase

Therefore the subfamilial classification and accepted genera below are just an estimate, and only three of the eight subfamilies are here followed.

Hebanthe (7), Helicilla (1), Henonia (1), Herbstia (1), Hermbstaedtia (5), Indobanalia (1), Irenella (1), Iresine (c. 70), Kyphocarpa (4), Lagrezia (12), Leucosphaera (1), Lithophila (3), Lopriorea (1), Manochlamys (1), Marcelliopsis (3), Mechowia (2), Nelsia (2), Neocentema (2), Nothosaerva (1), Nototrichium (2), Nyssanthes (2), Pandiaka (12), Pedersenia (7), Pfaffia (26), Pleuropetalum (3), Pleuropterantha (3), Polyrhabda (1), Pseudogomphrena (1), Pseudoplantago (2), Pseudosericocoma (1), Psilotrichopsis (3), Psilotrichum (18), Ptilotus (90), Pupalia (4), Quaternella (3), Rosifax (1), Saltia (1), Sericocoma (6), Sericocomopsis (2), Sericorema (2), Sericostachys (1), Siamosia (1), Stilbanthus (1), Tidestromia (6), Trichuriella (1), Volkensinia (1), Wadithamnus (1) and Woehleria (1); Chenopodioideae – Acroglochin (2), Agriophyllum (6), Allenrolfea (3), Anabasis (42), Anthochlamys (2), Aphanisma (1),

Chenopodiaceae subfamily Polycnemoideae sister to the rest. Genera of Polycnemoideae diversified during the Miocene and Pliocene, but are species-poor and appear relictual. Several classifications have been proposed, and up to eight subfamilies are sometimes recognised. Genera have also recently been revised, for instance, Chenopodium was found to be polyphyletic, with Atriplex, Dysphania, Einadia, Rhagodia and Spinacia embedded in it. Even though these genera share characters and a single genus concept could have been adopted, a greater number of genera are now accepted in this group (e.g. Blitum, Chenopodiastrum, Lipandra, Oxybasis), making the morphological differences between these genera minute. Similarly the Australian genus Maireana, North American Atriplex, Iresine and Gomphrena are all in need of generic recircumscription, and other genera in this family are likely to follow suit.

CARYOPHYLLALES

EUDICOTS

Archiatriplex (1), Arthrocnemum (5), Arthrophytum (9), Atriplex (c. 250), Axyris (5), Bassia (13), Beta (c. 12), Bienertia (2), Blitum (13), Borsczowia (1), Camphorosma (11), Ceratocarpus (2), Chenopodiastrum (5), Chenopodium (c. 110), Choriptera (3), Corispermum (65), Cornulaca (6), Cremnophyton (1), Cyathobasis (1), Cycloloma (1), Didymanthus (1), Dissocarpus (4), Dysphania (c. 32), Enchylaena (2), Eremophea (2), Exomis (2), Fadenia (1), Gamanthus (5), Girgensohnia (3), Grayia (1), Hablitzia (1), Halanthium (3), Halarchon (1), Halimocnemis (19), Halocharis (13), Halocnemum (1), Halogeton (5), Halopeplis (3), Halosarcia (c. 25), Halostachys (1), Halothamnus (21), Haloxylon (26), Heterostachys (2), Holmbergia (1), Horaninovia (7), Kalidium (5), Kirilowia (2), Krascheninnikovia (3), Lagenantha (2), Lipandra (1), Maireana (57), Malacocera (4), Microcnemum (1), Microgynoecium (1), Monolepis

(6), Nanophyton (3), Neobassia (3), Neokochia (2), Noaea (5), Nucularia (1), Ofaiston (1), Oreobliton (1), Oxybasis (5), Pachycornia (1), Panderia (1), Petrosimonia (11), Piptoptera (1), Rhagodia (11), Roycea (3), Salicornia (c. 10), Salsola (130), Sarcocornia (c. 15), Sclerolaena (64), Sclerostegia (5), Seidlitzia (7), Sevada (1), Spinacia (3), Suaeda (c. 110), Suckleya (1), Sukhorukovia (1), Sympegma (1), Tecticornia (3), Tegicornia (1), Threlkeldia (2), Traganopsis (1), Traganum (2) and Zuckia (1). Uses: The beet (Beta vulgaris subsp. vulgaris) is economically the most important species of this family. Sugar beet (B. vulgaris var. altissima) has become a competitor for sugar cane and is now responsible for 20% of global sugar production. Use of beets to produce sugar was developed in 16th century France. Beetroot, especially cultivars of B. vulgaris var. conditiva, have been a popular

vegetable since Mediaeval times. Fodder beet or mangold (B. vulgaris var. crassa) was developed in the 18th century to feed livestock. A beer was also brewed in England from fodder beet. All beets share a common wild ancestor, the sea beet (B. vulgaris subsp. maritima). Quinoa (Chenopodium quinoa) is an important grain originating in the Andes. It is rich in amino acids and has recently been popularised as a healthy cereal. It is a cultigen derived from C. berlandieri. Similarly C. macrocarpum, C. pallidicaule, Amaranthus caudatus and Lipandra polysperma are grown for their edible seeds on a smaller scale. Species of Salicornia are collected from saltmarshes and sold as samphire, a luxury vegetable. Similarly Salsola soda is cultivated as a salad herb in Mediterranean Europe. Spinach (Spinacia oleracea) is known to have been cultivated in Mesopotamia

Dysphania sp., near Coober Pedy, South Australia [333]

Beta vulgaris subsp. maritima, near Aberystwyth, Wales, UK [333]

Atriplex holocarpa, near Oodnadatta, South Australia [333]

Sarcocornia ramosissima, East Sussex, England, UK [333]

Sclerolaena longicuspis, near Oodnadatta, South Australia [333]

Maireana turbinata, near Coober Pedy, South Australia [333]

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CARYOPHYLLALES

Spinacia sativa, private allotment, Kingston upon Thames, Surrey, UK [333]

EUDICOTS

Chenopodium album, private allotment, Kingston upon Thames, Surrey, UK [333]

and was introduced to Europe around 1000 AD. It is rich in vitamins, iron and calcium. Several other species are also cultivated and cooked like spinach, e.g. Chinese spinach (Amaranthus tricolor), orache (Atriplex hortensis), chard (Beta vulgaris subsp. vulgaris var. vulgaris), blite or Good King Henry (Blitum bonus-henricus), goosefoot (Chenopodium album), huauzontle (C. berlandieri), mfungu (Celosia argentea), mukunawanna (Alternanthera sessilis) and vine spinach (Hablitzia thamnoides). Sand rice (Agriophyllum squarosum) seeds are an important food in Mongolia, and seeds of iodine bush (Allenrolfea occidentalis) are also edible. Leaves and seeds of Amaranthus hybridus (smooth pigweed) have been eaten for c. 6,000 years in Central America. Seeds of epazote or wormseed (Dysphania ambrosioides) and yerba del zorillo (D. graveolens) are used as a food flavouring in

452

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tropical America, and Hawaiian goosefoot (Chenopodium oahuense) is grown as a pot herb. Inflorescences of Aerva javanica (kapok bush) are used to stuff pillows in the Middle East. Kochia or summer cypress (Bassia scoparia) is cultivated in Japan for making brooms and is often grown as a summer bedding ornamental. Several other species are grown as ornamentals, especially cultivars of joyweed (Alternanthera ficoidea), cat tail (Amaranthus caudatus), cockscomb (Celosia cristata), globe amaranth (Gomphrena globosa), bloodleaf (Iresine herbstii) and pussytail (Ptilotus manglesii) are commonly grown as ornamentals. Etymology: Amaranthus is derived from Greek α (a), not, and μαραίνω (maraino), to wither, in reference to the everlasting quality of dried inflorescences.

Chenopodium quinoa, Peru (SK) [333]

334. STEGNOSPERMATACEAE Cuban-tangle family

Stegnospermataceae are a family of small trees and shrubs with hanging and climbing branches. Their leaves are alternate and simple with short petioles and no stipules. Leaf blades have an entire margin and pinnate venation. Inflorescences are axillary or terminal racemes, thyrses or condensed, umbellate cymes. The actinomorphic flowers are bisexual and have five, free sepals and five,

CARYOPHYLLALES

EUDICOTS

free, whitish petals. The five to ten stamens are fused basally around the ovary, and filaments bear dorsifixed anthers that open by longitudinal slits. The superior ovary is composed of two to five fused carpels that are initially tri- to pentalocular, but later becoming a single locule with a central column. Stigmas are as many as carpels and are free, rarely atop a short style. The fruit is a capsule that opens from the apex into three to five valves, each having a seed covered by a white or red aril.

Uses: Stegnosperma has high saponin content in its roots, and the plants were formerly used in Baja California as a soap substitute. Etymology: Stegnosperma is composed of the Greek words στεγνός (stegnos), dry, and σπέρμα (sperma), seed, referring to the aril that covers the seed and keeps it dry.

335. LIMEACEAE Lizard-foot family

condensed into small, dense clusters. The bisexual, actinomorphic flowers have five free sepals with a membranaceous margin. Petals are absent or three to five and fused to the stamen tube. The usually seven (sometimes five to ten) stamens have fused filaments and dorsifixed anthers that open by lengthwise slits. The superior ovary is composed of two fused carpels that form one locule with a single seed. The fruit separates into two woody capsules, each holding a kidney-shaped seed.

Distribution: The family is restricted to the Sonoran Desert, the Gulf Coast of Texas and Mexico and the Greater Antilles.

Distribution: The family is restricted to dry parts of Africa, Arabia and the Indian subcontinent.

Phylogeny and evolution: Stegnosperma used to be included in Phytolaccaceae, to which they are similar morphologically. Molecular studies found them to be widely separated from this family, and because there are differences in flower and pollen structure, they are now placed in a separate family.

Phylogeny and evolution: Another member of the disintegrated Molluginaceae (and previously in a polymorphic Aizoaceae), Limeum is now placed in its own family because it would make Molluginaceae widely polyphyletic.

Genera and species: A single genus, Stegnosperma, with three species encompasses this family: S. cubense, S. halimifolium and S. watsonii.

Usually these are prostrate annual and perennial herbs, often with a woody base, that are sometimes covered with glandular hairs. Leaves are alternate to subopposite, sessile and lack stipules. Blades are succulent, linear, lanceolate, orbicular or subulate, and the margins are entire. Inflorescences are bracteate, axillary cymes, sometimes

Stegnosperma cubense, Isla Juan Venado, León, Nicaragua (CD) [334]

Genera and species: A family of a single genus, Limeum, with 21 species. Etymology: Limeum is Latin for path, which is where this plant may be found.

Limeum africanum, South Africa (JA) [335]

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CARYOPHYLLALES

336. LOPHIOCARPACEAE Sandaarbossie family

EUDICOTS

pentalocular, each locule formed by one carpel (Corbichonia). They bear a style with four or five stigmas. The fruit is a spiny, warty or ridged achene (Lophocarpus) or a loculicidal capsule (Corbichonia).

337. KEWACEAE Suring family

Distribution: This family has a patchy distribution across southern and eastern Africa and southern Arabia to western India.

Lophiocarpaceae consist of herbs that are sometimes woody at the base. Leaves are alternate, simple, with entire margins and pinnate venation, lacking stipules. Inflorescences are terminal or axillary threeflowered cymules merged into spikes that are organised in clusters of two or three (Lophiocarpus), or they are terminal or leafopposed cymes (Corbichonia). Flowers are bisexual and actinomorphic. The five sepals are free, and there are 15–20 slender petals in Corbichonia, which are absent in Lophicarpus. Stamens are usually four (Lophiocarpus) or ten to 30 (Corbichonia), with free filaments and elongate, dorsifixed anthers. The superior ovary is unilocular and composed of four, fused carpels (Lophiocarpus) or

Phylogeny and evolution: Also a former member of Phytolaccaceae, Lophiocarpus was sometimes associated with Microtea (now Microteaceae). Lophiocarpus is related to neither family directly, but found to be sister to the morphologically different Corbichonia (formerly Molluginaceae), these two together forming a clade sister to a larger clade encompassing Kewaceae, Barbeuiaceae, Aizoaceae, Phytolaccaceae and Nyctaginaceae. Genera and species: This family includes two genera and six species: Corbichonia (2) and Lophiocarpus (4). Etymology: Lophiocarpus is composed of the Greek words λοφίος (lophios), a plume, and καρπός (karpos), a fruit.

Corbichonia decumbens, Lydenburg, South Africa (CD) [336] Kewa acida, Royal Botanic Gardens, Kew, UK [337]

454

Christenhusz, Fay & Chase

These are annual and perennial herbs that can be slightly woody at base. Leaves are alternate, usually more or less fasciculate at the branch ends, succulent, linear and terete. Stipules are present and fused to the base of the blade, more or less sheathing the stem. Inflorescences are terminal or apparently axillary, long-stalked false umbels. Flowers are bisexual and actinomorphic and have a whorls of five free sepals, of which three or four become petal-like (usually white or pink). True petals are absent. The five to 15 (rarely fewer) stamens have shortly fused filaments and dorsifixed anthers that open by longitudinal slits. The superior ovary is composed of three to five fused carpels topped by short, fleshy stigmatic crests.

Kewa salsoloides, Wolwekraal Nature Reserve, Western Cape, South Africa (CD) [337]

CARYOPHYLLALES

EUDICOTS

Kewa bowkeriana, Helsinki Botanical Garden, Finland [337]

The fruit is a membranaceous capsule that opens loculicidally. Distribution: Kewaceae are found mostly in southern Africa, but extend north into East Africa and Ethiopia and also along the south coast of St Helena and in western Madagascar. They usually grow in seasonally dry environments. Phylogeny and evolution: Like Mollugo, Macarthuria and Limeum, Hypertelis was originally placed in Aizoaceae, from where it was moved to Molluginaceae, a family that was based on plesiomorphic characters and later found to be widely polyphyletic in genetic studies. The majority of Hypertelis was found to be part of a well-supported clade that includes Aizoaceae, Barbeuiaceae, Lophiocarpaceae, Phytolaccaceae and Nyctaginaceae, apart from its type species, H. spergulacea, which goes with Molluginaceae. Therefore the new name Kewa was proposed for the eight species of Hypertelis that do not go with the type of the genus, and these are placed in their own family. Genera and species: The sole genus in this family is Kewa with eight species. Uses: The leaves of endemic Kewa acida were previously eaten as salad on St Helena.

Barbeuia madagascariensis, Fianarantsoa, Madagascar (CD) [338]

Etymology: Kewa is named for the village of Kew in southwestern London, where the Royal Botanic Gardens are situated.

338. BARBEUIACEAE Liane-Barbeu family

Barbeuiaceae are lianas with simple, alternate, spirally arranged leaves without stipules. Blades have entire margins and pinnate venation. Inflorescences are axillary fascicles (racemes) with actinomorphic, bisexual flowers. The five sepals are free, and petals are absent. There are 20 to numerous stamens with free filaments inserted on a ring-shaped disk, and dorsifixed, black anthers that open by lengthwise slits. The superior ovary is composed of two fused carpels, each forming a locule topped with two sessile flattened, papillose stigmas. Fruits are bilocular capsules that split in two, exposing one or two arillate seeds.

Distribution: This family is endemic to Madagascar. Phylogeny and evolution: Barbeuia was originally placed in Rosaceae and later moved to Phytolaccaceae, in which it formed its own subfamily (mainly because it has a capsule rather than a berry). Molecular studies have shown that it is not part of Phytolaccaceae, and thus it was segregated into its own family, pending further taxonomic research on this highly divided clade of Caryophyllales. Barbeuiaceae are estimated to have diverged c. 36–38 million years ago. Genera and species: Barbeuiaceae include a single species, Barbeuia madagascariensis. It is widespread across Madagascar, although specimens from southeastern Madagascar have ref lexed sepals in fruit and may represent a separate species. Etymology: Barbeuia is named in honour of French physician, botanist, author, translator and publisher Jacques BarbeuDubourg (1709–1779), who is best known for his translation of Benjamin Franklin’s oeuvre into French.

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CARYOPHYLLALES

339. GISEKIACEAE Stork’s-henna family

EUDICOTS

and five-parted. The five free sepals are petal-like, true petals being absent. The five to 20 stamens are free. The superior ovary is composed of five to 15 carpels that are fused at the base. The fruit is composed of separate achenes that each contain a black, swollen, kidney-shaped seed. Distribution: Gisekiaceae are found in the drylands of Africa, Madagascar, the Mascarenes, Arabia, southern and southeastern Asia east to Hainan Island.

Genera and species: The sole genus in this family is Gisekia with seven ‘species’, which may all have to be treated as a single variable species, G. pharnaceoides, because the traditional, morphologically defined taxa have not been found to be genetically distinct.

Annual and perennial herbs, these plants are generally creeping or prostrate, with the stems regularly branching. The usually opposite leaves are simple and more or less succulent, bearing white lines. Axillary or terminal dichasial inf lorescences are composed of umbellate partial inf lorescences. The bisexual or (rarely) functionally unisexual flowers are radially symmetrical

Phylogeny and evolution: Sometimes placed in Phytolaccaceae, Gisekia was later moved into Aizoaceae, from which it was segregated with several other genera in Molluginaceae, now found to be widely polyphyletic within Caryophyllales. Gisekia is now placed in its own family, sister to a clade of Nyctaginaceae, Petiveriaceae, Phytolaccaceae and

Uses: Gisekia is a weed of cultivated fields and is locally used medicinally. It may have anti-inflammatory properties.

Gisekia pharnaceoides, South Africa (JA) [339]

Mesembryanthemum crystallinum, South Australia [340]

Tetragonia tetragonoides, private allotment, Kingston upon Thames, Surrey, UK [340]

Carpobrotus edulis, naturalised on the cliffs at Sea Ranch, California, USA [340]

Gisekia pharnaceoides, Kundunchi Beach, Tanzania (CD) [339]

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Sarcobataceae, although it has also been suggested that Gisekia should be part of Petiveriaceae. It is estimated that Gisekiaceae evolved c. 15 million years ago in the Miocene of southern Africa.

Christenhusz, Fay & Chase

Etymology: Gisekia is named for the German physician, botanist and librarian Paul Dietrich Giseke (1741–1796), a contemporary and friend of Carolus Linnaeus.

CARYOPHYLLALES

EUDICOTS

Delosperma kofleri, Royal Botanic Gardens, Kew, UK [340]

Lithops hookeri, Royal Botanic Gardens, Kew, UK [340]

Sesuvium edmonstonei, Galápagos Islands [340]

valves as the styles, sometimes the fruit indehiscent or a berry, nut or circumscissile capsule.

including Agdestidaceae, Barbeuiaceae, Gisekiaceae, Kewaceae, Lophiocarpaceae, Nyctaginaceae, Petiveriaceae, Phytolaccaceae and Sarcobataceae. This would make a monophyletic group and may prevent further disintegration of families and proposals of new ones, although it may be difficult to find morphological characters that distinguish such a broadly circumscribed family.

340. AIZOACEAE Dewplant family

These succulent annual and perennial herbs and shrubs usually have opposite, rarely alternate, fleshy leaves. Leaves are simple and entire, terete to trigonous or flat, with or without a petiole, the petiole sometimes sheathing and stipule-like at its base, or the blade sessile and sheathing at its base. Flowers are mostly solitary or situated in loose dichasial cymes. The bisexual flowers are regularly symmetrical and usually have five, sometimes three to eight unequal (or equal) sepals. Sepals are basally fused with the filaments, forming a tube, the inner surface of which can be green or coloured, the stamen elements can be four to many, and if many, the outer whorl(s) developed into brightly coloured petals. Staminodes are present or not, but functional stamens are usually many, rarely one or a few. The usually inferior (rarely half-inferior or superior) ovary is bito penta- (rarely more) locular and topped by free stigmas, as many as the locules. Fruits are usually multi-celled capsules, which, when wet, open into as many winged or unwinged

Distribution: This widespread family occurs in arid and winter-rain, warm-temperate climates, in North America, Mesoamerica, the Caribbean, coastal South America, the Mediterranean coasts, Africa, southern Madagascar, the Mascarenes, Arabia, India, East and Southeast Asia, Malesia, Australasia and the Pacific. The greatest diversity is found in South Africa. Phylogeny and evolution: The great diversity in southwestern Africa is related to a rapid radiation in this region c. 5 million years ago, with the establishment of the warm-temperate, arid, winter-rain climate there. Four subfamilies (Aizooideae, Mesembryanthemoideae, Ruschioideae and Sesuvioideae) are sometimes accepted. Circumscription of Mesembryanthemum has been waxing and waning in the past, and many species traditionally placed there are now treated in Carpobrotus, Conophytum, Dorotheanthus, Sceletium and so on. Recently a molecular phylogenetic study resulted in the merging of several genera back into Mesembryanthemum (e.g. Aptenia, Aspazoma, Brownanthus, Dactylopsis, Phyllobolus, Psilocaulon and Synaptophyllum). However, generic circumscription is still uncertain, as is the number of species. The family may alternatively be expanded to encompass the entire clade

Genera and species: Aizoaceae have 113 genera and c. 1,900 species: Acrodon (5), Acrosanthes (5), Aethephyllum (1), Aizoanthemum (4), Aizoon (13), Aloinopsis (8), Amphibolia (5), Antegibbaeum (1), Antimima (101), Apatesia (3), Arenifera (4), Argyroderma (10), Astridia (12), Bergeranthus (12), Bijlia (2), Braunsia (c. 5), Carpanthea (1), Carpobrotus (13), Carruanthus (2), Caryotophora (1), Cephalophyllum (38), Cerochlamys (4), Chasmatophyllum (8), Cheiridopsis (29), Circandra (1), Cleretum (3), Conicosia (2), Conophytum (86–260), Corpuscularia (8), Cylindrophyllum (5), Cypselea (3), Delosperma (155), Dicrocaulon (7), Didymaotus (1), Dinteranthus (6), Diplosoma (2), Disphyma (5), Dorotheanthus (7), Dracophilus (2), Drosanthemum (107), Eberlanzia (4), Ebracteola (5), Enarganthe (1), Erepsia (29), Esterhuysenia (4), Faucaria (6), Fenestraria (1), Frithia (2), Galenia (29), Gibbaeum (28), Glottiphyllum (16), Gunniopsis (14), Hallianthus (1), Hereroa (28), Hymenogyne (2), Jacobsenia (3), Jensenobotrya (1), Jordaaniella (4), Juttadinteria (5), Khadia (6), Lampranthus (218), Lapidaria (1), Leipoldtia (8), Lithops (37), Machairophyllum Plants of the World

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CARYOPHYLLALES (4), Malephora (16), Mesembryanthemum (c. 100), Mestoklema (6), Meyerophytum (2), Mitrophyllum (6), Monilaria (5), Mossia (1), Namaquanthus (1), Namibia (2), Nananthus (7), Nelia (2), Neohenricia (2), Octopoma (8), Odontophorus (4), Oophytum (2), Orthopterum (2), Oscularia (23), Ottosonderia (1), Pleiospilos (4), Plinthus (6), Polymita (2), Psammophora (4), Rabiea (6), Rhinephyllum (10), Rhombophyllum (5), Ruschia (219), Ruschianthus (1), Saphesia (1), Schlechteranthus (2), Schwantesia (11), Scopelogena (3), Sesuvium (22), Skiatophytum (1), Smicrostigma (1), Stayneria (1), Stoeberia (5), Stomatium (40), Tanquana (3), Tetragonia (57), Titanopsis (3), Trianthema (27), Tribulocarpus (1), Trichodiadema (34), Vanheerdea (2), Vanzijlia (1), Wooleya (1), Zaleya (7) and Zeuktophyllum (2). Uses: New Zealand spinach, Tetragonia tetragonoides, is frequently cultivated and eaten as a cooked vegetable, especially in places where it is too hot to grow true spinach. Aizoon canariense is eaten by the Tuareg in North Africa, and mboga, Sesuvium portulacastrum, is a popular vegetable in the Old Phytolacca acinosa, private garden, Kingston upon Thames, Surrey, UK [341]

EUDICOTS

World tropics. The fruits of the hottentot fig or pigface, Carpobrotus edulis, can be eaten fresh, dried or in jams, but other Carpobrotus species such as C. deliciosus and C. muirii have tastier fruits. The succulent roots of Khadia acutipetala have been added to brews to make the alcoholic beverages known as khadi more potent. Popular ornamentals are the iceplants, Dorotheanthus bellidiformis, the camouf laged ‘living stones’ of the genera Conophytum, Gibbaeum, Lithops, Oophytum, Pleiospilos and Tanquana, and the peculiar ‘window plants’ Fenestraria and Frithia, which grow underground, and have windows at the tip of the leaves that focus the light into internal tissues. Many other species are also cultivated as ornamentals, and several naturalise easily, especially in coastal habitats where they can become troublesome weeds. Carpobrotus species, for example, are particularly invasive in Cornwall and California. Etymology: Aizoon is derived from Greek αείζωον, everlasting, composed of αεί (aei), always, and ζωόσ (zóos), alive; a plant that will live under almost any circumstance. Phytolacca dioica, male, Gibraltar [341]

341. PHYTOLACCACEAE Pokeweed family

These are perennial herbs, shrubs, trees and woody climbers, often with swollen, carrotlike below-ground stems in herbaceous taxa. Leaves are alternate and simple, with entire leaf margins and pinnate venation, lacking stipules. Inflorescences are terminal or axillary spikes, thyrses or racemes with bisexual or unisexual, actinomorphic or more or less zygomorphic (Anisomeria) flowers. The four or five sepals are free, and petals are absent. Stamens are five to 30 with free filaments and dorsifixed anthers that open by lengthwise slits. The superior ovary (inferior or partly inferior in Agdestis) is composed of three to 16 free or partially fused carpels,

Ercilla volubilis, Burncoose Nurseries, Cornwall, UK [341]

Phytolacca americana, in fruit, Michigan, USA [341]

Phytolacca dioica, female in fruit, Gibraltar [341]

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each topped with a short style and a stigma. Agdestis has four completely fused carpels topped with a fused style with four recurved, papillose stigmas. Fruits are a whorl of one-seeded berries, sometimes fused into a ring-shaped berry, rarely (in Agdestis) an achene with persistent sepals that function as wings for dispersal. Distribution: Phytolaccaceae are distributed in the Americas from southeastern Canada and the Pacific USA throughout the Neotropics to Uruguay, throughout tropical and East Africa, Madagascar and Southeast Asia and temperate East Asia. Agdestidoideae are restricted to Central America, southern Florida and the Caribbean region. Phylogeny and evolution: The delimitation of Phytolaccaceae has long been a matter of debate and it has varied greatly since they were separated from Chenopodiaceae (=Amaranthaceae). Phytolaccaceae in the traditional sense were found to be polyphyletic. They used to include Barbeuia (Barbeuiaceae), Gisekia (Gisekiaceae), Lophiocarpus (Lophiocarpaceae), Microtea (Microteaceae), Phaulothamnus (Achatocarpaceae) and Stegnosperma (Stegnospermataceae). Agdestis clematidea is still unplaced and may be close to Sarcobataceae, which is why it sometimes is placed in its own family. We have included it here as it is not yet certain were it should go and what the final word will be about this Rivina humilis, Yucatán, Mexico [342]

lineage of plants. Petiveriaceae were formerly included in Phytolaccaceae (as Rivinoideae), but they are, in some phylogenetic studies, sister to Nyctaginaceae, making Phytolaccaceae polyphyletic. Nevertheless these plants are morphologically similar. Cronquist (1981) wrote that “each of the several segregate families. .. appears to be a natural group, but collectively they all hang together with the rest of the Phytolaccaceae. I see no reason why they cannot be accommodated at the level of tribes or subfamilies.” A taxonomic revision accompanied by well-sampled phylogenetic studies is greatly needed. Fossil fruits similar to those of Phytolacca are known from the Upper Cretaceous of Mexico.

volubilis (climbing pokeweed) and Phytolacca dioica (ombú or tree pokeweed) are occasionally grown as garden ornamentals. Etymology: Phytolacca is composed of the Greek φυτών ( fyton), a plant, and λάκκα (lakka), lac, a scarlet resinous secretion of several insect species, formerly used as a red dye. Similarly, the berries of Phytolacca can be used to produce a red dye; therefore the name means ‘vegetable lac’.

342. PETIVERIACEAE Henwood family

Genera and species: Phytolaccaceae consist of five genera and c. 33 species in two subfamilies: Agdestidoideae – Agdestis (1); Phytolaccoideae – Anisomeria (3), Ercilla (2), Nowickea (2) and Phytolacca (c. 25). Uses: Youngs shoots and leaves of Indian pokeweed (Phytolacca acinosa) are commonly eaten in Asia. American pokeweed (P. americana) has antiviral and molluscicidal medicinal properties. Berries of several Phytolacca species yield a red dye, which can be used to colour food, wine and sweets. Cultivated frequently, they have become common weeds in temperate and subtropical gardens worldwide. Some species such as Agdestis clematidea (rockroot), Ercilla

Hilleria latifolia, Botanical Garden, Berlin-Dahlem, Germany [342]

These are trees, shrubs (often scandent), vines and herbs, sometimes spiny (Seguieria), occasionally smelling of garlic (Gallesia, Petiveria and Seguieria). Leaves are alternate, simple with pinnate venation and entire margins. Inflorescences are axillary or terminal panicles or racemes. Flowers are bisexual or unisexual (Ledenbergia) and usually actinomorphic, rarely zygomorphic (Hilleria). The four sepals (five in Seguieria)

Petiveria alliacea, Botanical Garden, Berlin-Dahlem, Germany [342]

Trichostigma peruvianum, Ruissalo Botanical Garden, Turku, Finland [342]

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CARYOPHYLLALES are free (the lower three somewhat fused in Hilleria), and petals are absent. The four to 65 stamens have free filaments and dorsifixed anthers that open by longitudinal slits. Four to six staminodes are present in the female flowers of Ledenbergia. The superior ovary is composed of a single carpel with a short style (sometimes flattened or absent) and a capitate, papillose, feathery or penicillate stigma. Fruits are variable: samaras (Gallesia and Seguieria), berries (Rivina and Trichostigma), utricles (Hilleria, Ledenbergia and Schindlera) and achenes (Monococcus and Petiveria). Distribution: Nearly restricted to the Neotropics, but one genus (Monococcus) is found in eastern Australia, New Caledonia and the New Hebrides. Rivina humilis is naturalised pantropically. Phylogeny and evolution: Previously included in Phytolaccaceae as subfamily Rivinoideae or Petiverioideae, it is now known that Petiveriaceae are more closely related to Nyctaginaceae than to the rest of Phytolaccaceae, although support for this is not yet strong. Genera and species: Petiveriaceae include nine genera and 20 species: Gallesia (1), Hilleria (3), Ledenbergia (2), Monococcus (1), Petiveria (1), Rivina (1), Schindlera (2), Seguieria (6) and Trichostigma (3). Uses: Guinea henwood (Petiveria alliacea) is used locally as a mouthwash to treat toothache. It smells strongly of garlic, and its fruits are strongly insecticidal; it is a good treatment against whitefly. Bloodberry (Rivina humilis) has red fruits that also yield a red dye. Hilleria latifolia is eaten as a potherb in West Africa, and bark strips of Trichostigma octandrum are used for basketry in the West Indies. Petiveriaceae are high in saponins and can be used as a substitute for soap. Rivina humilis and Trichostigma peruviana are occasionally grown as garden ornamentals. Etymology: Petiveria is named in honour of James Petiver (c.1665–1718), a London apothecary, who studied the plants and insects

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of Britain and India. He collected many specimens, and after his death his famous collection was purchased by Hans Sloane for the large sum of £4,000. It now resides in the Natural History Museum of London.

343. SARCOBATACEAE Greasewood family

This is a family of succulent, unisexual and bisexual shrubs with woody stems bearing leaves and one or two axillary thorns per node. The alternate or tufted leaves are sessile, fleshy, linear, nearly round in cross-section or somewhat flattened above. Inflorescences are unisexual, the male inflorescences terminal, peltately bracteate spikelets, and the female flowers are solitary or paired in the leaf axils. Flowers lack bracteoles; the male flowers do not have a perianth, and the stamens are sessile on the inflorescence stalk, arising around the stalk of the persistent, peltate, scale-like inflorescence bract. Female flowers have a fused, cup-like, shallowly lobed perianth that is fused to the ovary and forms a wing in fruit, topped with two stigmas. The fruit is a winged nutlet, the wing tinged red and the seed conical at the tip. Distribution: The family occurs in western North America, from southeastern British Columbia and southwestern Alberta in Canada through the dry areas of the western USA to Coahuila in northern Mexico, especially in saline habitats. It was first discovered during the Lewis and Clarke expedition in 1806. Phylogeny and evolution: This family was formerly treated as a tribe in Chenopodiaceae subfamily Salsoloideae, but it is now known that they are not directly related to

Sarcobatus vermiculatus, male inflorescence, Royal Botanic Gardens, Kew, UK [343]

Amaranthaceae (including Chenopodiaceae), but rather are closer to Phytolaccaceae and Nyctaginaceae. Genera and species: The sole genus is Sarcobatus with two closely related species, S. baileyi and S. vermiculatus. Uses: The hard yellow wood has been used as firewood by Native Americans and early settlers. The leaves are poisonous to grazing animals. Etymology: Sarcobatus is composed of the Greek words σαρκός (sarkos), flesh, and βάτος (batos), a bramble bush.

344. NYCTAGINACEAE Four-o’clocks family

CARYOPHYLLALES

EUDICOTS

Abronia latifolia, Sea Ranch, California, USA [344]

Boerhavia diffusa, Malacca, Malaysia [344]

Cryptocarpus pyriformis, Galápagos Islands [344]

Mirabilis nyctaginea, Helsinki Botanical Garden, Finland [344]

Bougainvillea spectabilis, Singapore [344]

Pisonia umbellifera, Royal Botanic Gardens, Kew, UK [344]

Perennial and annual herbs, shrubs, trees and spiny vines make up this family. Roots are sometimes fleshy or tuberous, and stems are often swollen at the nodes. Leaves are opposite, alternate or whorled, often unequal in size, without stipules, usually petiolate. Blades are simple, herbaceous or slightly fleshy, the margins entire or sinuate. The mostly terminal, occasionally axillary, inflorescences are cymes, umbels or whorls, often grouped into panicles, sometimes one-flowered or fascicled, bracteate. Bracts are variable, small or large, persistent or not, and sometimes forming a calyx-like, brightly coloured involucre. The bisexual (rarely unisexual) flowers are actinomorphic, rarely zygomorphic, the (three to) four to five (to seven) tepals are fused into a tubular or funnel-, urn- or bell-shaped perianth, the lower part surrounding the maturing ovary, the apex five- to ten-lobed. Stamens are usually three or five, sometimes one, ten or up to 40, the unequal filaments free or fused at the base. Anthers are nearly basifixed and open with longitudinal slits. Ovaries are superior, unilocular and composed of a single carpel topped with a (short to long) style and a globose- or otherwise-shaped stigma.

Fruits are achenes or nutlets enclosed by the modified persistent perianth base (anthocarp), which is ribbed or winged and often glandular.

(2), Phaeoptilum (1), Pisonia (40), Pisoniella (1), Ramisia (1), Reichenbachia (2) and Salpianthus (1).

Distribution: A pantropical family, extending into the temperate zones in North America, East Asia and Australia.

Uses: The tubers of mauca (Mirabilis expansa) and M. multiflora have been eaten for food since prehistoric times in tropical America and are still used for food in some traditional societies in Mesoamerica and the Andes. Extracts of mauca have been shown to suppress tumourous activity in cells. The young leaves of the lettuce tree or Moluccan cabbage, Pisonia ‘Alba’, are eaten in Indonesia. Species of Neea yield a black dye, and caparrosa (N. theifera) is used as a tea substitute. Several species are cultivated as ornamentals, especially four o’clock flower, Mirabilis jalapa, and cultivars of the hybrid bougainvilleas, Bougainvillea ×buttiana (= B. glabra × B. peruviana) and B. spectabilis.

Phylogeny and evolution: Nyctaginaceae form a clade with Phytolaccaceae, Gisekiaceae and Sarcobataceae, to which Aizoaceae are sister. It is not clear if the four families can be maintained separately. Petiveriaceae are closer to Nyctaginaceae than to Phytolaccaceae in which these genera were previously included. Nyctaginaceae diversified remarkably in the deserts of North America during the Oligocene and Miocene. Genera and species: This family includes 27 genera and c. 355 species: Abronia (24), Acleisanthes (16), Allionia (2), Andradea (1), Belemia (1), Boerhavia (c. 50), Boldoa (1), Bougainvillea (18), Caribea (1), Cephalotomandra (1), Colignonia (6), Cryptocarpus (1), Cuscatlania (1), Grajalesia (1), Guapira (c. 50), Leucaster (1), Mirabilis (c. 60), Neea (c. 70), Neeopsis (1), Nyctaginia (1), Okenia

Carnivory: Claims that the seeds of Pisonia grandis trap and kill seabirds for the added nutritional benefit of germinating near a decomposing corpse have been investigated. There were numerous reports of dead birds that apparently died after becoming entangled in the sticky infructescences of this coastal tree

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CARYOPHYLLALES species, but the possible benefit of nutrients to the germinating plants was outweighed by scavenging crabs that were attracted to the decomposing birds. The killing of birds by P. grandis seeds, however morbid, seems more an adaptation to distribute seeds effectively to various oceanic, guano-rich islands than an actual adaptation for carnivory. Similarly, the stems and leaves of Mirabilis longiflora bear viscid hairs that catch many insects, but the leaves have no power of absorption, the hairs being merely a protection against predation. Etymology: Nyctago is composed of Greek νυκτός (nyktus), night, and the suffix -ago, like, because some species flower at night. It is a later illegitimate synonym of Mirabilis, which means ‘wonderful’ in Latin. The family name is not based on Nyctaginia.

345. MOLLUGINACEAE Carpetweed family

EUDICOTS

opposite, often crowded in a basal rosette or in false whorls. Stipules are membranaceous or absent. Blades are often succulent, the margins entire and venation pinnate. Inflorescences are terminal (or seemingly axillary) cymes or false umbels, and sometimes flowers are solitary in the leaf axils. Flowers are bisexual (rarely unisexual) and actinomorphic, often small and unremarkable. The usually five (sometimes four) sepals are free or rarely slightly fused at the base, thin and membranaceous. Petals are usually absent, but in Glinus there can be up to eight (rarely more) petals (or petal-like staminodes), each with a split apex. Usually five (rarely fewer or up to 25) stamens are typically fused into a short tube at the base of filaments in fascicles or free. Anthers are dorsifixed, versatile and open by lengthwise slits. A superior ovary is composed of one to five fused carpels, each forming a locule, with as many styles or sessile stigmas as there are carpels. Fruits are loculicidal capsules, often membranaceous, rarely a nutlet (Adenogramma). Distribution: Molluginaceae are distributed across North, Central and South America, the West Indies, Spain, the Balkans, southern Russia, Turkmenistan, most of Africa and Arabia, Madagascar and Australia.

Annual and perennial herbs (often with a woody base) and cushion plants make up this family. The simple leaves are alternate, rarely

Phylogeny and evolution: The taxonomic placement of genera of Molluginaceae has been problematic in the past. They have been considered members of Aizoaceae,

Psammotropha mucronata, Umtamvuna Nature Reserve, Eastern Trigastrotheca pentaphylla, Cape, South Africa (CD) [345] Singapore (KH) [345]

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Glinus oppositifolius, Pilbara, Western Australia [345]

Nyctaginaceae or Phytolaccaceae but were segregated on the basis of some characters that were later found to be plesiomorphic. Phylogenetic studies have shown that the family in its original circumscription was polyphyletic, with Corbichonia (Lophiocarpaceae), Limeum (Limeaceae), Macarthuria (Macarthuriaceae) and Kewa (Kewaceae) not being part of Molluginaceae in the strict sense. Corrigiola and Telephium were sometimes placed here, but belong to Caryophyllaceae, despite their alternate leaf arrangement. The crown group of Molluginaceae is estimated to be c. 46–50 million years old. The type of Hypertelis belongs in Molluginaceae, but the rest of that genus is placed in a separate family Kewaceae, which is corroborated by the morphology of the species. Genera and species: Molluginaceae include 11 genera and c. 96 species: Adenogramma (11), Coelanthum (3), Glinus (10), Hypertelis (5), Mollugo (15), Paramollugo (9), Pharnaceum (25), Polpoda (2), Psammotropha (11), Suessenguthiella (1) and Trigastrotheca (4). Uses: Several species of Glinus and Mollugo, especially M. pentaphylla, are locally eaten as potherbs. Etymology: Mollugo is derived from Latin mollis, soft, and the suffix -ago, like. The name was given to the genus in reference to Galium mollugo (Rubiaceae), which it resembles due to its whorled leaves. Adenogramma glomerata, Papkuilsfontein, Northern Cape, South Africa (CD) [345]

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EUDICOTS

Calandrinia polyandra, Coober Pedy, South Australia [346]

Claytonia perfoliata, Royal Botanic Gardens, Kew, UK [346]

Claytonia sibirica, Royal Botanic Gardens, Kew, UK [346]

Lewisia rediviva, Royal Botanic Gardens, Kew, UK [346]

[346]

Montia fontana, Stott Park, Lake District, UK

Phemeranthus humilis, New Mexico, USA (DZ) [346]

nine per flower and then sepal-like. Flowers are bisexual and actinomorphic. There are two usually persistent sepals and four, five or six (or up to 19) petals that are free or sometimes basally fused. Stamens are as many as petals and opposite them (or one fewer and alternating with the petals in Lyallia), rarely numerous (up to 100). Filaments are free or basally fused to each other or to the petals, and anthers are basifixed and open by longitudinal slits. The superior ovary is composed of two to eight carpels that are fused to form a single locule. A more or less well-developed style has diverging branches (as many as carpels), each with a papillate stigma. Fruits are circumscissile or loculicidal capsules with many seeds or an indehiscent capsule with a single seed.

the Falkland Islands, several high mountains in East Africa, Kerguélen and other Antarctic islands, New Guinea, Australia, Tasmania and New Zealand.

346. MONTIACEAE Blinks family

Montiaceae are annual and perennial herbs, rarely with a woody base or somewhat woody cushion plants (Lyallia). They often have swollen roots and form a basal rosette of leaves with short internodes. Leaves are alternate, spirally arranged and often have broad clasping bases, closely imbricate or widely spaced, without stipules. Leaf blades are flat or terete, with entire leaf margins and an often obscure, pinnate venation. Inflorescences are terminal or axillary cymes, or flowers are solitary in the leaf axils. Bracts are free or sometimes fused around the stem. Bracteoles are absent or (in Lewisia) up to

Distribution: This family is widespread across the Northern Hemisphere, where it is found in temperate and Arctic North America, in the mountains in Mexico and the Antilles, throughout Europe and Anatolia and in northeastern Siberia and Kamtchatka. In the Southern Hemisphere it is found in the Andes,

Phylogeny and evolution: Montiaceae evolved c. 13 million years ago, and their current wide distribution is therefore the result of frequent long-distance dispersals. The family has radiated especially in seasonably inhospitable regions, such as areas that have prolonged winters or prolonged droughts. Lyallia is a Tertiary relict, and these sub-Antarctic cushion plants used to be placed in their own family Hectorellaceae. They share morphological characters with Calandrinia and anatomical characters with the other members of Montiaceae. Phemeranthus was previously included in Talinum (now Talinaceae) but belongs to this family. Both are segregate families of a disintegrated Portulaceae. Calyptridium and Philippiamra are now synonyms of Cistanthe; Hectorella now belongs to Lyallia; Parakeelya is part of Calandrinia; and Lewisiopsis is returned to its placement in Lewisia. Plants of the World

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CARYOPHYLLALES Genera and species: Montiaceae include c. 10 genera and c. 295 species: Calandrinia (c. 150), Cistanthe (35), Claytonia (27), Lenzia (1), Lewisia (19), Lyallia (2), Montia (12), Montiopsis (18), Phemeranthus (c. 30) and Schreiteria (1). Uses: The fresh, succulent leaves of some species of Calandrinia, Claytonia and Montia, especially miner’s lettuce or winter purslane (Claytonia perfoliata), are regionally eaten as salad. The root of bitter root, Lewisia rediviva, is starchy and was formerly a staple food for native Americans, important in local trade. Tubers of fairy potatoes (Claytonia lanceolata) were also locally eaten in the past. Species of Calandrinia, Cistanthe grandiflora, Claytonia and Lewisia are grown as garden ornamentals.

EUDICOTS

347. DIDIEREACEAE Octopus-tree family

Etymology: Montia is named in honour of Italian botanist Giuseppe Monti (1682–1760), who was professor of botany and prefect of the botanical garden of Bologna.

These usually unisexual (gynodioecious in Decarya, bisexual in Portulacaria) xerophytic trees and shrubs, sometimes vines, have pronounced long and short shoots. Stems are succulent when young and become woody with age, the short shoots often with spines. The alternate (opposite in Portulacaria and Ceraria) simple leaves are fleshy and entire, usually sessile or indistinctly petiolate and without stipules. Inflorescences are thyrses composed of dichasia, panicles, cymes or fascicles. The unisexual or bisexual flowers are usually subtended by two bracts that function as a calyx and can be showy. Tepals are four in two whorls, or five free persistent sepals and four to seven petals fused into a

Alluaudia procera, flowering stem, Irvine, California, USA [347]

Alluaudia procera, The Living Desert, Palm Desert, California, USA [347]

Survivors: Lewisia, often grown as a rockgarden ornamental, can survive submergence in boiling water, and has no problem with prolonged droughts of up to two years.

tube. The usually eight (sometimes four to ten or up to 60) stamens are generally basally fused to a ring-shaped nectar disc surrounding a pistil in male flowers and an ovary in female f lowers, which have staminodia instead of stamens, or (in Portulacaria) the stamens are fused to the corolla. Anthers of fertile stamens are dorsifixed and open by longitudinal slits. The superior ovaries are composed of usually three (sometimes two, four or six) carpels topped by a single style and a (strongly to scarcely) lobed stigma. The fruit is a dry indehiscent capsule enclosed in the pair of bracts, a thin-walled nut (Portulacaria), samara (Ceraria) or circumcissile capsule splitting open upwards into six valves (Calyptrotheca). Distribution: Didiereaceae can also be found in East Africa, but they are a dominant feature of the spiny forests of southern and southwestern Madagascar. Phylogeny and evolution: Many characters of Didiereaceae are shared with Cactaceae and Portulacaceae. In fact a number of genera formerly placed in Portulacaceae (Calyptrotheca, Ceraria and Portulacaria) are currently Didierea trollii, Adelaide Botanic Garden, South Australia [347]

Portulacaria afra, Prince Albert, Western Cape, South Africa (CD) [347]

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Ullucus tuberosus, Peru (SK) [348]

placed here, expanding the range of the family into East Africa; although they are no longer thought to be endemic to Madagascar, they remain a dominant dry-forest element.

Anredera cordifolia, Jardin des Plantes, Paris, France [348]

348. BASELLACEAE Malabar-spinach family

Genera and species: This family includes seven genera and 22 species: Alluaudia (6), Alluaudiopsis (2), Calyptrotheca (2), Ceraria (7), Decarya (1), Didierea (2) and Portulacaria (2). Uses: Stems of Didiereaceae are used for firewood by local people, endangering the populations of these slowly growing trees. Some species, especially Alluaudia trollii and Portulacaria afra, are grown as ornamentals. Etymology: Didierea is named for French botanist Alfred Grandidier (1836–1921), who is well known for his botanical explorations in East Africa, Zanzibar and Madagascar. His studies on Madagascar and his epic work on the natural and political history of the island drew the attention of the Franch government, which annexed Madagascar in 1890 as a result.

These are perennial vines and herbs that usually have thickened roots or tubers. Stems and leaves are succulent and mucilaginous. Leaves are alternate or nearly opposite at the base of the plant; they lack stipules and are often petiolate. Inflorescences are bracteate, axillary or terminal spikes, racemes or panicles that are usually indeterminant. The usually bisexual, actinomorphic flowers are subtended by one or two bracteoles. The two sepals are free or basally fused and coloured. The usually five (sometimes four or up to 13) petals are fused for half their length. The usually five (or four to nine) stamens

Basella alba, Helsinki Botanical Garden, Finland [348]

are fused with the petals, sometimes only at the base, sometimes almost completely so, or the filaments are broadened at the base and sometimes form a disc or tube. Anthers are basifixed or metafixed and open by longitudinal slits or pores. An annular nectary is usually present. The superior, unilocular ovary is sessile or shortly stalked and composed of three fused carpels, each topped with a style, which are basally fused or all fused into one. The stigma is slender, bifid to clavate or capitate to trilobed. The fruit is a thin-walled nutlet surrounded by the dry or fleshy perianth, and it is sometimes winged. Distribution: This family is distributed in tropical America and Africa, from Texas and Florida, throughout the West Indies and Central America to South America and the Galápagos Islands, and in Sub-Saharan Africa and Madagascar. The family has naturalised in Eurasia and Australia. Phylogeny and evolution: Basellaceae were formerly placed among Chenopodiaceae or Portulacaceae. However, the family was Plants of the World

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described in the 19th century and has since been supported by many studies. Morphological and molecular phylogenetic analyses placed Basellaceae close to Didiereaceae. Pollen morphology is similar to that of Didiereaceae and Portulacaceae.

connective. Female flowers lack a perianth, and the superior ovary is unilocular and topped with a style and three stigmas. The fruit is a thin-walled nutlet, several of which become embedded in a hardened inflorescence axis.

Genera and species: Basellaceae include four genera with 19 species: Anredera (12), Basella (5), Tournonia (1) and Ullucus (1).

Distribution: This family is endemic to Patagonia where it grows in salt marsh.

Uses: The tubers of ulluco (Ullucus tuberosus) were an important crop c. 4,000 years ago in pre-Columbian Peru and Ecuador, and they are still a significant source of protein and carbohydrates there. Similarly tubers or rhizomes of Anredera species are starchy and also eaten, and A. cordifolia (Madeira vine) is grown as an ornamental. Malabar spinach (Basella alba), native to Africa and Asia, is widely cultivated in tropical and subtropical areas as a vegetable. A red pigment from B. alba is used as food colouring in tropical Asia. Etymology: Basella is derived from vasala or basella-kira, Malabar names for this vegetable.

Phylogeny and evolution: Initially described as a species of Tetragonia (Aizoaceae) and later placed in Chenopodiaceae, the sole genus Halophytum is markedly different from Aizoaceae and Amaranthaceae in characters of the ovary, pollen and vascular anatomy. Molecular placement of this family is in a clade with Cactaceae, Portulacaceae and Basellaceae, and indeed this family shares pollen characters with the last. Genera and species: The family only has a single species: Halophytum ameghinoi. Etymology: Halophytum is composed of the Greek words άλας (halas), salt, and φυτών ( fyton), a plant.

350. TALINACEAE Fameflower family

This family of annual and perennial herbs (sometimes somewhat woody) often have tuberous roots. Leaves are alternate (rarely appearing opposite) and lack axillary hairs, scales or bristles. Blades are flat, succulent and sessile with entire margins. The inflorescence is an axillary or terminal panicle or thyrse, and rarely are the flowers solitary. The five sepals are free or slightly basally fused and fall off soon after the flower has opened. The five (to ten) petals are free and brightly coloured. Stamens are five to 30 and fused to the base of the petals. Anthers are basifixed and open by longitudinal slits. The superior ovary is topped with a (bi- or) tri-lobed stigma. Fruits

349. HALOPHYTACEAE Verdolaga family

Halophytum ameghinoi, La Rioja, Argentina (DA) [349]

Halophytaceae are succulent bisexual, branched, annual herbs. Their alternate (rarely appearing opposite or fascicled) leaves are sessile, lack stipules and are succulent and flattened on the top, rounded below. Inflorescences are unisexual, the males in dense bracteate spikes, the female flowers clustered in the axils of the upper leaves. Male flowers have four thin tepals and four stamens with long, exserted filaments. The dorsifixed anthers open by pores due to contraction of the

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Talinum paniculatum, private garden, Kingston upon Thames, Surrey, UK [350]

CARYOPHYLLALES

EUDICOTS

Portulaca oleracea, Yucatán, Mexico [351]

Portulaca pilosa in fruit (WA) [351]

are thin-walled capsules opening by three valves or nutlets.

351. PORTULACACEAE Purslane family

Distribution: This family is naturally distributed in tropical and subtropical America, Africa and Madagascar, but some species have escaped from cultivation elsewhere. Phylogeny and evolution: This family was previously included in Portulacaceae. Talinella is nested in Talinum. Talinopsis, previously placed with Talinum, belongs to Anacampserotaceae. Genera and species: Talinaceae consist of two genera with 28 species: Amphipetalum paraguayense and Talinum (27). Uses: Several species have edible leaves and are eaten like spinach. Waterleaf, Talinum fruticosum, and T. triangulare are both commonly used as a vegetable. Fameflower (T. paniculatum) is grown as an ornamental plant. Etymology: No derivation is given in the original publication, but Talinum may have been derived from the Senegalese name tali, traditionally used for Erythrophleum guineense (Fabaceae). However, the origin of Talinum is obscure and may alternatively have been derived from Ancient Greek ταλεία (taleia), blooming luxuriantly.

These are succulent annual and perennial herbs often with tuberous roots and usually with creeping or erect stems. Nodes and leaf axils have scales, bristles or hairs. Alternate or opposite leaves are simple, sessile and without stipules, and blades are succulent, flat or terete and entire. Inflorescences are terminal heads of solitary or clustered flowers subtended by a cluster (involucre) of leaves. Sepals are fused into a tube and persistent. Petals are four or five to seven and free or fused at the base only, the four to 100 stamens fused to the petals. Anthers are basifixed and open by longitudinal slits. The partly inferior to inferior ovary is unilocular at the apex and is topped by a short style and three to eight stigmas. The fruit is a thin-walled, circumscissile capsule with many seeds. Distribution: Portulacaceae can be found nearly worldwide but are most diverse in

Portulaca suffrutescens, New Mexico, USA

(DZ) [351]

the Neotropics. Their distribution in the more temperate areas is mostly due to naturalisation, although in some cases the origins of these plants are unknown. Phylogeny and evolution: Species in this clade have made several switches to C 4 photosynthesis. Previously, a larger number of genera were included in this family, but these are now placed elsewhere, mainly in Anacampserotaceae, Didiereaceae, Montiaceae and Talinaceae, leaving only Portulaca in this family. Portulaca is divided into two subgenera, one (subgenus Portulacella) restricted to Australia and the other more widespread. Genera and species: The sole genus in this family is Portulaca with c. 115 species. Uses: Purslane (Portulaca oleracea and several closely related species) can be eaten fresh or cooked and has a slightly acid flavour and a mushroom-like texture. It has been widely cultivated and is naturalised around the world. Portulaca grandiflora produces flowers of many colours and is often grown as a bedding plant. Etymology: Portulaca is the Latin name for the vegetable purslane, P. oleracea. The name was already used by Pliny and means ‘carrier of milk’. Plants of the World

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352. ANACAMPSEROTACEAE Love-plant family

EUDICOTS

fruit is a capsule that splits into three, five or six valves.

353. CACTACEAE Cactus family

Distribution: This is a family with a scattered distribution, in northern Mexico, southwestern USA, Bolivia, southern Africa, Somalia, Djibouti, Yemen and South Australia.

This is a family of perennial, succulent, rosette-forming herbs, some with woody bases and fleshy stems with short internodes. The opposite leaves have a tuft of axillary hairs or bristles and entire succulent blades. Flowers are in monochasia or dichasia or solitary in the leaf axils and are often surrounded by dry bracts. Sepals are fused and usually fall off soon after opening of the flower. The five petals are free and white or brightly coloured. There are five to numerous, basally fused stamens with basifixed anthers that open by longitudinal slits. The superior ovary is usually composed of three fused carpels with a single style and a capitate stigma. The Anacampseros sp., Eastern Cape, South Africa (JA) [352]

Phylogeny and evolution: Grahamia exhibits the putatively plesiomorphic characters of this family. They were previously included in Portulacaceae, but molecular studies have shown that they are sister to Cactaceae.

Etymology: Anacampseros (ανακάμψερος) is an ancient Greek name for a herb able to bring back lost love simply by touching it. It is composed of ανα (ana), return, καμπτο (kampto), bent, and έρως (eros), love.

These perennial evergreen trees, shrubs and vines can be diverse in shape or form. Some are leafy, but more frequently they are simple or branched leafless stem-succulents. These plants have stems that are columnar or globular and are solitary or in clumps, erect to scrambling or climbing. They are terrestrial, epipetric or epiphytic and are in the last case often pendent. Their roots are diffuse, sometimes with taproots or tubers. Stems remain green or become brown and woody with age and can be unsegmented or segmented, and segments often fall and make roots. Shoots or segments are spherical or clubshaped to cylindrical, sometimes flattened into

Avonia papyracea, Wolwekraal Nature Reserve, Western Cape, South Africa (CD) [352]

Jasminocereus thouarsii, Galápagos Islands [353]

Genera and species: A family of four genera and 27 species: Anacampseros (14), Avonia (11), Grahamia (1) and Talinopsis (1). Uses: Anacampseros telephiastrum is sometimes grown as an ornamental.

Hatiora salicornioides, Royal Botanic Gardens, Kew, UK [353]

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Echinocereus viereckii, Royal Botanic Gardens, Kew, UK [353]

Astrophytum ornatum, Ruissalo Botanical Garden, Turku, Finland [353]

cladodes. They can be smooth or tuberculate, the tubercles distinct and nipple-shaped or ridge-like protuberances or fused into vertical ribs. These ridges or ribs, when present, two to 30 or more; short shoots (called ‘areoles’) are positioned on the crests of these ribs or tubercles, usually bearing persistent spines and often also minute, barbed, deciduous spines (called ‘glochids’). The spirally arranged, alternate leaves are deciduous to persistent, vestigial or absent; when present they are usually sessile, but can be petiolate in some leafy genera such as Pereskia. They are flat or terete and lack stipules. Spines can be flexible and hair- or bristle-like to rigid and needle- or nail-like, terete to angled or flat. The bisexual (rarely unisexual), actinomorphic or sometimes zygomorphic flowers are formed one (to several) per areole, arranged in some leafy genera into panicles or cymes. The receptacle of the flower encloses and protrudes beyond the ovary, forming a floral tube (hypanthium). The five to 50 (or more) tepals intergrade gradually from bract-like or sepal-like outer tepals to petal-like inner tepals. Stamens are usually numerous (more than 50), fused with the inner surface of the flower tube. Anthers are bilocular and open by longitudinal slits. The inferior (or halfinferior) ovary can be covered with tubercles, areoles, spines and glochids and often has scales that are derived from tepals. The ovary is composed of up to 20 fused carpels forming a unilocular pistil with a single style and with a nectary around the base of the style which

carries a lobed stigma (each lobe representing a carpel). Fruits are usually succulent manyseeded berries, but some can split. Distribution: The family is widespread across the New World, from Canada throughout the USA and Mexico, Central America, the West Indies, the Galápagos Islands and South America. A single species, Rhipsalis baccifera occurs in tropical Africa, Madagascar, the Mascarenes, the Seychelles and Sri Lanka. Cactaceae are widely naturalised in other parts of the Old World tropics and subtropics. Phylogeny and evolution: Cactaceae are most closely related to parts of polyphyletic Portulacaceae s.l., and the latter have been dismembered to prevent the two from being merged. They share several characters, such as axillary hairs and indefinite numbers of stamens in bunches, that unite the group. The family might perhaps better be expanded to include the similar and closely related Anacampserotaceae, Portulacaceae and Talinaceae, although the botanical community appears to have little appetite for an expansion of this charismatic family. Traditionally, Cactaceae have been subdivided into four subfamilies, but at least Pereskioideae have been found to be polyphyletic, the genus Rhodocactus alone being sister to all other Cactaceae. Radiation of Cactaceae took place predominantly during the Miocene and Pliocene, and was most significant c. 8.5 million years ago. Generic

Pereskia weberiana, Royal Botanic Gardens, Kew, UK [353]

delimitation is not always clear, especially as hybridisation is common in some groups. Their generic taxonomy is inflated due to familiarity in horticulture resulting in an emphasis on only a few minor characteristics. Genera and species: This is a family of c. 94 genera and approximately 1,150 species (although more genera and species are listed by CITES): Acanthocereus (1), Ariocarpus (6), Armatocereus (6), Arrojadoa (4), Arthrocereus (4), Astrophytum (6), Austrocactus (3), Aztekium (2), Bergerocactus (1), Blossfeldia (1), Brachycereus (1), Brasilicereus (2), Browningia (8), Calymmanthium (1), Carnegiea (1), Cephalocereus (3), Cereus (20), Cipocereus (5), Cleistocactus (32), Coleocephalocereus (6), Copiapoa (19), Corryocactus (5), Coryphantha (48), Denmoza (1), Disocactus (10), Disocactus (11), Echinocactus (5), Echinocereus (c. 55), Echinopsis (c. 65), Epiphyllum (12), Epithelantha (2), Eriosyce (30), Escontria (1), Espostoa (9), Espostoopsis (1), Eulychnia (4), Facheiroa (3), Ferocactus (25), Frailea (9), Gymnocalycium (43), Haageocereus (19), Harrisia (7), Hatiora (5), Hylocereus (13), Jasminocereus (1), Leocereus (1), Lepismium (15), Leptocereus (4), Leuchtenbergia (1), Lophophora (3), Maihuenia (2), Mammillaria (135), Mammilloydia (1), Melocactus (33), Micranthocereus (9), Mila (1), Myrtillocactus (4), Neobuxbaumia (8), Neolloydia (2), Neoraimondia (2), Neowerdermannia (2), Obregonia (1), Opuntia (c. 110), Oreocereus (6), Ortegocactus (1), Pachycereus (12), Plants of the World

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Uses: The berries of many species are edible, although some are covered in spines that have to be removed before comsumption. Economically, the most important is Opuntia ficus-indica, of which the fruits (prickly pear or cactus fruit) are commonly found in fruit stalls around the world. Dragon fruit or pitaya (Hylocereus undatus), although of Mexican origin, is now mostly cultivated in Southeast Asia and exported to European and North American fruit stalls. It is used as an offering in Chinese temples and has a sweet but bland taste. Barbados gooseberry (Pereskia acutifolia) is cultivated for its edible fruits and leaves. The flat stems of

nopal (spineless forms of O. ficus-indica and other species) are considered a delicacy in Mexico and are eaten cooked or pickled as a vegetable (nopalitos). Similarly, the despined young stems of cadushi (Cereus repandus) are eaten in Curaçao. The pulp of the stem of some Echinocactus species is used in confectionery. Cochineal is produced from dried mealy bugs that live on Opuntia cochenillifera; they are used to make carmine, a scarlet dye that is now synthetically produced. The jumping cholla (Opuntia fulgida) is known in the southwestern USA and northern Mexico for its hard to remove stems (which the plant uses for vegetative reproduction); it is used to produce cholla gum, which has similar applications as gum arabic (Senegalia, Fabaceae). Some species of Cactaceae contain mescaline, which has hallucinogenic properties; these include Echinopsis pachanoi and especially Lophophora williamsii (peyote), the use of which can lead to colourful visions and a feeling of weightlessness. Many cactus species are cultivated as ornamentals, some specialists growing nothing else. Some species are popular houseplants; especially common are Christmas and Easter cacti

Etymology: Cactus is the Latinised form of the Greek κάκτος (kaktos), a name originally used for the cardoon (Cynara cardunculus, Asteraceae), a vegetable commonly eaten in the eastern Mediterranean. Linnaeus placed all known cacti in one genus, Cactus, complicating the nomenclature of Cactaceae because he even included a leafless orchid (Cactus parasiticus = Dendrophylax funalis, Orchidaceae). For nomenclatural stability, the generic name Cactus was rejected against Mammillaria and is now considered its synonym.

Opuntia bigelovii, Joshua Tree National Park, California, USA [353]

Opuntia littoralis, Santa Barbara, California, USA [353]

Lophophora williamsii (HB) [353]

Parodia (c. 50), Pediocactus (6), Pelecyphora (2), Peniocereus (15), Pereskia (17), Pereskiopsis (6), Polaskia (2), Pseudoacanthocereus (2), Pseudorhipsalis (6), Pterocactus (9), Quiabentia (2), Rebutia (26), Rhipsalis (35), Rhodocactus (1), Samaipaticereus (1), Schlumbergera (6), Sclerocactus (17), Selenicereus (11), Stenocactus (7), Stenocereus (22), Stephanocereus (2), Stetsonia (1), Strombocactus (1), Tacinga (6), Thelocactus (11), Uebelmannia (3), Weberbauerocereus (5) and Weberocereus (8).

Mammillaria varieaculata, Royal Horticultural Society Garden, Wisley, UK [353]

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(cultivars of Schlumbergera ×buckleyi), and ‘epiphyllums’ (Disocactus ×hybridus). Cactaceae are valuable garden plants for arid regions around the world, although some naturalise easily and can cause problems in nature conservation. International trade is regulated by CITES. Turks Island of Turks and Caicos received its name because of the abundant Melocactus intortus stands there that resemble Turk’s caps. Becoming 20 m tall, the saguaro (Carnegiea gigantea) is the largest member of the family. One of the smallest is Blossfeldia liliputana, a resurrection plant that is only a few millimetres across.

Cleistocactus winteri, Royal Botanic Gardens, Kew, UK [353]

CORNALES

EUDICOTS

CORNALES Families 354 to 360 are members of Cornales, an order that is estimated to have diverged c. 97–101 million years ago. The oldest fossils are from the Palaeogene and Late Cretaceous of Europe, where the Mastixioidean Flora originated. Fossils of this flora are also known from later dates in North America, but are absent from Asia. Fossil Alangium, Cornus, Davidia, Mastixia and Nyssa are all known in deposits from the Late Cretaceous onwards. The broadly circumscribed Cornaceae of some authors were found to be polyphyletic.

Inflorescences are axillary or produced on short lateral shoots, and they are pedunculate heads, racemes, panicles or spikes, sometimes subtended by one or two colourful bracts (Davidia). Flowers are actinomorphic and bisexual or unisexual. The five minute sepals are sometimes reduced to a rim or are absent. The five to ten petals are free or (in female flowers) fused to the ovary (Nyssa), or a perianth is absent altogether. Stamens are usually ten, sometimes up to 26 or as few as eight, in two (rarely three) whorls. Filaments are elongate, inserted around the disk or the style, and the basifixed anthers are often small, opening by lengthwise slits. The inferior ovary is composed of one or two carpels (six to ten in Davidia), absent in male flowers. They form a single locule topped

with a style that is immersed in the nectar disk and is simple or has two or three (or six to ten in Davidia) branches with a reflexed, revolute, spreading or punctiform stigma. Fruits are usually drupes, rarely samaras (Camptotheca).

Diplopanax stachyanthus, Siu Chai, Vietnam (CD) [354]

Nyssa sylvatica in autumn colour, Madison, Wisconsin, USA [354]

Nyssa sinensis, Fletcher Moss, Manchester, UK

Camptotheca acuminata in bud, Dali, Yunnan, China [354]

Davidia involucrata, Sarah Duke Botanical Garden, North Carolina, USA [354]

354. NYSSACEAE Tupelo-tree family

These are evergreen and deciduous trees with alternate or falsely opposite or pseudowhorled, petiolate leaves without stipules. Blades are pinnately veined (sometimes with several main veins branching near the base of the leaf) and have entire or toothed margins.

Distribution: This is a bicontinental family found in northeastern North and Central America and in southern India, Sri Lanka, East Asia and tropical Indomalesia. Phylogeny and evolution: Often included in a broadly circumscribed Cornaceae, Nyssaceae have been shown by molecular studies to include Camptothecaceae, Davidiaceae and Mastixiaceae, which are sister to a clade comprising Hydrostachyaceae, Loasaceae

[354]

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CORNALES and Hydrangeaceae; they are therefore widely separated from Cornaceae sensu stricto within Cornales. Probably originating in Europe in the Upper Cretaceous, where the family is now absent, the earliest representatives appear similar to extant Mastixia (called the Mastixioidean Flora). Tertiary fossils of this lineage are common in the Northern Hemisphere, Mastixia being abundant in Europe and Siberia c. 65–70 million years ago. Nyssaceae were also abundant in the Pliocene of North America, and fossil seeds are known that mix characters of modern Mastixia and Nyssa. Davidia (now restricted to East Asia) is known from Palaeocene fossils in North America and eastern Russia. It may be that these plants dispersed across the Bering Strait during the Late Cretaceous or Early Tertiary. The oldest Nyssa fossils are known from the Eocene of Europe, where the genus disappeared during the Pliocene, but Nyssa had migrated before that to North America, from where during the Miocene it made the long leap to East Asia, where it radiated. The placement of Davidia has been problematic in the past, but it is now clear that it belongs here. Diplopanax was previously placed in Araliaceae, but it is in fact a ‘living fossil’, identical to some Tertiary mastixioid fossils. Genera and species: Nyssaceae include five genera and c. 33 species: Camptotheca (2), Davidia (1), Diplopanax (2), Mastixia (c. 20) and Nyssa (c. 8). Uses: The fruits of ogeechee lime (Nyssa ogeche) from the southeastern USA are locally used for preserves. Camptotheca has compounds (especially camptothecin) used in the treatment of colon and ovarian cancers. Mastixia is used for plywood in India, and Nyssa wood is occasionally harvested as well, especially tupelo wood from black gum (Nyssa sylvatica), which is used for veneers etc. Happy tree (Camptotheca acuminata), handkerchief or dove tree (Davidia involucrata) and black gum are commonly cultivated ornamentals in the (warm-) temperate zones of the Northern Hemisphere.

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Etymology: Nyssa (Νήσσα) is a water nymph of Greek mythology who nursed the god Dionysos back to health. The name was probably given to this tree because it grows in water. In modern greek νήσσα (nissa) means ‘duck’.

355. HYDROSTACHYACEAE Waterspike family

Hydrostachyaceae are unisexual, highly modified, submerged, aquatic perennial herbs with a short disc-like stem that attaches to rocks in rapidly flowing water. Roots and the vascular system are highly reduced, and stomata are absent; all parts are covered in scale-like or fringe-like structures. The seaweed-like leaves are alternate, entire to two- or three-times pinnatifid, and the leaf base is enlarged and fused into intrapetiolar Hydrostachys imbricata, Amboniriana, Madagascar (CD) [355]

stipules. Inflorescences are dense, bracteate spikes with flowers in the axils. Petals and sepals are absent. Male flowers are interpreted as a single stamen, deeply divided into two thecae sessile on the bract (or composed of two fused stamens). Female flowers consist of a sessile superior ovary with two carpels forming a single locule and topped by two filiform styles. Fruits are septicidal capsules with numerous tiny seeds, which become mucilaginous when in contact with water and germinate when the slimy seeds have attached themselves to rocks. The plants only flower when water levels drop sufficiently during dry seasons. Vegetative reproduction through sprouts from adventitious roots is common. Distribution: This family is restricted to tropical Central and southern Africa and Madagascar. They resemble Podostemaceae in their aquatic habit, vegetatively resembling an alga, but they differ clearly from that family in the spike-like inflorescences. Phylogeny and evolution: Because of the aquatic nature of this family, the vegetative and floral characters have been strongly reduced, making placement among other Hydrostachys imbricata, Amboniriana, Madagascar (CD) [355]

CORNALES

EUDICOTS

Philadelphus coronarius, Transylvania, Romania [356]

Hydrangea hydrangeoides, Helsinki Botanical Garden, Finland [356]

Hydrangea febrifuga, San Francisco Botanical Garden, California, USA [356]

vascular plant families on the basis of morphology difficult. They were long associated with the superficially similar Podostemaceae, although vegetative and floral structures are markedly different. Molecular studies at first indicated that Hydrostachyaceae are most closely related to Hydrangeaceae in Cornales, although long-branch attraction was thought perhaps to be an issue in its specific placement within the order, and they are now thought to be sister to Loasaceae plus Hydrangeaceae.

climbers has opposite leaves that are joined by a line across the stem; leaves are rarely whorled or alternate. The stem line is formed by the sheathing petiole bases, and true stipules are absent. Blades have entire or toothed, seldom lobed margins, and venation is pinnate, sometimes pseudo-palmate at the base. Inflorescences are terminal cymes, sometimes corymbs, thyrses or panicles. Flowers are bisexual or unisexual and actinomorphic or somewhat zygomorphic in sterile flowers. The four, five or up to 12 sepals are free or basally fused, enlarged and petal-like in some species with sterile flowers. The four to 12 petals are basally or completely fused to form a hood (calyptra). The four to numerous stamens (usually twice as many as petals) are free or basally fused with f lat or round filaments, sometimes forked at the tip, sometimes the connective protruding beyond the anthers, which are basifixed and open by lengthwise slits. The (half-)inferior ovary is composed of two to 12 fused carpels, topped with a single style that is sometimes branched at the top or by free styles that are as many as there are carpels. Stigmas are papillate or smooth. Fruits are capsules or berries.

and northern South America, Southeast Asia, Malesia, Polynesia and Hawaii.

Genera and species: The sole genus in this family is Hydrostachys with 22 species. Etymology: Hydrostachys is derived from the Greek ύδωρ (hydor), water, and στάχυς (stachys), a spike.

356. HYDRANGEACEAE Hortensia family

This family of evergreen and deciduous, robust herbs, shrubs, small trees and woody

Distribution: This family is most diverse in the warmer temperate regions of North America, the Alps and the Caucasus in Europe and East Asia, with a few species in the tropics, mostly in montane regions of Central

Phylogeny and evolution: Hydrangea and allied genera were previously placed in Saxifragaceae, based on the two free carpels of some species, but are unrelated to that family. Recent molecular studies placed them in Cornales, close to Loasaceae. Hydrangeaceae are well represented in the Tertiary flora of mesic forests in the Northern Hemisphere, when they were more widespread than their current distribution, having died out in large areas during the ice ages. They probably evolved in the Late Cretaceous, from when there are several putative Hydrangeaceae fossils known. An estimated age for crown Hydrangeaceae may be 69–83 million years. In the latest phylogenetic studies, Hydrangea is polyphyletic and has been expanded to include genera such as Broussaisia, Decumaria, Dichroa, Pileostegia, Platycrater and Schizophragma, which all share many characters with Hydrangea. Genera and species: This is a family with 10 genera and c. 223 species in two subfamilies: Hydrangeoideae – Carpenteria (1), Deutzia (c. 60), Fendlerella (4), Hydrangea (c. 66), Kirengeshoma (2), Philadelphus (c. 80) and Whipplea (1). Jamesioideae – Fendlera (5) and Jamesia (2).

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Carpenteria californica, Santa Barbara Botanical Garden, California, USA [356] Kirengeshoma palmata, Royal Botanic Gardens, Kew, UK [356]

Deutzia gracilis ‘Nikko’, Royal Botanic Gardens, Kew, UK [356]

Hydrangea anomala, Mt Tianmu, Anhui, China [356]

Hydrangea macrophylla subsp. macrophylla, private garden, Kingston upon Thames, Surrey, UK [356]

Uses: Changshan (Hydrangea febrifuga) has been used as an antimalarial drug in East Asia for centuries. Steamed leaves of Hydrangea macrophylla subsp. serrata are used in a sweet drink called ‘ama-cha’ in Japan and Korea, where it is mainly used during a Buddhist ceremony in April. Hydrangeaceae are mildly poisonous, and care has to be taken when consuming them. The wood of Hydrangea paniculata is sometimes used to manufacture umbrella handles. Flowers of various mock oranges (Philadelphus coronarius, P. mexicanus, P. microphyllus etc.) were used for their jasmine-like scent in the perfume industry. Many species are valued garden ornamentals. A pot of flowering hortensia (Hydrangea macrophylla subsp. macrophylla cultivars) is traditionally given on Mother’s Day in some European countries. This species has been domesticated and cultivated in Japan for millennia and was not known to European gardeners before the mid-19th century, being popularised in the Netherlands and Russia 474

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simultaneously. Other popular garden hortensias are the Asian Hydrangea anomala subsp. petiolaris (climbing hortensia), H. aspera (rough hortensia), H. involucrata, H. heteromalla, H. paniculata (plume hortensia) and H. macrophylla subsp. serrata (mountain hortensia) and the American H. arborescens (smooth hydrangea) and H. quercifolia (oakleaf hydrangea). Apart from hortensia, several other Hydrangeaceae are frequently grown in gardens, including Carpenteria californica (bush anemone), Hydrangea barbara (woodvamp), Deinanthe coerulea (false hydrangea), Deutzia species and cultivars, Hydrangea, Kirengeshoma palmata (yellow waxbell), Philadelphus coronaria and other species of mock orange and Hydrangea hydrangeoides (climbing hydrangea). Etymology: Hydrangea is derived from the Greek ύδωρ (hydor), water, and αγγείο (angeio), a vase or vessel, referring to the seed capsules.

Fendlera rupicola var. wrightii, New Mexico, USA (DZ) [356]

357. LOASACEAE Blazingstar family

Loasaceae are a family of annual and perennial herbs, climbers, woody vines, shrubs and small trees to 10 m tall. Plants are often covered with stinging hairs and rough or glochid-like trichomes. Glandular hairs are also sometimes present. Leaves are opposite or sometimes alternate, in a basal rosette or distributed along the stem, with or without a petiole and with or without false stipules. Blades are simple and lobed or palmately or pinnately compound, pinnatisect, pinnatifid, bipinnatifid or trifoliate, the margins toothed

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with hydathode teeth or mucronate, rarely entire, and venation is pinnate, often with several main veins initiated from the base (palmate at base). Inflorescences are usually bracteate, terminal thyrses, rarely racemes or reduced to dichasia, or flowers are solitary. The bisexual f lowers are actinomorphic (usually) and occasionally cleistogamous. The four, five or up to eight sepals are fused into a conical to globose tube that is densely hairy (often stinging). Petals are as many as sepals and free and spreading or reflexed, linear, ovate or circular, spathe-like and boat-shaped or flat, and margins are entire, serrate or deeply incised. Petals are also often shortly clawed and have a tooth-like appendix on each side. Stamens are once or twice as many as petals or numerous with filaments that are inserted at the base of the petals, rarely the filaments petal-like or short.

Anthers are basifixed and open by lengthwise slits. Staminodia (sterile stamens) are common and highly modified, free or fused, filiform, petal-like or composed into complex nectar scales, often brightly coloured. Nectaries are separate glands or a ring- or cup-shaped nectar disk, sometimes absent. The inferior or nearly superior ovary is composed of one or three to seven fused carpels, each forming a locule or fused into a single locule. The style is filiform and topped with a single pointed or two- to five-lobed stigma. Fruits are straight or twisted capsules, opening along the sutures, or indehiscent capsule-like achenes of nuts. Distribution: A predominantly American family, found from the Great Lakes of North America south to Patagonia. They are especially diverse in tropical America at higher elevations. There are disjunct

populations on the Galápagos and in southwestern and northeastern Africa, Yemen and Polynesia (Marquesas Islands). Phylogeny and evolution: Affinities of Loasaceae have long been controversial. They were often associated with Passifloraceae or even placed in Turneraceae. Phytochemistry indicated that Loasaceae may be related to Cornales, and molecular studies have placed Loasaceae as sister to Hydrangeaceae. The family shares many characters with Jamesioideae of Hydrangeaceae, and thus this placement is not unreasonable. Even though four subfamilies have traditionally been recognised, not all are monophyletic, and five clades have been observed in the most recent phylogenetic studies. An age estimate for Loasaceae varies, from 63–67 to 31–33 million years old. The association between Plakothira

Author of the Green Bible - Arthur Cronquist (1919–1992) During the 1950s, American botanist Arthur Cronquist questioned the usefulness of the system laid out by Adolf Engler and Karl Prantl, which was the standard at the time. Originally a specialist on Asteraceae, he was flirting with the idea of creating a new taxonomic system. Several contemporary botanists had the same goal in mind; Rolf Dahlgren, Armen Takhtajan and Robert Thorne would each produce their own classification, but Cronquist’s became the most widely accepted. Cronquist became friends with Takhtajan, and they exchanged extensive information about each other’s schemes. A book on the evolution of flowering plants published in 1968 laid out what would become his system. His Integrated system of classification of flowering plants was published in 1981 and became the standard for family classification until the molecular revolution in the 1990s resulted in the consensus classification by the Angiosperm Phylogeny Group. His system was similar to that of Takhtajan, placing the flowering plants in two large classes, the dicots and monocots, but he opted for somewhat broader circumscriptions of orders and families. Apart from his classification system, he is also known for writing a manual to the plants of the northeastern USA, which was dubbed ’the green bible’. He passed away from heart failure while studying specimens of Mentzelia (Loasaceae), one of his favourite groups of plants, at Brigham Young University in Utah, one of his favourite herbaria. A species of that genus, M. cronquistii, was Arthur Cronquist (Copyright 1997, Hunt Institute) named in his honour.

Mentzelia multiflora, New Mexico, USA (DZ) [357]

Mentzelia aurea, Ruissalo Botanical Garden, Turku, Finland [357]

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Caiophora lateritia, Helsinki Botanical Garden, Finland [357]

Curtisia dentata in fruit, Kirstenbosch Botanical Garden, South Africa (CD) [358]

Curtisia dentata, South Africa (NH) [358]

Curtisia dentata, inflorescence, South Africa

(Polynesia), Klaprothia (South America) and Kissenia (Africa and Arabia) shows some intriguing biogeographical patterns that must involve dispersal over long distances.

although they are unlikely to become popular garden plants because of their stinging hairs.

Assegai family

Blades are pinnately veined, have a coarsely toothed margin and are covered below in rusty-coloured velvet hairs. Inflorescences are terminal thyrsoid panicles with many clusters of flowers and prophylls. The bisexual (rarely unisexual) flowers are actinomorphic. The four sepals are basally fused into a turbinate tube with triangular, hairy lobes, and the four petals are free and hairy. The four stamens alternate with the petals and have thin, free filaments, and the basifixed anthers are cordate at the base and open by lengthwise slits. A nectar disk tops the inferior ovary, which is composed of four carpels that are fused to form four locules, surrounding a short, conical style with a four-lobed stigma. Fruits are small tetralocular, four-seeded drupes that are crowned by the persistent calyx and style.

These evergreen trees have simple, opposite, petiolate leaves without stipules, but the petiole bases are fused across the stem.

Distribution: Curtisia dentata is found in forests and bushlands of southern and southeastern Africa.

Genera and species: This family has 20 genera and c. 308 species: Aosa (7), Blumenbachia (12), Caiophora (34), Cevallia (1), Chichicaste (1), Eucnide (13), Fuertesia (1), Gronovia (2), Huidobria (2), Kissenia (2), Klaprothia (2), Loasa (c. 36), Mentzelia (c. 80), Nasa (c. 100), Petalonyx (5), Plakothira (3), Presliophytum (3), Schismocarpus (1), Scyphanthus (2) and Xylopodia (1). Uses: Seeds of Mentzelia albicaulis are edible, and the plants have locally (in western North America) been smoked instead of tobacco. Because of the attractive flowers of Loasaceae, some species of Blumenbachia, Caiophora, Eucnide, Loasa and Mentzelia are occasionally cultivated as ornamentals, 476

Loasa tricolor, Ruissalo Botanical Garden, Turku, Finland [357]

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Etymology: The derivation of the genus Loasa is unknown, but it is probably from a native South American language.

358. CURTISIACEAE

(NH) [358]

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Phylogeny and evolution: Curtisia was previously included in Cornaceae, with which it shares pollen morphology and fruit characteristics. It was excluded from them because it lacks the peculiar vascular bundles that cross the septa in the ovaries of Cornaceae. Molecular analysis has shown that Curtisiaceae are probably most closely related to Grubbiaceae. The distinctive fruits of Curtisia are known from Eocene sediments in southern England. Genera and species: The only species in this family is Curtisia dentata. Uses: Assegai wood is a valued timber for furniture and other fine woodwork. Etymology: Curtisia is named for English botanist, nurseryman and entomologist William Curtis (1746–1799), who was prefect of the Chelsea Physic Garden and founder and editor of The Botanical Magazine (later Curtis’s Botanical Magazine), the longestrunning botanical journal.

359. GRUBBIACEAE Koolhout family

Grubbiaceae are deciduous and evergreen shrubs with opposite, simple, petiolate and nearly sessile leaves without stipules. Petiole bases are connected by a transverse ridge across the stem. Leaf blades are inrolled, leathery and linear, with pinnate (but obscure) venation. Inflorescences are axillary, sessile, two- or three-flowered conelike cymes subtended by bracts. Flowers are actinomorphic, bisexual and subtended by two bracteoles. Depending on the interpretation of these reduced flowers, the perianth is fourparted in one or two whorls, four sepals or two sepals and two petals; they are all hairy outside and pink or red inside. The eight stamens are in two whorls, with one opposite the perianth that is basally connected to it and has longer filaments. Anthers are dorsifixed, the filaments slightly exceeding the thecae and terminated by a blunt tip. They open by lengthwise slits below to form two valves. A hairy nectar disk surrounds the style. The inferior ovary is composed of two fused carpels that form one or two locules. The ovaries are sometimes fused with those of the other flowers in the triad. A single style tops the ovary and is terminated by a one- or two-lobed stigma. Fruits are drupes that are fused into a small cone-like structure.

Geissolomataceae, which are found in the same area but are unrelated. Early molecular studies placed Grubbia as an early branch of Ericales, but more recently it has been shown that Grubbiaceae are sister to Curtisiaceae in Cornales. Grubbia shares several general cornalean characters, but it does not share unique characters with Curtisiaceae other than perhaps fused leaf-bases. The two families could potentially be merged but are maintained separate pending further study. Ophira and Strobilocarpus are now treated as part of Grubbia. Genera and species: The sole genus in this family is Grubbia with three species: G. rosmarinifolia, G. rourkei and G. tomentosa. Etymology: Grubbia honours Swedish botanist Michael Grubb (1728–1808), who collected plants in South Africa.

360. CORNACEAE Dogwood family

Distribution: This family is endemic to the Cape Province in South Africa. Phylogeny and evolution: Many affinities have been suggested in the past for Grubbiaceae, but their wood anatomy shares characters with those of Bruniaceae and Grubbia tomentosa, Kogelberg Nature Reserve, Cape Province, South Africa (CD) [359]

Grubbia rosmarinifolia, South Africa (NH) [359]

Deciduous and evergreen trees, shrubs, vines and rhizomatous, perennial herbs make up this family. They are glabrous or have T-shaped or stellate hairs and can be thorny or not. Opposite or less commonly alternate, petiolate leaves are without stipules. Blades have entire, serrate or lobed margins and pinnate or a basally palmate venation that curves along the leaf margin. Leaf veins of Cornus have xylem thickenings that can be pulled out when tearing the leaf without breaking connections between the veins. Inflorescences are cymes, heads, corymbs, umbels or panicles, sometimes with colourful (or green) involucral bracts with part or the entire inflorescence posing as a single flower or bracts absent. The actinomorphic

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Alangium ridleyi, Singapore (MN) [360]

Cornus kousa var. chinensis, Mt Huangshan, Anhui, China [360]

Alangium platanifolium, Royal Botanic Gardens, Kew, UK [360]

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Cornus mas, Gordes, France [360]

Cornus officinalis, East Bergholt Place Arboretum, UK [360]

Cornus suecica, Isokari, Finland [360]

flowers are bisexual, rarely unisexual, and plants then dioecious. The four, five, or up to ten sepals are free, sometimes no more than small teeth, sometimes margins toothed or lobed. Petals are sometimes absent, but usually four, five or up to ten, free or fused at the base. There are four, five or up to 40 stamens that surround a circular nectar disk on the ovary. Filaments are free, glabrous, villous or bearded on the inner side. Anthers are sagittate, linear and basifixed or dorsifixed, opening by lengthwise slits, opening explosively in Cornus canadensis. The inferior ovary is composed of two (rarely four) carpels that are fused to form one to four locules. Styles are free (two to four) or fused into a single style, topped with a dry, non-papillate, usually capitate stigma. Fruits are drupes or berries, often crowned with the sepals and disk.

Distribution: Cornaceae are mostly distributed across the North Temperate zone, extending south into the mountains of Central and South America, the mountains of East Africa, Madagascar and the Mascarenes and widespread across temperate and tropical South and East Asia, extending into eastern Australia and the Pacific islands.

reduced this family to the single genus Cornus and its close relative Alangium, which was in earlier classifications placed in its own family Alangiaceae. Cornus and Alangium fossils have been known since the Upper Cretaceous, and molecular clock studies have indicated an age of c. 80 million years, with diversification c. 66 million years ago. Cornus had more locules per fruit during the Palaeocene and Eocene than the two modern taxa. Alangium originated in North America from where it dispersed during the Eocene into East Asia, south to Africa, Madagascar and the Mascarenes, with its major diversification taking place in South and Southeast Asia during the Neogene. Herbaceous members of Cornus (C. canadensis and C. suecica) were previously placed in the genus Chamaepericlymenum, and up to seven additional genera are sometimes

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Phylogeny and evolution: Cornaceae have been variably circumscribed in the past, but were not found to be monophyletic in the traditional broader sense. They used to include genera such as Aucuba (Garryaceae), Curtisia (Curtisiaceae), Davidia, Mastixia and Nyssa (Nyssaceae), Griselinia (Griseliniaceae), Helwingia (Helwingiaceae), Melanophylla and Torricellia (both Torricelliaceae), Metteniusa (Metteniusaceae) and Kaliphora (Montiniaceae). Molecular evidence has

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proposed instead of the single-genus concept, but this makes an otherwise monophyletic genus unnecessarily complicated; subgeneric classification is more appropriate. Genera and species: Cornaceae in the strictest sense include two genera with c. 86 species: Alangium (c. 21) and Cornus (c. 65). Uses: Several species of Cornus have edible fruits. Cornelian cherry (C. mas) is the most common, especially in Eastern Europe, and its drupes are higher in vitamin C than oranges. It is used in jams, syrups and cordials and is the

basis of vin de cornouille, a French alcoholic drink. East Asian C. kousa, commonly grown as an ornamental, also has edible fruits, although they are mealy in texture. Fruits of Japanese cornel (C. officinalis) have somewhat better texture and flavour. The wood of Cornus is hard and finegrained. It is useful for making tool handles, arrows, skewers, walking canes, furniture inlays and other small items requiring hard and durable wood. Muskwood (Alangium villosum) has scented wood locally used in northeastern Australia. Several species (e.g. C. florida and C. officinalis) are used medicinally for their

high tannin content, and Alangium salviifolium is used as an ipecac substitute in India. Cornus florida has molluscicidal saponins. Several species are grown as ornamentals, especially Alangium platanifolium and many species and cultivars of Cornus. Cornus alba ‘Sibirica’ has red stems that are used in winter decorations and basketry. Etymology: Cornus, meaning ‘tough’ (referring to the wood), is the Latin name for cornelian cherry, C. mas. It is called ‘dogwood’ because the stiff branches of C. sanguinea were used as dags or skewers.

ERICALES Families 361 to 382 form the diverse order Ericales. The oldest fossils of Ericales are from 90-million-year-old rocks in North America, and a good diversity existed there during the Late Cretaceous. Diverse age estimates for diversification of Ericales have been suggested, varying from c. 118 to c. 85 million years. Ericales make up nearly 6% of the extant diversity of plants, the majority of this diversity in Ericaceae. They most notably occur in warm temperate regions. Some families (e.g. Ebenaceae, Lecythidaceae and Sapotaceae) can dominate the understorey of tropical rainforests. Relationships within Ericales are poorly understood, although several wellsupported clades can be recognised. Previous to molecular phylogenetics, several families were included in Theales, but that order included families that are now in Caryophyllales, Malpighiales and Malvales.

361. BALSAMINACEAE Balsam family

This is a family of annual and perennial herbs, rarely slightly woody, with succulent stems that usually root at the lower nodes and everywhere in creeping species. Roots occasionally make tubers, and stems may form rhizomes in some species. They are usually terrestrial, but aquatics and epiphytes are also known. Leaves are simple, usually alternate and spirally arranged, occasionally opposite or whorled, without stipules, but sometimes with glands at the base or on

the petiole. Blades are pinnately veined and have an entire to serrate margin, with the teeth often glandular. Inflorescences are axillary racemes, false umbels or fascicles or flowers are solitary in leaf axils, pedunclate or not. Flowers are bisexual, zygomorphic and usually upside-down (resupinate). The three (or five) free sepals are petal-like, the lower one enlarged, sac- or funnel-shaped and usually tapering to a nectar spur with a swollen or pointed tip, rarely bilobed or the spur missing. The five petals are all free (Hydrocera) or the lower four united into lateral pairs (Impatiens). The upper petal is flat or hooded, often with a crest. The five stamens alternate with the petals and are fused into a ring surrounding the ovary and falling off in a single unit. Filaments are flat with scale-like appendages inside; anthers are basifixed and open by a slit or pore. The superior ovary is composed of four or five fused carpels, each forming a locule, topped by a single short or absent style with one to five stigmas. Fruits are

an indehiscent, fleshy berry (Hydrocera) or a four- or five-valved, elastically opening fleshy capsule, with seeds dispersed explosively from the opening valves, often upon touch (as referred to in the generic name Impatiens and the English common name ‘touch-me-not’ for I. noli-tangere). Distribution: The family has a broad distribution, being found in temperate North America, temperate and subtropical Eurasia, Sub-Saharan Africa, Madagascar and the Seychelles. They have their greatest diversity in tropical Asia and Africa. Hydrocera is confined to India and Southeast Asia. Phylogeny and evolution: An interpretation of morphology suggested a relationship with Tropaeolaceae and Geraniaceae, but molecular and chemical studies placed this family in Ericales, which were redefined to accommodate a broader number of families. Within the order, they fall within a clade Plants of the World

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Impatiens balsamina, Aurangabad, India [361]

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Impatiens pseudoviola, private garden, Kingston upon Thames, Surrey, UK [361]

with Marcgraviaceae and Tetrameristaceae, both woody families. With Tetrameristaceae, they share the petal-like sepals and structure of the nectar glands, anthers and styles. Diversification in Balsaminaceae has been estimated to have started c. 30 million years ago, with diversification of Impatiens itself somewhat later, during the Early Miocene (c. 23 million years ago). Climate change in Southeast Asia has caused much fragmentation and sparked some of the diversification in this family c. 5 million years ago. Genera and species: Balsaminaceae are a family of two genera with c. 1,000 species: Hydrocera triflora and Impatiens (c. 1,000), Uses: The fruit of giant or Himalayan balsam (Impatiens glandulifera) is sometimes eaten. Originally from the Himalayas, it was introduced as a garden ornamental in North America and Europe, where it has become invasive, growing quickly and crowding out native species, especially along water courses and lake shores. Impatiens niamniamensis is occasionally grown as an ornamental pot plant, and its young shoots are eaten as a vegetable in its native Congo. Impatiens tinctoria has tubers that are the source of a red dye used in cosmetics. It has beautiful fragrant flowers and is a valuable ornamental. Other popular ornamental species are garden balsam (I. balsamina), formerly 480

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Impatiens hians, Helsinki Botanical Garden, Finland [361]

used in medicine, busy lizzy (I. walleriana and cultivars), which due to a mildew disease has now largely been replaced by New Guinea hybrids (I. hawkeri cultivars and hybrids with I. linearifolia or other species). Etymology: Balsamina is the Latin diminutive for balsam. It is a synonym of Impatiens, which is Latin for impatient, referring to the explosive capsules when touched.

362. MARCGRAVIACEAE Shingle-vine family

Marcgraviaceae are terrestrial, epiphytic and hemi-epiphytic vines and shrubs, rarely small trees. Leaves are simple, alternate and lack stipules, with blades that have entire margins (rarely minutely crenate) and pinnate venation. Inflorescences are upright or pendent terminal racemes, sometimes resembling umbels or spikes, with bracts that are transformed into leaf-, bladder-, spur-, sac-, tube-, boat-, pitcheror cup-shaped nectaries. Pedicels usually bear

Hydrocera triflora, Singapore (WA)

[361]

two sepal-like prophylls. Flowers are bisexual and actinomorphic. The four or five sepals are unequal in size and free or nearly so. The three to five petals are free or fused, united into a cap in Marcgravia. Stamens are three to numerous with free or basally fused filaments in one or two whorls. Anthers are basifixed and open on the inside with lengthwise slits. The superior ovary has two to 20 complete or incomplete locules, with a distinct style or stigmas sessile. Fruits are dehiscent berries or capsules. Distribution: An exclusively Neotropical family, Marcgraviaceae can be found from central Mexico and the Antilles to northern Argentina and southern Brazil. Phylogeny and evolution: Many characters of Marcgraviaceae (wood anatomy, pollen morphology) are shared with Theaceae and Pentaphylacaceae. They also share some characters with Tetrameristaceae, and both were believed to be closely related to Theaceae in the past. Molecular studies placed these families in a clade with herbaceous Balsaminaceae. Apart from distinctive Marcgravia (two-ranked leaves, tetramerous f lowers, calyptrate petals), delimitation of these genera may need to be revisited; Caracasia may have to be merged with Ruyschia. The peculiar inflorescence bracts attract

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EUDICOTS

Schwartzia costaricensis, Volcán Barva, Heredia, Costa Rica (CD) [362]

Marcgravia umbellata, Guadeloupe [362]

Marcgravia rubra, Dominican Republic (WJ) [362]

a great variety of pollinators, including hummingbirds and bats; Marcgravia evenia has a concave bract that bats use for echo location. The name shingle vine refers to the stems of some species that have leaves appressed to the trunk of the host tree. Leaves on flowering branches are normally oriented. Genera and species: This is a family of eight genera and c. 120 species: Caracasia (2), Marcgravia (c. 60), Marcgraviastrum (15), Norantea (2), Ruyschia (5), Sarcopera (10), Schwartzia (18) and Souroubea (19). Uses: Local tribes in Peru and Ecuador eat the fruit and drink the sap of some Marcgravia species called ‘purum hijos’. Schwartzia adamantium is said to indicate the presence of diamonds, but no research has confirmed this. Etymology: Marcgravia honours German naturalist and astronomer Georg Marggraf (also spelled Marcgrave or Markgraf, 1610–1644), who was known for his travels in Brazil.

363. TETRAMERISTACEAE Punah family

This is a family of terrestrial trees, shrubs and mangroves with fluted trunks. Leaves are simple, alternate, with or without a petiole, no stipules, entire leaf margins and pinnate venation. Pelliciera bears salt glands on the leaf blades. Inflorescences are axillary umbellike or corymb-like racemes or one to three flowers in the leaf axils, with two prophylls inserted directly below the sepals. Flowers are bisexual and actinomorphic. The four or five petal-like sepals are free, with numerous flaskshaped nectar glands at their base, appearing like pores or slits on the inner surface. The

Norantea guianensis, Serra de Maranguape, Ceará, Brazil [362]

four or five petals are free, and the four or five stamens alternate with the petals and are shortly basally fused, forming a tube. Anthers are basifixed and open on the inside with lengthwise slits. The superior ovary is composed of two, four or five fused carpels, each forming a locule. Fruits are a four- or five-seeded, leathery berry (Pentamerista and Tetramerista) or a turnip-shaped, ridged, indehiscent capsule with a single seed and spongy tissue (Pelliciera). Distribution: This is a widely disjunct family, with Pelliciera occurring along the Pacific coast in Central America and western Colombia and Ecuador, Pentamerista endemic to the Guianas and Tetramerista only in Southeast Asia. Phylogeny and evolution: The family has often been associated with Marcgraviaceae and in a broader sense with Theaceae. Molecular analyses have shown that Pelliciera (often formerly placed in its own family Pellicieraceae) is their closest relative, with Plants of the World

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Pentamerista neotropica, Colombia (MF) [363]

Pelliciera rhizophorae, Caldera, Puntarenas, Costa Rica (CD) [363]

Tetramerista glabra, Klias Forest Reserve, Sabah, Malaysia (CD) [363]

Fouquieria diguetii, The Living Desert, Palm Desert, California, USA [364]

Fouquieria columnaris, Rancho Santa Ana Botanical Garden, California, USA [364]

Fouquieria splendens, Palm Springs, California, USA [364]

which they share the flask-shaped glands at the base of the petal-like sepals that function as nectaries. Together Tetrameristaceae form a clade with Balsaminaceae (which has a nectary in a spur of a modified petal-like sepal) and Marcgraviaceae, with which they share leaf anatomy. The fossil record of Pelliciera appears to go back to the Tertiary and Cenozoic thermal maximum when it was more widespread, although the identity of these fossils may have to be reassessed. The range may have contracted due to changes in temperature and water salinity.

Genera and species: This family includes three genera with five species: Pelliciera rhizophorae, Pentamerista neotropica and Tetramerista (3).

364. FOUQUIERIACEAE

Christenhusz, Fay & Chase

Ocotillo family

Uses: Punah (Tetramerista glabra) has an edible fruit, and its timber is used for construction beams. Etymology: Tetramerista is composed of the Greek words τετρα (tetra), four, and μέρος (meros), part, referring to the four-parted flowers of that genus. The other genera in this family are pentamerous (five-parted).

These are spiny shrubs and trees with outwardly arched branches. Leaves are

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I see you! How big a prey can a carnivorous plant consume? We know of f lies and other insects becoming trapped in the leaves of Venus’ fly trap (Dionaea muscipula, Droseraceae). Many plants have sticky glands as a defence mechanism against herbivory, and some have found ways to use insects for their nutritional benefits (e.g. Byblidaceae, Droseraceae, Drosophyllaceae, Dioncophyllaceae, Roridulaceae, Solanaceae etc.). Bigger prey has been known to be trapped in large pitchers of Nepenthes, and seabirds, especially juveniles, become trapped and die on distant islands due to the weight of numerous sticky seeds of Pisonia. Anything even bigger? Maybe a dog, a deer, a small child or even an adult human? If we believe popular myths, there certainly are. larger victims. Man-eating plant feature in popular books (the Moomins, Harry Potter), films (Day of the Triffids) and musicals (Little Shop of Horrors) and many more sources, but from where does this fable originate? In a book about creatures of the sea and on land, Buel (1887) wrote a detailed description under the heading “A maneating plant”. He stated that it is “not content with the myriad of large insects which it catches and consumes, but its The spiny forest at Ifaty, Madagascar with Alluaudia procera (Didiereaceae), Jialiang Gao, Wikimedia Commons)

voracity extends to making even humans its prey”. He further described it as “having a short, thick trunk, from the top of which radiate giant spines, narrow and flexible, but of extraordinary tenaciousness, the edges of which are armed with barbs or dagger-like teeth. Instead of growing upright, these spines lay their outer ends upon the ground, and so gracefully are they distributed that the trunk resembles an easy couch with green drapery around it. The unfortunate traveler, ignorant of the monstrous creation which lies in his way or curious to examine the strange plant ... is without suspicion of his certain doom. The moment his feet are set within the circle of the horrid spines, they rise up, like gigantic serpents, and entwine themselves about him until he is drawn upon the stump, when they speedily drive their daggers into his body and thus complete the massacre. The body is crushed until every drop of blood is squeezed out of it and becomes absorbed by the goreloving plant, when the dry carcass is thrown out and the horrid trap set again”. What Ocotillo (Fouquieria splendens) near Palm Desert, California, USA

Ya te veo (page 476), from Buel JW. 1887. Sea and land. An illustrated history of the wonderful and curious things of nature existing before and since the deluge. Historical Publishing Company, Philadelphia & St Louis, USA.

could this plant be? Buel also stated that he found no “no description of it in the most elaborate works on botany”. “Yet hundreds of responsible travellers declare they have frequently seen it”, making one think that no botanist survived the collection of a specimen. Maybe it needs to receive a proper scientific account in case it is eventually found, or have these numerous accounts been based on an entirely different plant? The final reference is to South America as ya te veo (I see you) with a mention that the branches are constantly moving in the air. This made one wonder which plants look like the illustration and occur in the Americas, and immediately Fouquieria (although it is found strictly in the southern part of North America) comes to mind, with their short trunks and snakelike prickly branches. The references to Africa must be Madagascar as Alluaudia (Didiereaceae) can resemble Fouquieria (and they are often called it “Madagascan ocotillo” in Californian horticulture).

Plants of the World

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simple and alternate, nearly sessile among the spines and without stipules, petiolate on long shoots. They have entire margins and pinnate venation. Inflorescences are terminal or axillary spikes, racemes or panicles, sometimes corymbose, each flower bisexual, actinomorphic and subtended by two prophylls. The five sepals are free, and the five petals are fused into a tube. Stamens are usually ten (rarely up to 23) and unevenly exserted, with the filaments slightly fused to the base of the petal tube, and the anthers have pointed tips and are bilobed at the base, opening by lengthwise slits. The superior ovary is composed of three fused carpels, with a terminal style, branched at the tip into three. Fruits are loculicidal capsules with a central column and winged seeds.

Uses: Ocotillo (Fouquieria splendens) and boojum tree (F. columnaris) are frequently planted in xeriscaped gardens in the USA or planted as a living fence in their native area. Because of their light weight, ocotillo branches are used as walking sticks, and flowers are sometimes dried and used for tisanes. Etymology: Fouquieria is named for French physician Pierre Éloi Fouquier (1776–1850), who was personal physician to the Kings Charles X and Louis-Philippe of France.

365. POLEMONIACEAE Jacob’s-ladder family

Distribution: Fouquieriaceae are restricted to the arid zones of southwestern North America: from southern California, northern Arizona and eastern Texas south to southeastern Oaxaca (Mexico). Phylogeny and evolution: The family is closely related to Polemoniaceae, with which they share many chemical, vegetative and floral characters. They are remarkably similar in habit to woody members of Polemoniaceae, such as Acanthogilia and Cantua. They also share a biogeographical link; the family is found in the area where Polemoniaceae are most diverse. Fouquieriaceae separated from Polemoniaceae c. 61–65 million years ago. Genera and species: The sole genus in this family is Fouquieria with 11 species. Cobaea scandens, Royal Botanic Gardens, Kew, UK [365]

484

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Polemoniaceae are annual and perennial herbs, sometimes climbing and woody shrubs and small trees. Leaves are alternate or opposite, simple or pinnately or palmately compound, petiolate or not, without stipules. Blades have pinnate or sometimes palmate venation, and margins are entire to deeply lobed. Inflorescences are terminal dichasia organised in racemes, panicles or heads, and flowers are rarely solitary. Flowers are bisexual and usually actinomorphic, but

Cantua buxifolia, California, USA [365]

occasionally zygomorphic. The usually five (occasionally four or six) sepals are often fused and persistent in fruit. Petals are as many as sepals (usually five) and fused into a salveriform (or funnel-shaped) or bilobed corolla. The usually five (sometimes three, four or six) stamens alternate with the petals. Filaments are fused to the petal tube and unequal in length, topped with basifixed to dorsifixed anthers that open by lengthwise slits. A nectar disc surrounds the superior ovary, which is composed of (two or) three fused carpels, each forming a locule. It is topped by a style with (two or) three stigmatic branches. Fruits are usually loculicidal, rarely septicidal or indehiscent, capsules, sometimes explosively so. Seeds are smooth and sometimes winged. Distribution: This is a family with their greatest diversity in southwestern North America, but extending throughout North America into the western Canadian Arctic and south into Mesoamerica and along the Andes to southern South America. They are widespread in the Palaearctic, extending south into the mountains of Europe, the Caucasus, Pamir, Himalayas and Japan. Phylogeny and evolution: Polemoniaceae diversified c. 35 million years ago with a fossil similar to Gilia known from the Mid Eocene of Utah. The placement of Acanthogilia is still in doubt. It may be sister to all other Polemoniaceae or to just the Cobaea assembly. Because relationships between the small number of genera are not clear, classification into three subfamilies

Gilia inconspicua, New Mexico, USA (DZ) [365]

Giliastrum acerosum, New Mexico, USA (DZ) [365`]

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Ipomopsis aggregata, New Mexico, USA (DZ) [365]

Phlox divaricata, Tennessee, USA [365]

(Acanthogilioideae, Cobaeoideae and Polemonioideae) is not followed here.

Etymology: Polemonium is the Latinised form of πολεμονιών ( polemonion), the Ancient Greek name for the plant, possibly originally derived from πόλεμος (polemos), war, but its exact derivation is unknown.

Genera and species: Polemoniaceae include 18 genera and c. 350 species: Acanthogilia (1), Aliciella (21), Allophyllum (6), Bonplandia (1), Cantua (12), Cobaea (18), Collomia (c. 15), Eriastrum (14), Gilia (c. 40), Giliastrum (9), Gymnosteris (2), Ipomopsis (c. 30), Langloisia (3), Linanthus (c. 55), Loeselia (c. 14), Navarretia (31), Phlox (69) and Polemonium (28). Uses: Species of Ipomopsis, Loeselia and Polemonium have been used by Native Americans as a soap substitute. Cobaea flowers are used in Mesoamerica to make a cough syrup, and several species of the family may have anti-inf lammatory and anti-tumour properties. Several species are occasionally grown as ornamentals, especially species and selections of Cobaea scandens (cup-and-saucer vine), Collomia grandiflora (mountain trumpet), Eriastrum (woollystar), Ipomopsis (scarlet trumpet), Linanthus grandiflorus (mountain phlox), Phlox species (especially P. paniculata, garden phlox, and P. subulata, moss phlox, cultivars) and Polemonium (Jacob’s ladder). Cantua buxifolia (qantu) was sacred to the Incas and is now the national flower of Peru; it is also cultivated occasionally.

366. LECYTHIDACEAE Cannonball-tree family

This is a family of trees, sometimes large (to 55 m in Bertholletia excelsa) and sometimes shrubs, with simple, alternate leaves that usually crowd the tips of branchlets (opposite in Abdulmajidia). They are usually shortly petiolate, sometimes with small stipules, although usually not stipulate, and blades are pinnately veined and usually have an entire, sometimes serrate, margin. Inflorescences are terminal or axillary or formed on stems/ trunks (plants cauliflorous); they are racemes, fascicles, thyrses or panicles composed of racemose or spicate branches, rarely reduced

Polemonium caeruleum, private garden, Kingston upon Thames, Surrey, UK [365]

to a solitary flower. Flowers are often large, actinomorphic or zygomorphic and bisexual. Sepals are fused into a two- to six-lobed bellshaped tube, rarely without lobes, which is fused to the ovary. The three to six (rarely eight, 12 or 18) petals are free, sometimes absent. In some taxa without petals there are six to 28 petal-like staminodes that form a false corolla. The ten to numerous (>1,000) stamens are basally fused into a staminal ring (free in Foetidia), and the ring is actinomorphic or elongate on one side, forming a strap-like structure that arches over the top of the ovary, enclosing the flower opening. Anthers are basifixed and open by lateral or inward slits or apical pores. The inferior or half-inferior ovary is composed of two to eight carpels, each forming a locule and topped by a usually short style with an entire, rarely lobed stigma. Fruits are fleshy drupes or woody capsules, sometimes indehiscent, loculicidal or opening by a circumcissile lid (operculum), sometimes large. Seeds are winged (Cariniana, Couroupita) or not. Distribution: Lecythidaceae is a pantropical family with high diversity in the American tropics, but small centres of diversity are also found in tropical Africa, Madagascar, the Mascarenes, Asia, northern Australia and some Pacific islands. Plants of the World

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Barringtonia asiatica, New Caledonia [366] Barringtonia asiatica in fruit, New Caledonia [366] Couroupita guianensis, Rio de Janeiro Botanical Garden, Brazil [366]

Eschweilera itayensis, Rio Tigre, Nauta, Loreto, Peru (CD) [366]

Phylogeny and evolution: Previously, Lecythidaceae were placed in Myrtales, although an association with Theaceae has been suggested on morphological grounds as well. On the basis of wood anatomy, pollen morphology and floral structure, the family never was a good match for Myrtales. An association with Scytopetalaceae (originally in Malvales) was realised because they have similar floral and wood anatomy, and this family is now included in Lecythidaceae. Asteranthos seems to be intermediate morphologically between the two. Molecular studies have shown that Scytopetalaceae are embedded in Lecythidaceae, the latter sometimes divided into six subfamilies (Napoleonaeoideae, Scytopetaloideae, 486

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Gustavia augusta, Ecuador [366]

Foetidioideae, Planchonioideae, Barringtonioideae and Lecythidoideae). Few fossils are known from this family, the most reliable being a Miocene fossil flower from Trinidad, but fossils of wood and fruits of lecythidaceous plants have been reported from various areas in the Upper Cretaceous. Molecular estimates have dated the crown group at 46–65 million years old. Genera and species: This family has 25 genera and 355 species: Abdulmajidia (2), Allantoma (8), Asteranthos (1), Barringtonia (65), Bertholletia (1), Brazzeia (3), Careya (4), Cariniana (9), Chydenanthus (2), Corythophora (4), Couratari (19), Couroupita (3), Crateranthus (3), Eschweilera (94),

Napoleonaea imperialis, Fairchild Tropical Botanical Garden, Florida, USA [366]

Foetidia (18), Grias (12), Gustavia (42), Lecythis (27), Napoleonaea (10), Oubanguia (3), Petersianthus (2), Pierrina (1), Planchonia (8), Rhaptopetalum (11) and Scytopetalum (3). Uses: Economically, the most important crop of this family is the Brazil or Pará nut (Bertholletia excelsa), a popular snack around the world that is still harvested from the wild (logging is now banned because trees take many years to get to fruiting age). Pará nut oil is used commercially in food and cosmetics. Several other species of Barringtonia, Eschweilera and Lecythis also have edible seeds (especially sapucaia nuts, L. zabucajo), although some are poisonous, and Barringtonia is often used as a fish

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poison. Seeds of patana oak (Careya arborea) are eaten in Sri Lanka, and its leaves are sometimes fed to silk worms. Grias has avocado-like fruits that are eaten in Peru, and anchovy pear (G. cauliflora) is high in vitamin C and eaten in Mesoamerica and Jamaica. The pulp of Gustavia speciosa fruits is eaten in Colombia. In New Guinea and northern Australia, fruits and seeds of cocky apple (Planchonia careya) are popular. Colombian mahogany or albarco is the wood of Cariniana pyriformis, which has been overexploited and is now rare. Planchonia valida is used in the construction of houses in Indonesia, and essia (Petersianthus macrocarpus) is an important local timber in West Africa. The wood of Eschweilera (manbarklak) is resistant to marine borers and so is perfect for constructing boats and harbour quays. Many Lecythidaceae have attractive flowers, and for this reason they are sometimes cultivated in tropical gardens or as street trees in the tropics, most commonly the fish poison tree (Barringtonia asiatica), the emperor tree (Napoleonaea imperialis) and the cannonball tree (Couroupita guianensis). Gustavia species are attractive, and several would make good garden plants in the tropics. Etymology: Λήκυθος (lekythos) is a type of Greek pot (ampulla) used for storing olive oil. The name Lecythis was given to this genus in reference to its pot-shaped fruits, e.g. the so-called ‘monkey pots’.

tree of rainforests in the mountains of East Africa.

367. SLADENIACEAE Ribfruit family

These are evergreen trees with alternate, simple, petiolate leaves without stipules. Leaf blades have a pinnate venation and toothed or entire margins. Inflorescences are axillary cymes with lanceolate bracts. Flowers are bisexual and actinomorphic, subtended by two bracts. The five sepals are free and persistent. The usually five petals are free (Sladenia) or fused at the base (Ficalhoa). The eight to 13 (Sladenia) or 15 (Ficalhoa) stamens form a single whorl in which they are free or fused to the base of the petal tube, with short basally thickened filaments and basifixed, hairy anthers that open by apical pores or slits. There is no nectary, and the superior ovary is conical, composed of three or five fused carpels. The ovary is topped by a fused, slightly three-parted style. Fruits are loculicidal capsules (Ficalhoa) or a schizocarp (Sladenia). Distribution: Sladenia is restricted to a small area in Indochina. Ficalhoa is a

Sladenia celastrifolia, Kunming Botanical Garden, China [367]

Phylogeny and evolution: Sladenia was usually associated with Theaceae or Pentaphylacaceae and was also included in Actinidiaceae, from which it was excluded and placed in its own family. Ficalhoa had a similar history, being included in Ericaceae initially and later moved to Actinidiaceae from where it was removed and placed with Sladenia in Theaceae. Molecular studies placed the two genera together in a position separate from Pentaphylacaceae, with which they share some characters. Despite their geographical separation they have similarities, and 100-million-year-old fossil wood from Sudan closely matches the wood of extant Sladenia, providing a phytogeographical link between the two genera. Genera and species: Sladeniaceae include only two genera and three species: Ficalhoa laurifolia, Sladenia celastrifolia and S. integrifolia. Etymology: Sladenia is named “after Major Sladen, the energetic head of the Yunnan expedition”. Sir Edward Bosc Sladen (1827–1890) was a British army officer, working in India; he headed a political mission to the Chinese frontier in 1868, during which this genus was first collected.

Ficalhoa laurifolia, Tanzania (RB) [367]

Plants of the World

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Eurya japonica, Royal Botanic Gardens, Kew, UK [368]

EUDICOTS

Ternstroemia tepezapote, San Francisco Botanical Garden, California, USA [368]

Adinandra dumosa, Singapore [368]

368. PENTAPHYLACACEAE Saintedwood family

This is a family of bisexual and unisexual evergreen trees and shrubs with alternate leaves arranged spirally or in a plane (distichous). Leaves are petiolate and lack 488

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Ternstroemia impressa in fruit, University of California Botanical Garden, Berkeley, USA

[368]

Cleyera integrifolia, San Francisco Botanical Garden, California, USA [368]

stipules and have pinnate venation and entire or toothed margins. when toothed with a small hair on the tip of each tooth. Inflorescences are axillary fascicles, or flowers are solitary, rarely in axillary racemes. Flowers are unisexual or bisexual and actinomorphic. The five sepals are fused or free and persistent in fruit. The five petals are free or basally fused, persistent or not. The five to numerous stamens are free, fused to each other or with the base of the petal tube. Filaments are flattened or spreading, often thickened at base and incurved at the tip. Anthers are basifixed and open on the inside by lengthwise slits. A connective is often present and elongate above the anthers. The superior (rarely inferior) ovary is composed

of usually three to five carpels topped with a simple or apically branched style, or each carpel topped with a style; stigmas are usually free. Fruits are berries or a loculicidal capsule. Distribution: This is a pantropical family, widespread across the American and Asian tropics. They extend into temperate East Asia (north to Korea and southern Japan), south to northern Australia, and the Canary Islands and Madeira. Phylogeny and evolution: Previously included in Theaceae, this family was found to be closer to a clade including Primulaceae, Ebenaceae and Sapotaceae than to Camellia (Theaceae)

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and its relatives. Pentaphylax itself was often not included in Theaceae because of differences in anther morphology, but molecular studies placed them as sister to Ternstroemiaceae as the clade was previously called (Pentaphylacaceae is a conserved name that thus has priority). They are closely related to Sladeniaceae, which could also be included on morphological grounds. Pentaphylacaceae differ little from Theaceae s.s., the minor differences being in palynology and wood anatomy. Pentaphylacaceae appear to have been common throughout Europe in the lauraceous forests during the Late Cretaceous and Tertiary, and several reliable fossils from Baltic amber and formations in southern Sweden and Central Europe are known. In Europe, the family is now restricted to Macaronesia (Visnea). Genera and species: This family includes 12 genera with c. 330 species: Adinandra

(c. 80), Anneslea (3), Archboldiodendron (1), Balthasaria (1), Cleyera (8), Eurya (c. 70), Eurydendron (1), Freziera (57), Pentaphylax (1), Symplococarpon (9), Ternstroemia (c. 100) and Visnea (1). Uses: Anneslea fragrans produces timber with beautiful markings, used for furniture and carving. Cleyera wood is also used for carving and making wooden combs in Japan. Some species are occasionally grown as ornamentals in specialist collections, especially Adinandra, Cleyera, Eurya and Ternstroemia. Sakaki (Cleyera japonica) is a sacred tree in Shintoism and is therefore often planted near shrines. Etymology: Pentaphylax is composed of the Greek words πέντε (pente), five, and φύλαξ ( fylax), a guardian, in reference to the five anthers that seem to ‘stand guard’ around the ovary.

369. SAPOTACEAE Sapodilla family

Trees, shrubs and lianas make up this family. Stems are sometimes swollen, sometimes spiny; they always have latex, which is usually white, rarely yellow (Chromolucuma congestifolia) or blue (Niemeyera acuminata). Hairs are always T-shaped (malpighiaceous), rarely simple (Delpydora). Leaves are simple, alternate, spirally arranged or in a plane (distichous), rarely opposite or in whorls, with or without stipules and petioles sometimes with

Sideroxylon borbonica, Réunion [369]

Manilkara zapota, Turbaco, Bolívar, Colombia [369]

Argania spinosa, Morocco (HR) [369]

Mimusops sechellarum, Seychelles [369]

Plants of the World

489

ERICALES

Chrysophyllum cainito, Royal Botanic Garden, Sydney, Australia [369]

EUDICOTS

Synsepalum dulcificum, Matthaei Botanical Garden, Ann Arbor, Michigan, USA [369]

a pair of small stipels (stalked glands). Blades are pinnately veined and have entire or (rarely) prickly toothed margins. Inflorescences are axillary or cauliflorous panicle-like fascicles, or flowers are solitary. Flowers are bisexual or unisexual and actinomorphic. Sepals are four to six in a single whorl, four to 11 in a spiral, or four to eight in two whorls, free or fused into a tube. The four to 18 petals are fused into a bell- or cup-shaped tube, with shallow or deep lobes. There are four to 43 stamens fused at various heights in the petal tube, rarely free, rarely fused into a stamen tube, sometimes partly staminodial and then filament- or petallike. Anthers are basifixed or dorsifixed, open on the inside by lengthwise slits, and often are below a protruding connective. A nectar disk is often present surrounding the ovary base and sometimes fused with it. The superior ovary has one to 30 locules and is topped with a simple style that has a simple or slightly lobed stigma. Fruits are berries or (sometimes) a drupe, sometimes roughly opening by a valve, usually fleshy, sometimes leathery or woody. Seeds are usually smooth and shiny, usually with a large hilum, a good character to recognise seeds of this family. Distribution: Sapotaceae are a pantropical family usually found in humid forests, but some extending into arid regions and temperate regions in southern North America, warm-temperate South America, Morocco, subtropical East Asia, Australia and northern New Zealand. 490

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Phylogeny and evolution: Sarcosperma is sister to the rest of Sapotaceae, in which two clades (given subfamily rank: Chrysophylloideae and Sapotoideae) can be recognised. Some generic delimitations are still uncertain, and widely different conclusions were drawn on the basis of morphology in the past; however, molecular studies have helped to elucidate relationships and have aided delimitation. The age of the family is estimated to be c. 92 million years. The biogeographical patterns show a high level of long-distance dispersal; they must have reached New Caledonia at least nine times since the island emerged from the sea c. 32 million years ago. This puzzles biogeographers because seeds of this family are generally large and dispersed mostly by larger animals, although fruit bats may have played a role. Genera and species: Sapotaceae include c. 58 genera with 1,273 species in three subfamilies: Sarcospermoideae – Sarcosperma (11); Chrysophylloideae – Aubregrinia (1), Boerlagella (1), Breviea (1), Capurodendron (23), Chromolucuma (5), Chrysophyllum (87), Delpydora (2), Diploon (1), Ecclinusa (11), Elaeoluma (4), Englerophytum (14), Magodendron (2), Micropholis (38), Niemeyera (3), Omphalocarpum (27), Pichonia (10), Pleioluma (32), Planchonella (106), Pouteria (210), Pradosia (26), Pycnandra (45), Sarcaulus (5), Sersalisia (2), Synsepalum (35), Tridesmostemon (2),

Pouteria campecheana, seed and fruit, Merida, Yucatán, Mexico [369]

Tsebona (1), Van-royena (1) and Xantolis (14); Sapotoideae – Amorphospermum (1), Argania (1), Aulandra (3), Autranella (1), Baillonella (1), Burckella (13), Diploknema (7), Eberhardtia (3), Faucherea (11), Gluema (2), Inhambanella (2), Isonandra (10), Labourdonnaisia (7), Labramia (9), Lecomtedoxa (6), Letestua (1), Madhuca (115), Manilkara (79), Mimusops (45), Neohemsleya (1), Neolemonniera (3), Northia (1), Nesoluma (3), Palaquium (120), Payena (20), Sideroxylon (78), Tieghemella (2), Vitellaria (1) and Vitellariopsis (5). Uses: Many species of Sapotaceae produce edible fruits. Sapodilla (Manilkara zapota), which gave the name to the family, has long been cultivated in Mexico and is now grown for its fruits worldwide. Star-apple (Chrysophyllum cainito) from the Caribbean is also a popular dessert fruit. Other species commonly grown for their fruits are damson-plum (Chrysophyllum oliviforme), mahua (Madhuca longifolia), African pear (Manilkara obovata), Spanish cherry (Mimusops elengi), abiu (Pouteria caimito), canistel (P. campecheana), lúcuma (P. lucuma) and mamey sapote (P. sapota). Miracle berry (Synsepalum dulcificum) contains miraculin, which affects the taste buds, making sour and salty foods taste sweet. The shea tree (Vitellaria paradoxa) has oil-rich seeds that can be pressed into shea butter, which finds many uses, especially as an emollient in cosmetics, and can replace palm

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oil in food. Argan oil, high in vitamin E, is pressed from the seeds of Moroccan (near) endemic Argania spinosa, which is highly valued in cosmetics (facials, aftershave) and as a food oil with a delicious flavour. The tree also produces a valuable gum. It is mostly harvested from wild populations, although it has been introduced to southern Spain and Libya. Madhuca species have oil-rich seeds that are called illipe nuts, used previously to make margarine. Seeds of many American Pouteria species have similar oils that are yet to be exploited. Latex is abundant in many species, and especially the sap of Manilkara and Palaquium was formerly used to insulate marine cables and to make machine belts and rubber soles and was the chewing gum of the Aztecs. As with most other tropical trees, timbers are harvested from many species, and used for plywood, flooring or construction. Species are sometimes cultivated, especially maulsari or bullet wood (Mimusops elengi), which has fragrant flowers used for garlands in India. Historical curiosities: Northia seychellana, on Mahé known as capucin, was named by Joseph Hooker for Marianne North, who painted this species in the Seychelles and

exhibited it in a remarkable gallery at Kew with numerous other paintings, which can still be seen there. On another Indian Ocean Island, Mauritius, flocks of dodos almost certainly ate and distributed sapotaceous fruits around the rainforest. One of these species is called tambalacoque (Sideroxylon grandiflorum), endemic to Mauritius, but because the wood is highly valued, there were only a few trees left in the 1970s. It was postulated that this decline and lack of regeneration was because the seed needed to pass through the digestive tract of a dodo to germinate, but other animals such as tortoises may have helped as well. Germination is now achieved by damaging the seed coat or feeding them to a turkey, and the species is no longer endangered. Several other species are thought to be adapted to distribution by now extinct or rare megafauna, but some regeneration of most of these happens despite the absence of these animals. Etymology: Sapota is the Latinised form of Aztec Nahuatl tzapotl, meaning ‘soft fruit’, and was thus adopted by the Spanish (as sapote) for several unrelated soft fruits found in the region. Sapota is a name nomenclaturally rejected against Manilkara.

Euclea undulata, Hluhluwe Game Reserve, KwaZulu Natal, South Diospyros kaki, Işhan, Turkey [370] Africa (CD) [370]

370. EBENACEAE Persimmon family

Ebenaceae are evergreen and sometimes deciduous shrubs and trees with clear sap and hardwood that is often black (ebony). Hairs are simple or two-armed, sometimes also with glandular or multicellular hairs. Leaves are simple, usually alternate (rarely opposite or whorled), organised in planes along the stem (distichous), petiolate and without stipules. Leaf blades have fine, pinnate venation and entire margins, often bearing nectaries on the lower surface. Inflorescences are axillary cymes, racemes, fascicles or panicles. Flowers are unisexual, occasionally bisexual, and actinomorphic. The three to five (to eight) sepals are fused at least basally, persistent and often enlarged in fruit. There are as many petals as sepals, usually four or five, fused into a tube that is often short in male flowers and

Diospyros whyteana, Royal Botanic Gardens, Melbourne, Australia [370]

Diospyros virginiana, Royal Botanic Gardens, Kew, UK [370]

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491

ERICALES enlarged and urn-shaped in female flowers. In most Lissocarpa, the four-lobed petal tube is adorned with a corona with eight small lobes. Male flowers have eight or 12 to 100 stamens, usually three to five times as many as petals (rarely as many as petals), the filaments are fused at the base to the petal tube, free or fused into pairs, threes or bunches, often unequal in length. Anthers are basifixed, often with a pointed connective and thecae opening longitudinally. Pistillodes are often present in the middle of male flowers. Female flowers often have staminodes. The superior (inferior in Lissocarpa) ovary is composed of two to eight fused carpels, usually forming eight locules. The terminal style is short with branches longer than the rest of the style and as many branches and stigmas as there are carpels. Fruits are usually berries (rarely capsules), with an often astringent endocarp, and up to 16 seeds per fruit. Seeds are shiny with a small, apical hilum.

differs from Diospyros in its corona and inferior ovary, among other features. The other three genera recognised commonly (Maba, Rhaphidanthe and Tetraclis) are now recognised as subgeneric or sectional ranks under Diospyros. The family is placed with strong support in a clade with Primulaceae and Sapotaceae. Tertiary fossils of Ebenaceae are known, but leaves are usually so non-descript that family placement is often uncertain. Crown Ebenaceae are dated to c. 54 million years old. There is a remarkable radiation of Diospyros on New Caledonia, representing four independent lineages. Genera and species: Ebenaceae includes four genera and c. 580 species: Diospyros (c. 500), Euclea (c. 16), Royena (c. 56) and Lissocarpa (8). A taxonomic revision of the family is greatly needed.

used in South Africa to make vinegar. Guarrie bark is insecticidal and produces a dye. Apart from their fruits, Ebenaceae are best known for the timber, ebony, used mainly for carving, musical instruments, furniture inlays, cabinetry and decorative veneer. Most species of Diospyros yield a black or white ebony, but the true ebony wood is mainly harvested from Indian ebony (D. ebenum) and tendu (D. melanoxylon). Cape ebony (Eurya pseudebenus) also has valuable black timber used for furniture. Etymology: Ebenus is Latinised from Greek έβενος (hebenos), black wood or ebony. The genus Ebenus of Kunze on which Ebenaceae is based, is a homonym of the earlier Ebenus of Linnaeus (Fabaceae) and is a later synonym of Diospyros. Διόσπυροσ (diospyros) in turn is Greek for ‘divine wheat’, in reference to the starchy fruit of the date plum, D. lotus.

371. PRIMULACEAE

Phylogeny and evolution: Generic inclusions and circumscriptions have varied in Ebenaceae, although generally only pantropical Diospyros and African Euclea have been accepted. Royena was found to be sister to Diospyros and has been accepted as separate from Diospyros. Lissocarpa (originally described in Styracaceae) was found to be sister to the rest and has in the past been placed in its own subfamily or family. It

Uses: Many species of Diospyros have edible fruit, but only the kaki or sharon fruit (D. kaki and cultivars) is widely grown. Locally, common persimmons (D. virginiana) from North America and date plum (D. lotus) from Eurasia are also consumed (the latter usually dried or after frost), and many other species have edible fruit, although they are usually astringent, unless over-ripe and after a frost. Other species eaten are yellow persimmon (D. australis), velvet apple (D. blancoi), sand apple (D. chamaethamnus), gold apple (D. decandra), black sapote (D. digyna), jackalberry (D. mespiliformis) and Texas persimmon (D. texana). Sea guarrie (Euclea racemosa subsp. schimperi) has a fruit that is

Maesa lanceolata, Royal Botanic Gardens, Kew, UK [371]

Clavija sp., Royal Botanic Gardens, Kew, UK [371]

Deherainia smaragdina, Royal Botanic Gardens, Kew, UK [371]

Distribution: Ebenaceae are a pantropical family, extending north into temperate eastern North America, southeastern Europe, southwestern Asia and temperate East Asia.

492

EUDICOTS

Christenhusz, Fay & Chase

Primrose family

This varied family includes annual and perennial herbs, shrubs, trees, mangroves (Aegiceras) and vines. The simple or rarely pinnatifid or pinnate leaves are usually

ERICALES

EUDICOTS

Samolus valerandi, Biarritz, France [371]

Bonellia, Mérida, Yucatan, Mexico [371]

Bonellia in fruit, Mérida, Yucatan, Mexico [371]

alternately arranged, sometimes opposite or in whorls, distributed spirally along the stem, in rosettes or crowded near the stem apex. Blades have a pinnate, palmate or pinnateparallel venation, and margins are entire or variously toothed, crenate, serrate, lobed or pinnately dissected. Inflorescences are terminal or axillary racemes, spikes, corymbs or (multiple) umbels, sometimes scapose, reduced to axillary fascicles or with solitary flowers. Bracts are present or not. Flowers are bisexual or unisexual and actinomorphic or rarely slightly zygomorphic. The usually five, but frequently three to nine, sepals are fused into a short, often bell-shaped tube or the lobes are nearly free. Petals are as many as sepals or absent (Lysimachia maritima) and fused into a bell- or urn-shaped tube or the lobes nearly free. Corolla lobes are entire, emarginate, dissected or fringed, sometimes reflexed and twisted. Stamens are in a single whorl, as many as and opposite the petals, the filaments entirely or partly fused with the petal tube. Anthers are dorsifixed and open inside by lengthwise slits or on top with porelike slits, free or rarely fused together around the style. Staminodes are present or not. The superior (inferior or half-inferior) ovary is composed of five carpels, fused to form a

single locule with a free central column. The ovary is topped by a single, hollow style (sometimes heterostylous), with a punctate, capitate, truncate or discoid stigma. Fruits are fleshy drupes or capsules that open by a valve, a circular lid (operculum), loculicidally (Samolus) or by disintegration. Cotyledons are usually two but reduced to one in Cyclamen.

the families Maesaceae, Myrsinaceae, Primulaceae and Theophrastaceae. Molecular studies have shown that Primulaceae in the traditional sense are not monophyletic, with Samolus falling with Theophrastaceae and Ardisiandra, Coris, Cyclamen, Lysimachia, Stimpsonia and Trientalis being related to Myrsinaceae. Moreover, Myrsinaceae were also not monophyletic in the traditional sense, with Maesa sister to the entire lineage. One could keep the family names, but their content would be highly altered, so they were all merged into the single family Primulaceae, with the main clades recognised at subfamilial level. The family is united by having usually spirally arranged leaves, fused, usually pentamerous flowers, with stamens opposite the petal lobes, an ovary with free-central placentation and a hollow style with an entire stigma. Generic circumscription is poor at best, and phylogenetic studies have indicated that Primula should be recircumscribed to include Cortusa, Dionysia, Dodecatheon, Kaufmannia and Sredinskya. Androsace should include Douglasia, Vitaliana and Pomatosace to be monophyletic. Additionally Anagallis, Asterolinon, Glaux and Pelletiera are embedded in and have been transferred to

Distribution: A nearly cosmopolitan family found from the Arctic to the tropics. Maesoideae are restricted to the Palaeotropics. Theophrastoideae are mostly distributed in the Neotropics, with Samolus in temperate North and South America, southern Africa, Europe, West Asia, Japan, Australia and New Zealand. Primuloideae are mostly in the temperate Northern Hemisphere, Arctic and mountains, with some outliers in the far south of South America, the Falkland Islands, the Horn of Africa and Indonesian mountains. Myrsinoideae are most diverse in the tropics, but their herbaceous members extend them into the temperate zones and the Arctic and Antarctic. The family is absent from the great deserts of the world. Phylogeny and evolution: This family was previously treated as an order including

Plants of the World

493

ERICALES

EUDICOTS

Primula meadia, Helsinki Botanical Garden, Finland [371]

Soldanella alpina, private garden, Kingston upon Thames, Surrey, UK [371]

Lysimachia, Myrsine is expanded to include Rapanea and Suttonia, and Badula is embedded in Oncostemum. Generic delimitation in woody Myrsinoideae in general is poorly studied at best, and some realignments are likely in the future. Not all combinations have been made, for instance Cortusa and Dionysia have not been transferred to Primula, but we have anticipated some of these findings in the list of genera below. The fossil history of this family is sparse; fossil Myrsinoideae-type pollen appearing first during the Oligocene of Australasia, and seeds of Androsace are known from the Siberian Miocene. The crown group of Primulaceae has been estimated to be c. 46–61 million years old. Genera and species: Primulaceae include c. 53 genera and about 2,790 species, in four subfamilies: Maesoideae – Maesa (c. 150); Theophrastoideae – Bonellia (22), Clavija (c. 55), Deherainia (2), Jacquinia (13), Neomezia (1), Samolus (15), Theophrasta (2) and Votschia (1); Primuloideae – Androsace (c. 160), Bryocarpum (1), Hottonia (2), Omphalogramma (15), Primula 494

Christenhusz, Fay & Chase

Primula vulgaris, Cornwall, England, UK [371]

(c. 475) and Soldanella (16); Myrsinoideae – Aegiceras (1), Amblyanthopsis (2), Amblyanthus (3), Antistrophe (5), Ardisia (c. 500), Ardisiandra (3), Badula (c. 14 +3 extinct), Conandrium (2), Coris (1), Ctenardisia (5), Cybianthus (c. 165), Cyclamen (22), Discocalyx (c. 50), Elingamita (1), Embelia (c. 130), Emblemantha (1), Fittingia (6), Geissanthus (c. 55), Grenacheria (6), Heberdenia (1), Hymenandra (17), Labisia (7), Loheria (6), Lysimachia (c. 175), Monoporus (9), Myrsine (c. 300), Oncostemum (c. 90), Parathesis (c. 95), Pleiomeris (1), Sadiria (4), Solonia (1), Stimpsonia (1), Stylogyne (c. 60), Systellantha (2), Tapeinosperma (55), Trientalis (2), Vegaea (1) and Wallenia (c. 20). Uses: Fruits of Clavija are eaten as a forest snack in Latin America. Jacquinia flowers were used to decorate temples by the Mayan people. Atki (Maesa indica) leaves are an ingredient in Indian curries. Maesa lanceolata seeds produce an oil that is used for cooking in Ethiopia. The fruits of Maesa are bactericidal and used to combat intestinal parasites. False black pepper (Embelia ribes) is sometimes used to adulterate black pepper

Omphalogramma delavayi above Dali, Yunnan, China [371]

because it looks similar. It is used against flatulence and tapeworms in India. Many genera have species that are popular in horticulture, especially Androsace (rock jasmine), Ardisia (coralber r y), Cyclamen (cyclamen), Jacquinia, Lysimachia (yellow loosestrife, pimpernel), Myrsine (Cape myrtle), Omphalogramma, Soldanella (snowbell) and Primula (primrose; and its former segregates Cortusa, Dionysia and Dodecatheon still sold under their old names), especially as rock-garden, border or pond-margin plants. Hottonia palustris (water violet) and Samolus valerandi (brookweed) are popular aquarium or terrarium plants. Cultivars of Cyclamen persicum, Primula auricula, P. malacoides, P. obconica, P. sinensis and P. vulgaris, and several types with complex ancestry are important in the pot and bedding plant industry. Wild collected Cyclamen fall under CITES legislation aimed at preventing overcollecting. Etymology: Primula is deminutive from Latin primus, the first, referring to the early spring flowers of some species.

ERICALES

EUDICOTS

Ardisia crenata, Réunion [371]

Ardisiandra wettsteinii, Vuria Hill, Taita Hills, Kenya [371]

Androsace delavayi, Yunnan, China [371]

Cyclamen coum, Royal Botanic Gardens, Kew, UK [371]

Myrsine africana, Yale Rock, Taita Hills, Kenya [371]

Hottonia palustris, Twickel Estate, Delden, the Netherlands [371]

Coris monspeliensis, Royal Botanic Gardens, Kew, UK [371]

Lysimachia arvensis, Ruissalo Botanical Garden, Turku, Finland [371]

Lysimachia vulgaris, Savonlinna, Finland [371]

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495

ERICALES

372. THEACEAE Tea family

Theaceae are evergreen and sometimes deciduous trees and shrubs that have simple, alternate, spirally or distichously arranged leaves without stipules. Leaf blades are pinnately veined and usually toothed, with a gland on each tooth, rarely the margin entire. Inflorescences are reduced to solitary axillary flowers that are usually large and showy with several bracteoles below the flower integrading with the calyx. Flowers are bisexual and actinomorphic. The five (or more) sepals are free or basally fused, usually persistent in fruit, sometimes unequal in size. The free or basally fused petals are five, rarely numerous, and in a single whorl or spirally arranged. The 20 or more stamens are free or often fused to the base of the petals, sometimes in five fascicles. Anthers are dorsifixed and versatile, rarely basifixed, opening by lengthwise slits. The superior ovary is composed of usually five (sometimes three to ten) fused carpels, topped by a simple branched style or free styles with lobed stigmas. Fruits are loculicidal capsules, rarely a drupe, with a persistent columella. Camellia azalea, Kunming Botanical Garden, Yunnan, China [372]

496

Christenhusz, Fay & Chase

EUDICOTS

Distribution: This family is disjunct between tropical and subtropical America (extending north to the southeastern USA) and tropical and warm temperate East Asia, where their greatest diversity occurs. Phylogeny and evolution: A 91-millionyear-old fossil from eastern North America has been attributed to Theaceae but could instead be Pentaphylacaceae. Leaf fossils from the Late Cretaceous and Tertiary are similarly ambiguous. Theaceae seeds are known from fossil beds, showing that there was a greater diversity and distribution of Theaceae in the Mid Eocene of the Northern Hemisphere. Fossil wood attributed to Schima is known from Borneo, and Camellia fossils are known from Europe and North America, where the genus does not currently occur. Molecular studies have shown that Theaceae are sister to Symplocaceae, Styracaceae and Diapensiaceae, whereas Pentaphylacaceae (formerly included in Theaceae) are more closely related to Primulaceae, Ebenaceae and Sapotaceae. Genus and species delimitation in Theaceae are poorly understood, and some genera form intergeneric hybrids in cultivation. Camellia, Gordonia and Schima have many taxa that are difficult to distinguish. Laplacea and Polyspora are probably distinct from Gordonia, a clade that needs further taxonomic investigation. Genera and species: Theaceae include nine genera and c. 240 species: Apterosperma (1), Camellia (c. 120), Franklinia (1, extinct Camellia saluensis, Royal Botanic Gardens, Kew, UK [372]

in the wild), Gordonia (2), Laplacea (2), Polyspora (c. 30), Pyrenaria (42), Schima (c. 15) and Stewartia (30). Uses: Economically the most important species in the family is the tea plant (Camellia sinensis), the young leaves of which are processed into tea. When tea dries, the leaves oxidise and turn darker, so the sooner it is dried (usually by heat) the lighter the tea. White, yellow and green tea are made from freshly dried leaves, whereas oolong and black tea are fermented and then dried. Tea was originally used for medicinal purposes in China, brewed into a drink at least since 350 BC, and indeed it has many beneficial properties. Tea was introduced into Japan in the 8th century AD. A special variety was grown there, and a separate ceremonial culture surrounding the drink evolved during the 15th century. In the 1830s, seeds were taken from Japan to Java, where most of the Japanese tea is now grown. Drinking tea in Britain and Europe became popular in the 17th century, and the British East India Company introduced tea to India and Sri Lanka, from where it maintained a monopoly on the import of tea to Europe. Tea is now mostly grown in China, India, Kenya, Sri Lanka, Turkey, Vietnam, Iran, Indonesia, Japan and Argentina, but plantations can also be found in unexpected places like British Columbia (Canada), Turkey, the Republic of Georgia and Cornwall and Wales (UK). Tea oil is pressed from the seeds of Camellia oleifera and is used mainly in

Camellia sinensis, Réunion [372]

ERICALES

EUDICOTS

Camellia yunnanensis var. trichocarpa in fruit, Kunming Botanical Garden, Yunnan, China [372]

Schima argentea, Kunming Botanical Garden, Yunnan, China [372]

Franklinia alatamaha, private garden, Kingston upon Thames, Surrey, UK [372]

cosmetics (hair oil), lubrication, stamp pads, soap and sometimes cooking. Other species used for oil are tsubaki oil (from C. japonica) and the somewhat inferior oil from yuletide camellia (C. sasanqua). Several species are popular garden ornamentals, especially Camellia japonica and cultivars, and other Camellia species (especially C. reticulata, C. saluensis and C. sasanqua). Members of the genera Franklinia, Gordonia and Stewartia are fairly frequently encountered in subtropical and temperate gardens.

Etymology: The word tea is derived from Chinese 茶 (tu), used to describe a bitter herb. In Mandarin, this character is now usually pronounced as cha; however, in southern China, it is called te, and this word was spread from Amoy (Xiamen) throughout Southeast Asia and into Portuguese, English and other European languages. Latin thea is derived from this, and became the genus given to the tea plant by Linnaeus. Thea is now considered a synonym of Camellia, a genus that was named for George Joseph Kamel (1661–1706), who studied the flora of the Philippines.

Franklin’s extinct shrub: When exploring the Altamaha River in the southern US state of Georgia, botanists John and William Bartram from Philadelphia found some ‘very curious shrubs’ in 1765. They returned several times to the site in the following years and collected material and seeds, which they grew in their garden in Philadelphia, and they sent plants to John Fothergill in London, who funded some of the expeditions. They decided that it was a new genus and named it for a family friend, the famous Benjamin Franklin. Franklinia alatamaha had a very limited distribution in the wild, and pressure on the land from cotton farming, fires and floods resulted in destruction of the population by the 1840s. By that time, the plant had found some popularity as a garden plant, all descendents of the Bartram collections, which are now occasionally grown in warm temperate gardens in Europe and North America.

373. SYMPLOCACEAE Sweetleaf family

These are evergreen, rarely deciduous, trees and shrubs. The alternate, spirally or distichously arranged leaves are petiolate, without stipules. Leaf blades are serrate, dentate or crenulate, rarely nearly entire. Inflorescences are usually axillary (rarely internodal), bracteate panicles, thyrses, fascicles, glomerules, racemes or spikes or flowers are solitary. Flowers are usually

Stewartia pseudocamellia, Helsinki Botanical Garden, Finland [372]

bisexual and actinomorphic, supported by two bracteoles below the sepals, rarely bractless or with several bracts in the leaf axil. The (three to) five petals are basally fused and persist in fruit. The five (sometimes three to 11) petals are basally fused, split to the middle or almost to the base. The usually numerous, sometimes four or five, stamens are fused to the base of the petals. Filaments are fused to some degree, often grouped in five bundles opposite the petals, sometimes hairy. Anthers are basifixed or versatile and open by lengthwise slits. The inferior ovary is composed of two or three fused carpels, forming up to five incomplete locules. A nectary usually surrounds the single, hollow style, which is topped by a two- to five-lobed stigma. Fruits are drupes. Distribution: Like Theaceae, Symplocaceae have a disjunct distribution in the Americas and East Asia. They are found in southeastern North America, the Antilles and throughout Central and South America and in East Asia from Korea, Japan and northern China, south to the Indian Subcontinent, Sri Lanka, throughout tropical Asia, east to Melanesia, eastern Australia and New Caledonia. Cordyloblaste is restricted to eastern Asia. Phylogeny and evolution: Symplocaceae have long been associated with Styracaceae, and the two were usually placed near Theaceae. Morphological placement is congruent with molecular findings, and the Plants of the World

497

ERICALES

EUDICOTS

Symplocos paniculata, Mt Huangshan, Anhui, China [373]

three families are now placed close together in Ericales, with Symplocaceae sister to Styracaceae and Diapensiaceae. Within Ericales, Symplocaceae are distinguished by their exstipulate leaves, actinomorphic flowers with fused petals, stamens connate with petals, inferior ovaries, simple styles and drupaceous fruits. These features and recent phylogenetic studies based on DNA sequence data and morphology leave no doubt that Symplocaceae constitute a clade, with two genera now accepted, Hopea now being treated as a subgenus of Symplocos. Symplocaceae have an abundant fossil fruit and pollen record in the Tertiary of Europe, North America, East Asia and New Zealand, so the current distribution is most probably relictual. Genera and species: This family includes only two genera with c. 260 species: Cordyloblaste (2) and Symplocos (c. 260). Uses: Bogotá tea (Symplocos theiformis) is used as a tea substitute in Colombia. Bark and leaves of S. racemosa and S. tinctoria are used to produce red and yellow dyes, respectively. Wood of some species is used for turning. Etymology: Symplocos is derived from the Greek συμπλοκή (symploke), a connection, in reference to the fused stamens. 498

Christenhusz, Fay & Chase

Symplocos paniculata, Arboretum Poortbulten, De Lutte, the Netherlands [373]

374. DIAPENSIACEAE Pincushion-plant family

Low-growing, often mat-forming, evergreen, perennial herbs, sometimes with woody base, form this family. Stems are often creeping and rooting at nodes, sometimes with taproots and usually mycorrhizal. Leaves are simple, alternate and without stipules, usually crowded on the stems with short petioles, or blades are larger and long-petiolate in a basal rosette. Leaf blades have pinnate or palmate venation, sometimes (Diapensia, Pyxidanthera) with secondary veins that curve basally and become fused towards the base but do not rejoin the primary vein. Leaf margins are entire, toothed or serrate. Inf lorescences are terminal racemes, or flowers are solitary. Flowers are bisexual and actinomorphic. The five sepals are free or fused into a short tube, persistent in fruit. The five petals are fused into a short basal tube or nearly free (Galax). The five stamens are

fused basally to the petal tube and alternate with the lobes. Filaments are free from each other or are fused to form a basal ring, often alternating with five, scale- or spoon-like sterile stamens (these absent in Diapensia and Pyxidanthera). Anthers are basifixed and open by longitudinal or transverse slits. The superior ovary is composed of three carpels, each forming a locule and topped with a single style and a trilobed stigma. Fruits are loculicidal capsules. Distribution: This family has a mostly Arctic distribution, extending into temperate eastern North America and Eurasia (e.g. Scotland, Lapland, Siberia, Eastern Himalaya, Taiwan, Korea and Japan). Diapensia lapponica was only discovered in the UK in 1951 on a Scottish mountain, making Diapensiaceae the most recent family to be added to the flora of the UK. Phylogeny and evolution: Diapensiaceae have mycorrhizal associations and are herbs or small shrubs, and so an association with Ericaceae was always assumed. However, molecular results place this family as sister to Styracaceae, which were previously associated with Theaceae. A phylogenetic analysis based on morphology and molecular data indicated that Galax is sister to all other genera, and Pyxidanthera is sister to the rest.

ERICALES

EUDICOTS

[374]

Berneuxia thibetica, Royal Horticultural Society Alpine Show, London, UK

Shortia uniflora by M. Smith, from Curtis’s Botanical Magazine vol. 133: plate 8166 (1907) [374]

Diapensia himalaica, Yunnan, China [374]

Diapensia lapponica, Korkea Jehkas, Finnish Lapland (HV) [374]

Genera and species: Diapensiaceae include five genera with 12 species: Berneuxia (1), Diapensia (4), Galax (1), Pyxidanthera (1) and Shortia (5).

375. STYRACACEAE Storax family

Uses: The leaves of Galax urceolata (beetleweed) stay fresh for a long time and are gathered for the florist trade. It is also sometimes cultivated for this purpose. Other species are only rarely grown in specialist alpine collections. Etymology: Diapensia (Διαπένσια) is an ancient Greek name that was used for another unidentified but unrelated plant, probably from δια (dia), through, and πέντε (pente), five.

Evergreen and deciduous trees and shrubs comprise Styracaceae. Stellate or scale-like hairs are present on leaves, which are simple, alternate, petiolate and exstipulate. Blades have pinnate venation and entire, serrate

or rarely lobed margins. Inf lorescences are axillary (or falsely terminal) racemes or panicles, but f lowers are sometimes solitary. Flowers are usually bisexual and actinomorphic with a floral tube (hypanthium) surrounding the ovary. The four or five (rarely two to nine or absent) sepals are free or fused, sometimes forming a tube with minute apical teeth. The four or five (rarely to eight) petals are basally fused. Stamens are usually twice the number of petals (rarely equal the number or up to four times as many), and filaments are fused to the petal tube and flattened at the base, sometimes also forming a tube (corona). Anthers are basifixed and open inside by lengthwise slits. The partly to completely inferior ovary is composed of two to four Plants of the World

499

ERICALES

Sinojackia xylocarpa, Royal Botanic Gardens, Kew, UK [375]

EUDICOTS

Halesia carolina, Arboretum Poortbulten, De Lutte, the Netherlands [375]

carpels, fused to form a single locule. The single style is usually hollow and topped with a truncate or minutely lobed stigma. Fruits are loculicidal capsules, samaras, nuts or drupes. Distribution: This family occurs in the Neotropics and Asian tropics, but with some in temperate regions of North America (California, southeastern USA, Mexico), the West Indies, southern Europe, western Asia and in temperate East Asia (Himalayas, northern China, South Korea and Japan). Phylogeny and evolution: Before DNA studies, Styraceae were placed close to Ebenaceae, but molecular results have shown that they are sister to Diapensiaceae and together this pair are sister to Symplocaceae. Styracaceae have been shown to be monophyletic on the basis of morphological and DNA sequence analyses, and within the family two clades can be recognised, one composed of Styrax and Huodendron (the bulk of the species), with the other genera their sister. South American Pamphilia is now included in Styrax. The fossil record of Styracaceae extends back to the early Eocene, and diversification of the family has been estimated to have occurred c. 55 million years ago. It was once more widespread across the Northern Hemisphere, and the current occurrence in California, southern Europe and western Asia is relictual. Styrax and Halesia both occur in eastern North America and eastern Asia. 500

Christenhusz, Fay & Chase

Pterostyrax hispida, Helsinki Botanical Garden, Finland [375]

Genera and species: This family has 11 genera and c. 160 species: Alniphyllum (3), Bruinsmia (2), Halesia (2), Huodendron (4), Melliodendron (1), Parastyrax (2), Perkinsiodendron (1), Pterostyrax (4), Rehderodendron (5), Sinojackia (5) and Styrax (c. 130). Uses: Benzoin, a balsamic resin, is harvested commercially from several Asian species (Styrax benzoin, S. paralleloneurus and S. tonkinensis). It is used in the pharmaceutical, perfume and candy industries. Species of Halesia, Pterostyrax and Styrax are sometimes cultivated for their abundant white flowers in spring. Other genera are sometimes found in specialist collections. Etymology: Στύραξ (Styrax) is the Greek name for benzoin resin or storax, a resin used in classical times as a perfume, traditionally harvested from Liquidambar orientalis (Altingiaceae). Several Asian species of Styrax also yield such resin, but not S. officinalis native to Greece, so this is possibly a confusion of names.

376. SARRACENIACEAE American-pitcherplant family

Styrax japonica, Royal Botanic Gardens, Kew, UK [375]

Sarraceniaceae are perennial carnivorous herbs (rarely shrubs) with underground rhizomes and alternate rosette-forming, pitchershaped leaves without stipules. Leaves form hollow tubes with a dilated clasping petiole and variously coloured, blotched, spotted or striped blades. The pitcher often has wings or ridges along its length and a hood-like tip. The mouth has an outcurved rim that is thickened and smooth. The tube is minutely glandular and hairy inside. Linear or scale-like leaves are sometimes also formed early or late in the season. Inflorescences are terminal scapes bearing solitary flowers or in few-flowered racemes (Heliamphora). Flowers are bisexual and actinomorphic, usually nodding. The perianth is composed of a single petal-like whorl of four (to six) free tepals (Heliamphora) or two whorls with five free sepals and five free petals (Darlingtonia, Sarracenia). The ten to 20 or numerous (to 100) stamens are free or in fascicles, in Sarracenia placed on ten stalks. Filaments are short and anthers are basifixed or versatile and open laterally by lengthwise or pore-like slits on the anther appendages (Heliamphora). The superior ovary is composed of three (Heliamphora) or five fused carpels, each forming a usually incompletely separated locule. The terminal style is slightly trilobed with a truncate stigma (Heliamphora) with five short branches (Darlingtonia) or apically expanded into an umbrella-like structure with stigmas under each of the five tips (Sarracenia). Fruits are loculicidal capsules with numerous (up to 1,000), small, winged seeds.

ERICALES

Sinojackia xylocarpa, Royal Botanic Gardens, Kew, UK [375]

EUDICOTS

Halesia carolina, Arboretum Poortbulten, De Lutte, the Netherlands [375]

carpels, fused to form a single locule. The single style is usually hollow and topped with a truncate or minutely lobed stigma. Fruits are loculicidal capsules, samaras, nuts or drupes. Distribution: This family occurs in the Neotropics and Asian tropics, but with some in temperate regions of North America (California, southeastern USA, Mexico), the West Indies, southern Europe, western Asia and in temperate East Asia (Himalayas, northern China, South Korea and Japan). Phylogeny and evolution: Before DNA studies, Styraceae were placed close to Ebenaceae, but molecular results have shown that they are sister to Diapensiaceae and together this pair are sister to Symplocaceae. Styracaceae have been shown to be monophyletic on the basis of morphological and DNA sequence analyses, and within the family two clades can be recognised, one composed of Styrax and Huodendron (the bulk of the species), with the other genera their sister. South American Pamphilia is now included in Styrax. The fossil record of Styracaceae extends back to the early Eocene, and diversification of the family has been estimated to have occurred c. 55 million years ago. It was once more widespread across the Northern Hemisphere, and the current occurrence in California, southern Europe and western Asia is relictual. Styrax and Halesia both occur in eastern North America and eastern Asia. 500

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Pterostyrax hispida, Helsinki Botanical Garden, Finland [375]

Genera and species: This family has 11 genera and c. 160 species: Alniphyllum (3), Bruinsmia (2), Halesia (2), Huodendron (4), Melliodendron (1), Parastyrax (2), Perkinsiodendron (1), Pterostyrax (4), Rehderodendron (5), Sinojackia (5) and Styrax (c. 130). Uses: Benzoin, a balsamic resin, is harvested commercially from several Asian species (Styrax benzoin, S. paralleloneurus and S. tonkinensis). It is used in the pharmaceutical, perfume and candy industries. Species of Halesia, Pterostyrax and Styrax are sometimes cultivated for their abundant white flowers in spring. Other genera are sometimes found in specialist collections. Etymology: Στύραξ (Styrax) is the Greek name for benzoin resin or storax, a resin used in classical times as a perfume, traditionally harvested from Liquidambar orientalis (Altingiaceae). Several Asian species of Styrax also yield such resin, but not S. officinalis native to Greece, so this is possibly a confusion of names.

376. SARRACENIACEAE American-pitcherplant family

Styrax japonica, Royal Botanic Gardens, Kew, UK [375]

Sarraceniaceae are perennial carnivorous herbs (rarely shrubs) with underground rhizomes and alternate rosette-forming, pitchershaped leaves without stipules. Leaves form hollow tubes with a dilated clasping petiole and variously coloured, blotched, spotted or striped blades. The pitcher often has wings or ridges along its length and a hood-like tip. The mouth has an outcurved rim that is thickened and smooth. The tube is minutely glandular and hairy inside. Linear or scale-like leaves are sometimes also formed early or late in the season. Inflorescences are terminal scapes bearing solitary flowers or in few-flowered racemes (Heliamphora). Flowers are bisexual and actinomorphic, usually nodding. The perianth is composed of a single petal-like whorl of four (to six) free tepals (Heliamphora) or two whorls with five free sepals and five free petals (Darlingtonia, Sarracenia). The ten to 20 or numerous (to 100) stamens are free or in fascicles, in Sarracenia placed on ten stalks. Filaments are short and anthers are basifixed or versatile and open laterally by lengthwise or pore-like slits on the anther appendages (Heliamphora). The superior ovary is composed of three (Heliamphora) or five fused carpels, each forming a usually incompletely separated locule. The terminal style is slightly trilobed with a truncate stigma (Heliamphora) with five short branches (Darlingtonia) or apically expanded into an umbrella-like structure with stigmas under each of the five tips (Sarracenia). Fruits are loculicidal capsules with numerous (up to 1,000), small, winged seeds.

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Distribution: This is a family with a disjunct distribution. Sarracenia is found in eastern North America, extending from southern Canada and Newfoundland south to Florida. Darlingtonia occurs in northwestern North America (northern California and Oregon), and Heliamphora is restricted to table mountains (tepuis) in the Guayana Highlands northern South America. Phylogeny and evolution: Because of their derived morphology, relationships have been difficult to discern using traditional methods of classification. They have been suggested to be related to other carnivorous families such as Droseraceae and Nepenthaceae, but apart from gross morphology there are few similarities among these families. Floral characteristics are in agreement with Theaceae and Actinidiaceae, and indeed molecular studies have placed them in Ericales, among which they are found in a clade with Roridulaceae and Actinidiaceae. Evolutionary origins are obscure, and the fossil record is highly questionable, especially the allocation of 124 million year old Archaeamphora from China to this family. Some authors have suggested that Darlingtonia californica, Royal Botanic Gardens, Kew, UK [376]

Heliamphora may be primitive in the family because of its less complex pitcher structure and lack of digestive glands, although this is likely to be affected by specific adaptations to their habitats. Molecular analyses have placed Darlingtonia as sister to the rest of the family.

Uses: Several species and hybrids of Sarraceniaceae are grown as botanical curiosities as either indoor potplants or garden bog-plants. Sarracenia purpurea and perhaps some other species have become naturalised in Europe (e.g. Ireland, UK, the Netherlands, Switzerland and Sweden) and East Asia (Japan).

an insect will lose its footing and fall into the pitcher. Only the species of Sarracenia are full-f ledged carnivores, producing digestive enzymes in the pitcher liquid and absorbing released nutrients. Most species of Heliamphora rely on bacteria in the pitcher to digest prey, which is then absorbed, although at least one species of Heliamphora (the shrubby H. tatei) also produces enzymes. Some species of Sarracenia also produce nectar around the rim of the opening that is reputed to contain the narcotic substance coniine. Nectar production appears to be more important in attracting prey than does the colour of the pitchers. Species with a hood over the pitcher keep out rainwater, necessitating liquid production from within the pitchers and preventing f lying prey from escaping.

Carnivory: American pitcher plants have leaves modified into tubular pitfall traps (pitchers) that attract, catch and digest (most but not all) small invertebrate prey. They are characteristic of nutrient-poor, acid or ultramafic bogs. Their pitchers have no moving parts but have wax scales on the upper surfaces that increase the chances that

Etymology: Sarracenia is named for French physician Michel Sarrazin, Latinised as Sarracenus (1659–1734). He was a naturalist in Quebec, Canada, where he first discovered these plants and sent a specimen to Tournefort, who named the genus in his honour. Sarrazin is famed for his early works on the Canadian flora.

Genera and species: Sarraceniaceae include three genera with 34 species: Darlingtonia (1), Heliamphora (25) and Sarracenia (8).

Heliamphora nutans, Royal Botanic Gardens, Kew, UK [376]

Sarracenia minor, Chelsea Flower Show, UK

[376]

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377. RORIDULACEAE Flycatcher-bush family

EUDICOTS

outward toward the top with a capitate, erect, papillate stigma. The fruit is a three-valved, loculicidal capsule. Distribution: This family is restricted to the Cape Province of South Africa.

These are evergreen, insect-trapping shrubs with a taproot and few-branched stems. Leaves are alternate, crowded at the stem tips and spirally arranged. They lack stipules and petioles and are linear, with entire or shortly lobed margins. Blades are covered with stalked glands that exude a sweet sticky liquid that traps insects. Inflorescences are terminal few-flowered tight clusters. The bisexual flowers are actinomorphic. The five sepals and five petals are free, the petals showy, pink, purple or white. The five stamens are opposite the petals, the filaments free and anthers fixed dorsally near the base, incurved in bud and opening by four apical pores or short slits. Anthers have a basal swelling that produces nectar, and the stamens are initially pendent but when touched swing upwards, releasing pollen. The superior ovary is composed of three carpels, fused to each form a locule. The terminal style is straight or tapering

Roridula dentata, Hampton Court Flower Show, UK [377]

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Phylogeny and evolution: Roridulaceae have been variously associated with a number of families on the basis of gross morphology. Especially relationships with Droseraceae, Byblidaceae and Clethraceae have been suggested in the past. Molecular studies have shown that the family is not closely related to any of these, but is not far from Clethracae in Ericales, in which they are sister to Sarraceniaceae. Roridulaceae are estimated to have diverged c. 90 million years ago and are a relictual element of the Cape flora. Genera and species: The family has a single genus, Roridula, with two species: R. gorgonias and R. dentata. Carnivory: The leaves of Roridula resemble those of the sundews (Drosera) in being covered with long glandular hairs and excreting a sticky liquid that traps insects. Despite their morphological resemblance to sundews, they lack the specialised digestive and absorptive glands on their leaves that other carnivorous plants have to digest prey and take up the released nutrients, which Roridula gorgonias, Royal Botanic Garden, Kew, UK [377]

suggests that these are not fully carnivorous species. However, two species of Hemiptera (true bugs), Pameridea roridulae and P. marlothii, are closely associated with Roridula, living their entire lives on these plants without getting trapped. They eat the dead insects and occasionally drink sap if insects are scarce. It is in fact the faeces of these bugs that provide these plants with nutrients, probably mostly through their roots but probably also directly across their thin leaf cuticle. This mutualist relationship between a plant and an insect is unique. Pameridea bugs benefit from the trapped prey, and Roridula benefits by taking up nutrients from droppings at their roots with their unusually shaped cuticular caps. The population density of Pameridea bugs on plants of Roridula is negatively correlated with the density of a spider, Synaema marlothii, a species that feeds on both trapped prey and Pameridea. Pameridea also sucks the sap from Roridula, which reduces the benefit of the nutrients when the population of Pameridea becomes too large. Synaema marlothii is important in this mutualistic triangle to stabilise the populations of Pameridea and maintain the balance. Roridula has been termed a protocarnivore by some. Etymology: Roridula is a diminutive of the Latin roridus, dewy, in reference to the sticky exudate of the glands on the leaves. Capsid bug Pameridea roridulae on Roridula gorgonias, Kogelberg Nature Reserve, Cape Province, South Africa (CD) [377]

ERICALES

EUDICOTS

Actinidia chinensis ‘Tonshan’, Wuhan Botanical Garden, China (CD) [378]

Actinidia deliciosa, University of British Columbia Botanical Garden, Vancouver, Canada (CD) [378]

Actinidia kolomikta, Helsinki Botanical Garden, Finland [378]

Saurauia madrensis, University of California Botanical Garden, Berkeley, USA [378]

Clematoclethra lasioclada, National Botanic Gardens of Ireland, Glasnevin [378]

378. ACTINIDIACEAE

free and often cup-shaped. The usually five (sometimes three to nine) petals are free or more or less fused at the base. The ten, 20 or indefinite (to about 240) stamens are free or fused to the petal bases. Anthers are dorsifixed and open outside by lengthwise slits or near the tip by pores. There is no nectar disk. A superior ovary is composed of three, five or many carpels, fused into many locules. Styles are as many as carpels and free and radiating or all fused into a single style. Fruits are usually berries or occasionally a leathery, dehiscent capsule.

in temperate and subtropical East Asia, and Clematoclethra is endemic to China. Saurauia is widespread.

Kiwifruit family

This family consists of woody vines, trees and shrubs with alternate, simple, petiolate leaves without stipules. Blades have pinnate venation, and margins are toothed, serrate or finely serrate. Inflorescences are axillary cymes or paniculate thyrses, sometimes reduced to solitary flowers. The pedicellate actinomorphic f lowers are bisexual or unisexual, sometimes only functionally so. The five sepals (rarely three to eight) are

Distribution: Actinidiaceae are found in tropical and subtropical America (Mexico, Mesoamerica, northwestern South America) and East Asia from the Russian Far East and Japan throughout China, Korea, the Himalayas, tropical East Asia and Malesia to New Guinea, Melanesia, northeastern Australia and Fiji. Actinidia is only found

Phylogeny and evolution: A family previously associated with Dilleniaceae or Theaceae, they are now understood to belong to Ericales on the basis of embryology, floral characters and molecular studies, the last placing them in a clade with the carnivorous Sarraceniaceae and Roridulaceae, although support for this relationship is not strong. Late Cretaceous fossil flowers attributed to Saurauia are known from the eastern USA, c. 80 million years old, and leaves similar to Saurauia are known from Eocene fossil beds in North America. Fossil Actinidia-like leaves are known from the Tertiary of Japan. Fossil Saurauia and Actinidia seeds have been found in abundance in Europe from the Maastrichtian onwards, showing that the current distribution of the family is likely to be relictual. Plants of the World

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Genera are still taxonomically poorly studied, and numbers of species given below are therefore rough estimates. The 25 species of Clematoclethra are sometimes treated under a single name, C. scandens. Genera and species: This family includes three genera with c. 360 species: Actinida (36), Clematoclethra (c. 25) and Saurauia (c. 300). Uses: Kiwifruit (Actinidia deliciosa) originated in central China, and its original name was Chinese gooseberry. It was introduced in 1904 to New Zealand, a single large-fruited clone called ‘Hayward’ being the most common. After 1949 (and the Chinese Communist revolution) it became difficult to market them under the name Chinese gooseberry because anything Chinese was unpopular in the west at the time. The name kiwifruit was coined in New Zealand after the resemblance to the flightless bird found in that country. In China, four main cultivars are grown (particularly along the Yangtze River), whereas in New Zealand, six others are popular. A yellow variety with smooth skin, marketed as ‘Kiwi Gold’, is a selection from A. chinensis ‘Hort 16 A’, which is sweeter and less acidic and finds greater popularity. Other species with edible fruit but of minor economic importance are kokuwa or hardy kiwi (A. arguta), wild kiwi (A. chinensis) and silver vine (A. polygama), some of which have been tried as commercial crops, notably in

Clethra arborea, Royal Botanic Gardens, Kew, UK [379]

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Soviet Russia, because of their high vitamin C content. Several Actinidia species (e.g. A. kolomikta, A. polygama) are attractive to cats and have a euphoric effect on these animals. They are often used as a cat treat in Asia. Some Actinidia species are grown as ornamental vines, especially A. kolomikta, of which the male plants naturally have white and pink blotches on their leaves. Crying flowers: In the lowlands along river banks on the island of Bougainville (Papua New Guinea) the native people reported that the flowers of karakara or tokitoki, identified as Saurauia purgans (a synonym of S. schumanniana), make ‘a peculiar crying sound when opening’. This report from 1945 has never been reconfirmed, but it may be worth investigating. Etymology: Actinidia is derived from the Greek ακτινος (aktinos), a ray, in reference to the radiating styles.

379. CLETHRACEAE Lily-of-the-valley-tree family

Clethra arborea in fruit, Royal Botanic Gardens, Kew, UK [379]

Clethraceae are a family of shrubs and trees with deciduous and evergreen, simple, petiolate alternate leaves without stipules. Blades have pinnate venation and toothed to serrate margins, the teeth sometimes glandular. Inflorescences are terminal or axillary simple racemes or racemes clustered in panicles, fascicles or umbels. Flowers are usually bisexual and actinomorphic, rarely slightly zygomorphic. The five (rarely six) sepals are free or fused up to three quarters, equal in size or the outer sometimes larger. The five (seldom six) petals are free or basally fused (up to to half) and then the corolla urn-shaped. The ten (or 12) stamens are in two whorls and free or slightly fused at the base to the petals. Anthers are ventrifixed, swinging when a flower opens, eventually opening by apical pore-like slits. There is sometimes a nectar disk surrounding the ovary. The superior ovary is composed of three or five fused carpels, each forming a locule. A simple, hollow style is topped by three or five linear stigmas. Fruits are indehiscent or loculicidally opening capsules enclosed by a persistent calyx. Distribution: The family has an unusual disjunct distribution, with a centre of diversity in tropical America extending into southeastern Brazil and another in the Malesian Archipelago and warm temperate East Asia (Indochina, China, South Korea and southern Japan). There are also two species in eastern North America and one species on Madeira.

Clethra alnifolia, New York Botanic Garden, USA [379]

Clethra barbinervis in fruit, East Bergholt Place Arboretum, USA [379]

ERICALES

EUDICOTS

Phylogeny and evolution: Clethraceae were usually considered to include only Clethra, but molecular studies have shown that Purdiaea, a Neotropical genus with its greatest diversity in Cuba previously included in Cyrillaceae, is better placed in Clethraceae. Clethraceae have been generally accepted as part of Ericales, in which they are sister to Cyrillaceae plus Ericaceae. The fossil record of Clethraceae extends back to the Middle Eocene in Europe, Japan and North America, suggesting that the current distribution is relictual. The clade including Clethraceae, Cyrillaceae and Ericaceae has been estimated to have evolved c. 58 million years ago. Genera and species: Clethraceae include two genera with c. 95 species: Clethra (83) and Purdiaea (12). Uses: The leaves of Clethra barbinervis are eaten in China. Timber of some larger species is used to make furniture in tropical America. Only a few species are grown as garden ornamentals, mainly the North American bush pepper, C. alnifolia, and the Madeiran lily-ofthe-valley tree, C. arborea, because of their scented flowers. Etymology: Clethra is the Latinised form of Greek κλήθρα (klethra), an alder, in reference to the similar leaves.

380. CYRILLACEAE Leatherwood family

This family includes evergreen and sometimes deciduous shrubs and trees with alternate, simple leaves without stipules crowded at the ends of branches. Leaf blades can be petiolate or sessile and have entire margins with a pinnate venation.

Cyrilla racemiflora var. parvifolia, Florida, USA

Cyrilla racemiflora, Green Swamp, North Carolina, USA [380]

Inf lorescences are terminal or axillary racemes. Flowers are subtended by paired small leaves (prophylls) on the pedicels and are actinomorphic and bisexual. The five (sometimes four to eight) sepals are fused at least at the base, the outer are larger than the inner ones with the calyx persistent in fruit. Petals are as many as sepals (usually five) and free, the petals glandular in Cyrilla. The five (Cyrilla) or ten (Cliftonia) free stamens are broader at the base in Cliftonia and become petal-like in the lower half; the stamens of Cyrilla are round and subulate. Anthers are dorsifixed, not inverted when the flower opens (compare Clethraceae), and the thecae open on the inside by longitudinal slits. A circular nectar disk surrounds the ovary base. The superior ovary is composed of two to fived fused carpels, each forming a locule and topped with a single hollow style and a twoto five-lobed stigma, or with as many short style branches as there are carpels. Fruits are single-seeded, dry and drupe-like or samaras.

Central America and northern South America. Cyrilla is mostly found on acid, sandy or peaty soils along water bodies or in bogs, wet prairies, pine barrens and tropical cloud forests. Cliftonia is restricted to southeastern North America from Louisiana to Florida, where it grows in habitats similar to those of Cyrilla.

(WJ) [380]

Distribution: Cyrillaceae are restricted to warm-temperate and tropical regions of the Americas, in the coastal regions from southeastern Virginia to southeastern Texas, southern Mexico (Oaxaca), the Caribbean,

Phylogeny and evolution: On morphological and embryological grounds, Cyrillaceae have long been considered closely allied to Ericaceae, a placement that was confirmed by molecular studies. These have placed Cyrillaceae (excluding Purdiaea) sister to Ericaceae. The original delimitation between Cyrillaceae and Clethraceae was based on the type of fruit, which resulted in Purdiaea being placed in the former. Molecular analyses have shown that this genus is closer to Clethra, which agrees with the general morphology of these plants. Abundant fossil wood and pollen of Cyrillaceae were found in Upper Oligocene formations in Vermont. Similar fossils are known from Tertiary deposits in Germany. Late Cretaceous fossil fruits from Europe attributed to Cyrillaceae may refer to Clethraceae, the two being difficult to distinguish when fossilised.

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ERICALES Genera and species: There are two genera in the family, each with a single species: Cliftonia monophylla and Cyrilla racemiflora.

EUDICOTS

381. ERICACEAE Heather family

Uses: Both species are occasionally grown as garden ornamentals in specialist collections. Etymology: Cyrilla is named in honour of Italian physician Domenico Maria Leone Cirillo (1739–1799), who was physician to King Ferdinand IV of Naples and an acclaimed botanist and entomologist. He was caught up in political turmoil between royalists and French Parthenopean republicans in Naples who seized power for six months in 1799. Together with other republicans, he was executed after the King returned to power.

Ericaceae include evergreen and deciduous shrubs, trees, vines, creepers and perennial herbs. They can be epiphytic, epilithic or terrestrial and often make associations with mycorrhizal fungi, sometimes lacking chlorophyll (mycoheterotrophic). Leaves are

Enkianthus chinensis, Mt Huangshan, Anhui, China [381]

Monotropa uniflora in fruit, Wisconsin, USA [381]

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Pterospora andromedea, New Mexico, USA (DZ) [381]

simple and alternate, opposite or whorled, often clustered at the tips of branches or spirally arranged along the stem, usually without stipules. Blades are entire or serrate at the margins, usually with pinnate venation, sometimes achlorophyllous and scale-like, or with the margins recurved around the midvein and the leaf needle-like (ericoid). Inf lorescences are terminal or axillary racemes, usually with prophylls and sometimes bracteoles, or flowers can be solitary. Flowers are usually bisexual, sometimes unisexual, and actinomorphic or zygomorphic. The four or five (rarely two to seven) sepals are basally fused. The four to five (or three to seven) petals are fused, rarely free or forming a cap; usually the petals form a bell-, cup- or urn-shaped corolla. The usually five or ten (sometimes two Empetrum nigrum, Punkaharju, Finland [381]

Hypopitys monotropa, red autumn form, North Carolina, USA [381]

ERICALES

EUDICOTS

Chimaphila umbellata, Old Mission Peninsula, Michigan, USA [381]

Orthilia secunda in fruit, Punkaharju, Finland [381]

Arctostaphylos manzanita, Rancho Santa Ana Botanical Garden, California, USA [381]

to 16) stamens are typically free from the petal tube (rarely fused with it). Anthers are variable in shape and ornamentation but are usually dorsifixed or nearly basifixed, inverting during their development, often with two appendages or a protruding connective and thecae opening on the inside or at the tip with pores or short to long slits. A nectary is often present inside the stamen whorl(s) around the ovary. The superior to inferior ovary is composed of usually four or five (rarely up to 12) fused carpels topped with a single, hollow style and a punctate or lobed stigma. Fruits are berries, drupes or capsules. Distribution: Ericaceae have a nearly worldwide distribution, but they occur mostly in the northern and southern temperate and subtropical zones, extending north into the Arctic and south into the sub-Antarctic

Arctostaphylos uva-ursi, Nauvo, Finland [381]

islands. They are absent from the great deserts and sparse or absent from the American and African lowland tropics. There are species radiations in Mesoamerica and the Andes, South Africa (Erica), the Himalayas and East Asia (especially Rhododendron), Malesia and Australia (Styphelioideae). Phylogeny and evolution: Paleoenkianthus is a c. 90-million-year-old (Late Cretaceous) fossil, which has a bee-pollinated flower, similar to many modern Ericaceae. Age estimates based on fossils and molecular results are often in disagreement, depending on the taxa included in the analysis. They have been a dominant element of the heaths and tundra in the Pleistocene ice ages, probably benefitting from the evolution of large grazing mammals, although Ericaceae have evolved to

Pyrola picta, New Mexico, USA (DZ) [381]

Arbutus unedo, Kos, Greece (AP) [381]

inhabit a great number of niches, varying from rainforest canopies, mountain slopes and moist broadleaf forests to seasonally dry, sandy scrubland. Their current distribution may have some relictual and vicariant elements, and long-distance dispersal is common, which can be observed, for example, in the bipolar distribution of Empetrum. Previously Empetraceae, Epacridaceae, Pyrolaceae and Monotropaceae were recognised, but they are all closely related to traditional Ericaceae, with Enkianthus sister to the rest. Therefore, seven subfamilies are now accepted, with Empetraceae placed in Ericoideae and former Epacridaceae treated as Styphelioideae. Also Cassiope and Harrimanella are sometimes given their own subfamilies, depending on their placement in the phylogenetic analysis. Plants of the World

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Kalmia polifolia, Helsinki Rhododendron catawbiense, North Botanical Garden, Finland [381] Carolina, USA [381]

Erica cinerea, Wales, UK [381]

Astroloma cf. ciliatum (Christenhusz 6350), Mt Benia, Western Australia [381]

Cassiope selaginoides, Yunnan, China [381]

Epacris longiflora, Australian National Botanic Gardens, Canberra [381]

Vaccinium corymbosum, Royal Botanic Gardens, Kew, UK [381]

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Rhododendron japonicum, Royal Botanic Kalmia buxifolia, North Carolina, USA [381] Gardens Edinburgh at Dawyck, UK [381]

Erica canaliculata, Royal Botanic Gardens, Kew, UK [381]

Richea dracophylla, Australian National Andersonia lehmanniana, Mt Botanic Gardens, Canberra [381] Benia, Western Australia [381]

Agapetes serpens, Royal Botanic Gardens, Edinburgh, UK [381]

Agarista buxifolia, Réunion [381]

ERICALES

EUDICOTS

Molecular phylogenetics has caused some generic reorganisations, such as the merger of Azalea, Ledum and Menziesia with Rhododendron, Leiophyllum and Loiseleuria in Kalmia, the inclusion of Agarista in Agauria, the designation of Pernettya as a synonym of Gaultheria, and the expansion of Erica to include many more genera. On the other hand, Arctostaphylos is split to accept Arctoüs and Xylococcus, Hypopitys is separated from Monotropa, and Dielsiodoxa is segregated from Monotoca. Leucopogon was found to be polyphyletic and hence several genera were accepted as segregates. There are still many questions related to generic circumscription, and it is likely that changes will be made in the future when the DNA of more species from large genera have been sequenced. Pseudogonocalyx paradoxa, previously placed here, is a species of Schoepfia (Schoepfiaceae, Santalales). Genera and species: Ericaceae are a family with 126 genera and 4,250 species in seven subfamilies: Enkianthoideae – Enkianthus (17); Monotropoideae –Allotropa (1), Cheilotheca (2), Hemitomes (1), Hypopitys (1), Monotropa (1), Monotropastrum (2), Monotropsis (1), Pityopus (1), Pleuricospora (1), Pterospora (1) and Sarcodes (1); Pyroloideae – Chimaphila (5), Moneses (2), Orthilia (1) and Pyrola (35); Arbutoideae – Arbutus (10), Arctostaphylos (c. 55), Arctoüs (5), Comarostaphylis (10), Ornithostaphylos (1) and Xylococcus (1); Ericoideae – Bejaria (16), Bryanthus (1), Calluna (1), Cassiope (12), Ceratiola (1), Corema (2), Daboecia (1), Diplarche (2), Elliottia (4), Empetrum (2), Epigaea (3), Erica (c. 860), Harrimanella (2), Kalmia (11), Kalmiopsis (1), Ledothamnus (7), Phyllodoce (7), Rhododendron (c. 1,000), Rhodothamnus (2) and Therorhodion (2); Styphelioideae – Acrothamnus (6), Acrotriche (15), Agiortia (3), Andersonia (c. 35), Androstoma (1), Archeria (7), Astroloma (28), Brachyloma (7), Budawangia (1), Coleanthera (3), Conostephium (6), Cosmelia (1), Croninia (1), Cyathodes (3), Cyathopsis (1), Decatoca (1), Dielsiodoxa (5), Dracophyllum (c. 48), Epacris (c. 40), Lebetanthus (1), Leptecophylla (13), Leucopogon (c. 230), Lissanthe (9),

Lysinema (5), Melichrus (4), Monotoca (17), Montitega (1), Needhamiella (1), Oligarrhena (1), Pentachondra (5), Planocarpa (3), Prionotes (1), Richea (11), Rupicola (4), Sphenotoma (6), Sprengelia (4), Styphelia (15), Symphysia (15), Trochocarpa (12) and Woollsia (1); Vaccinioideae – Agapetes (c. 400), Agarista (30), Andromeda (1), Anthopteropsis (1), Anthopterus (11), Cavendishia (c. 130), Ceratostema (23), Chamaedaphne (1), Costera (9), Craibiodendron (5), Demosthenesia (11), Didonica (4), Dimorphanthera (75), Diogenesia (13), Diplycosia (c. 100), Disterigma (35), Eubotrys (2), Gaultheria (134), Gaylussacia (49), Gonocalyx (>9), Lateropora (3), Leucothoë (6), Lyonia (35), Macleania (37), Mycerinus (5), Notopora (5), Oreanthes (7), Orthaea (35), Oxydendrum (1), Paphia (26), Pellegrinia (5), Pieris (7), Plutarchia (11), Polyclita (1), Psammisia (c. 60), Rusbya (1), Satyria (25), Semiramisia (4), Siphonandra (2), Sphyrospermum (22), Tepuia (7), Themistoclesia (25), Thibaudia (60), Utleya (1), Vaccinium (c. 140) and Zenobia (1). Uses: Blueberries (highbush blueberry, Vaccinium corymbosum cultivars and hybrids with other Vaccinium species) and American cranberries (V. macrocarpon) are the most widely grown fruit crops. Additional minor fruit crops, usually gathered from the wild, sometimes cultivated, are lowbush blueberry (V. angustifolium), New Jersey blueberry (V. caesariense), evergreen blueberry (V. darrowii), Cascade bilberry (V. deliciosum), Andean blueberry or mortiño (V. floribundum), square-twig blueberry (V. membranaceum), Canadian blueberry (V. myrtilloides), billberry or European blueberry (V. myrtillus), Alaskan blueberry (V. ovalifolium), wild cranberry (V. oxycoccus), red huckleberry (V. parvifolium), deerberry (V. stamineum), bog bilberry (V. uliginosum), rabbit-eye blueberry (V. virgatum), lingon berry (V. vitis-idaea) and many others. Several other genera of Ericaceae also have edible fruits that can be welcome forest snacks during hikes, but care has to be taken because there are a few that are poisonous (e.g. Agarista salicifolia, Comarostaphylis discolor and Gaultheria myrsinoides). Acrotriche

depressa, called native currant in Australia, is used in chocolates and other confectionery, and Astroloma humifusum also has sweet edible berries. Berries of the strawberry tree (Arbutus unedo) are sometimes eaten in the Mediterranean and used in jams, drinks and liqueurs such as medronho (a type of Portuguese brandy). Wintergreen (Gaultheria procumbens) is the source of a flavouring taken from the leaves, which after fermentation for three days yield an essential oil used in drinks, chewing gum and ice-cream. The leaves can be used to make a tea and were an important source of salicylic acid before the chemical synthesis of aspirin was developed. Berries of several species of Gaultheria (G. procumbens, G. shallon) can be eaten fresh or in preserves. The leaves of Labrador tea (Rhododendron tomentosum, formerly Ledum palustre) can be made into a herbal tea and were used in brewing beer during the Middle Ages. It also deters clothes moth and can be placed in closets for that purpose. Juice of crowberry (Empetrum nigrum) is sometimes made and is said to have a beneficial effect on the kidneys. The berries can be eaten fresh but have a watery taste. Fruits of several species in other genera (e.g. Lyonia, Macleania etc.) are also eaten locally. Common heather (Calluna vulgaris) shoots have been used for brooms, and formerly were the source of a yellow dye for wool. It has also been used for millennia in Scotland as a replacement for hops when brewing beer. Berries of cranberry (Vaccinium macrocarpon) and stems of Gaultheria shallon and Kalmia latifolia are frequently used for ornament in flower arrangements. Ericaceae are popular ornamental plants in temperate gardens, with Rhododendron species topping the list. Numerous species, selections, cultivars and hybrids are offered for sale, especially the Japanese or kurume azaleas (R. kaempferi × R. kiusianum), which are popular pot plants in many parts of the world. In addition to Rhododendron, species and cultivars of Agapetes, Andromeda, Astroloma, Calluna, Cassiope, Daboecia, Enkianthus, Erica, Gaultheria, Gaylussacia,

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ERICALES

Cavendishia micayensis, New York Botanical Garden [381]

Gaultheria shallon, Sea Ranch, California, USA

Mitrastemon matudae, Tres Picos, Chiapas, Mexico (CD) [382]

Kalmia, Leucothoë, Pieris, Vaccinium and Zenobia are often cultivated in gardens. Some Styphelioideae have become popular as ornamentals in Australia and New Zealand.

botanists all survived the ordeal and were rescued a couple of days later.

The superior ovary has a single locule and is conical with a groove below the style, which is only slightly constricted and topped with a thick conical slightly bilobed stigma. Flowers produce nectar that collects in the scale leaves to attract pollinators. The fruit is a fleshy woody capsule that opens along a horizontal slit between the style and the ovary, exposing the sticky seeds.

Frisian cranberries: Cranberries (Vaccinium macrocarpon) were traditionally imported from North America into Europe, where a sauce of the berries accompanied game, especially at Christmas. A story goes that several barrels of berries had blown off a ship in the North Sea during an autumn storm, and these washed ashore on the Frisian island of Terschelling. There a beachcomber, Mr. Cupido, found them and thinking it was wine, pulled the barrels up into the dunes. When he realised they were berries and not wine, he threw them into the boggy dune valley, where they naturalised. Today this little island has a thriving cranberry economy, making everything from sauce and chutney to wine. Lost in the world of the poisonous blueberry: On an expedition to the Venezuelan tepuis in the 1980s, several botanists were stranded due to a helicopter malfunction and ate local blueberries (a variety of Vaccinium puberulum). This caused a sudden drop in blood pressure, nausea, vomiting and finally loss of consciouness in those who consumed the fruit. The ones that did not noted down the interesting effect on the sufferers. Similar symptoms are known from species of Agarista, Kalmia and Rhododendron, caused by andromedotoxin, but this compound was not found in typical Venezuelan V. puberulum, which has edible fruit. It is the only species of Vaccinium that is known to be toxic. The 510

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[381]

Etymology: Erica is the classical Latin name for heather, probably from Ancient Greek ερεικε (ereike, heather). It is also sometimes used as a name for a woman, but in that case it may alternatively be the feminine form of the Old Norse name Erik.

382. MITRASTEMONACEAE Nippledaisy family

These are mushroom-like root endoparasites without chlorophyll that often cause ‘witches brooms’ to form and when flowering may create a ‘fairy ring’ around the host tree. Stems are confined to the xylem of the host root tissue but eventually break out and completely surround the roots when older. Leaves are opposite, sessile, cream-coloured scales subtending the single-f lowered inflorescence, becoming larger towards the apex of the latter. The bisexual flowers are actinomorphic and have a reduced, lobed, collar-shaped, fused perianth. Stamens are fused into a tube (androphore) that has a fertile zone at the tip, composed of cavities that bear pollen in several vertical rings. The tube is pushed open by the developing gynoecium.

Distribution: The family is disjunct between Central America–Colombia and tropical Asia (Sumatra, Borneo, New Guinea, Indochina, Taiwan and Japan). They grow exclusively on the roots of Fagaceae (i.e. Castanopsis, Lithocarpus, Quercus and Trigonobalanus). Phylogeny and evolution: Because they are endoparasites with little visible vegetative tissue, the family was traditionally included in Rafflesiaceae, but molecular studies place these odd plants in Ericales, although their closest relatives in this order are not yet certain. They were found to be early divergents in Ericales, and thus their placement here close to Ericaceae is not likely to be correct. Genera and species: The single genus Mitrastemon includes two species: M. matudae in Mesoamerica and M. yamamotoi in East Asia. Etymology: Mitrastemon is composed of the Greek words μήτρα (mitra), a mitre or cap, and στήμονα (stemona), a stamen or thread. The generic name was originally spelled Mitrostemma, but it was later corrected by the original author as Mitrostemon, which is now the conserved spelling.

ICACINALES

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ICACINALES Families 383 and 384 form the order Icacinales. Together with Metteniusales and Garryales they form a grade leading up to the rest of the lamiid (asterid I) clade. Icacinaceae, previously unplaced to order, have been found to be polyphyletic, with some members included in Cardiopteridaceae and Stemonuraceae (Aquifoliales) and Pennantiaceae (Apiales), and the former Apodytaceae and Emmotaceae being merged with formerly monogeneric Metteniusaceae (Metteniusales). This association has only recently been discovered and no age estimates are yet available for this assemblage of families. These tropical trees and vines are difficult to identify, and their putative synapomorphies may be plesiomorphic characters that cannot be used to distinguish these clades.

383. ONCOTHECACEAE Kanak-laurel family

This family includes rounded, evergreen trees or shrubs. The leathery, simple leaves are alternate, spirally arranged and crowded at the branch tips. They are petiolate or almost sessile and lack stipules. Blades are pinnately veined and have entire margins that sometimes

Oncotheca humboldtiana, New Caledonia (JM) [383]

have minute, glandular teeth near the leaf tip. Inflorescences are axillary thyrses composed of bracteate cymes with short pedicels (the flowers nearly sessile). The minute flowers (c. 2 mm across) are bisexual and actinomorphic. The five sepals are persistent in fruit, more or less equal and free to the base. The five petals are fused and form a short cup. The five stamens are fused to the petals by their filaments and are placed between the petal lobes. The basifixed anthers have a thick connective and open by lengthwise slits toward the outside. The connective has an apical appendage in Oncotheca balansae. The superior ovary is composed of five fused carpels, each forming a locule. The ovary is topped with five free styles. Fruits are fleshy drupes with a large seed and thin flesh.

Oncotheca humboldtiana, New Caledonia (JM) [383]

Distribution: This family is endemic to New Caledonia. Phylogeny and evolution: Oncothecaceae has been variously placed, often in Theales by former authors, but they have been placed in Icacinales in the molecular studies and appear to be most closely related to Icacinaceae and could perhaps be merged with that family. Genera and species: The sole genus of this family is Oncotheca, with two species: O. balansae and O. humboldtiana. Etymology: Oncotheca is composed of the Greek words onkos, a tumor, and theca, a covering or in botany the anther of a stamen, referring to the thickened connectives.

Oncotheca balansae, New Caledonia (JM) [383]

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ICACINALES

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Phytocrene bracteata, female flowers, Java, Indonesia (MN) [384]

Cassinopsis tinifolia, Umtamvuna Nature Reserve, Eastern Cape, South Africa (CD) [384]

Icacina mannii by W. Fitch, Curtis’s Botanical Magazine, vol. 102: plate 6260 (1876) [384]

Phytocrene bracteata, male flowers, Singapore

Iodes ovalis, Singapore (WA) [384]

Pyrenacantha malvifolia, Royal Botanic Gardens, Kew, UK [384]

384. ICACINACEAE

stipules. Blades are entire or palmately lobed, sometimes toothed, and venation is pinnate or reticulate, or palmate in some climbing taxa (e.g. Hosiea). Inflorescences are axillary or leaf-opposed cymes, panicles, racemes, spikes, heads, or flowers can be solitary, often bracteate. Flowers are actinomorphic and usually bisexual, rarely zygomorphic or unisexual. The three to five (sometimes absent) sepals are fused to form a tube with blunt lobes, sometimes free, and persistent in fruit. Petals are as many as sepals (usually four or five, rarely absent), basally fused, rarely free. Stamens are the same number as sepals and alternate with the petal lobes. Filaments are free or fused basally to the petals. Anthers are dorsifixed and open inside by lengthwise slits. Female flowers often have

staminodes. The superior ovary is composed of three (rarely two, four or five) fused carpels each making a locule or all fused into a single locule. The simple style is apical and topped by as many stigmas as carpels. Fruits are fleshy or dry drupes.

(WA) [384]

False-yam family

This is a family of trees, shrubs and vines that climb by stem tendrils, often with large underground, woody rootstocks. Leaves are simple, usually alternate and spirally arranged, rarely opposite (Iodes), mostly petiolate and without 512

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Distribution: A pantropical family found in the American, African and Asian tropics, Madagascar, northern Australia and on Pacific islands. Phylogeny and evolution: A crown age of 115 million years has been estimated. In the Eocene, Icacinaceae were widespread and diverse in the Northern Hemiphere, with genera now known only from tropical

METTENIUSALES

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Asia found in fossil beds of North America, suggesting a boreotropical hypothesis. The family has previously been difficult to place due to the lack of obvious distinctive characteristics, and as traditionally circumscribed Iacacinaceae were found to be polyphyletic. They included genera that are now placed elsewhere, in Cardiopteridaceae and Stemonuraceae (Aquifoliales) and Pennantiaceae (Apiales). Former Apodytaceae and Emmotaceae, often included in Icacinaceae, are correctly placed in Metteniusaceae with which they form a clade. Cassinopsis is an oddball in this family, with its spiny stems and imbricate (instead of valvate) petals and is tentatively placed here.

Chlamydocarya and Polycephalium belong to Pyrenacantha, and Polyporandra is merged with Iodes. Genera and species: Icacinaceae in the strict sense include 25 genera and c. 150 species: Alsodeiopsis (11), Casimirella (7), Cassinopsis (6), Desmostachys (7), Hosiea (2), Icacina (6), Iodes (29), Lavigeria (1), Leretia (1), Mappia (4), Mappianthus (2), Melliodendron (1), Merrilliodendron (1), Miquelia (8), Natsiatopsis (1), Natsiatum (1), Nothapodytes (5), Phytocrene (11), Pittosporopsis (1), Pleurisanthes (6), Pyrenacantha (29), Rhyticaryum (12), Sarcostigma (2), Sleumeria (1) and Stachyanthus (6).

Uses: Tubers and seeds of false yam (Icacina oliviformis) can be ground into an edible flour, eaten locally in Africa. Leaves of Iodes scandens are cooked with taro in Melanesia. The fatty acids from the tubers of some species of Casimirella can be used to relieve pain. Stems of Miquelia vines produce drinking water when cut. Several species are locally harvested for timber. Etymology: Icacina was named because of its vegetative resemblance to Chrysobalanus icaco (Chrysobalanaceae). Icaco in turn is derived from ikaku, the Arawak name for coco plum.

METTENIUSALES This is a recently recognised order, which now include a number of genera formerly placed in Icacinaceae. Fossil pollen is known from the Palaeocene of Greenland and Arkansas.

385. METTENIUSACEAE Urupagua family

Poraqueiba sericea, Rio Tigre, Nauta, Loreto, Peru (CD) [385]

This is a family of evergreen trees and shrubs (climbing in some Rhaphiostylis) with simple, alternate, petiolate leaves arranged spirally along the stem, lacking stipules. Margins are entire, sometimes toothed (Calatola), with a pinnate venation. Inflorescences are bracteate, axillary cymes, racemes, panicles, fascicles, spikes or compound thyrses. Flowers are bisexual, rarely unisexual, and actinomorphic (but zygomorphic in female f lowers of Calatola). The five (or four) sepals are free or slightly fused at the base

Metteniusa tessmanniana, Selva Bananito, Limon, Costa Rica (CD) [385]

Apodytes dimidiata, Karatara River, Western Cape, South Africa (CD) [385]

and persistent or deciduous in fruit. The five (or four) petals are free, fused at the base or completely fused and often fleshy or sometimes hairy outside (Metteniusa). The five (or four) stamens alternate with the petals and are attached to them or are free and elongate (Metteniusa), sometimes hairy. Anthers are dorsifixed or basifixed, in Metteniusa they are multilocular, with the thecae in four rows opening by slits individually and broadly recurving to form a circle. Connectives are often large. The

Apodytes abbottii, Umtamvuna Nature Reserva, Eastern Cape, South Africa (CD) [385]

Emmotum nitens, Chapada Diamantina, Bahía Brazil [385]

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METTENIUSALES superior ovary is composed of fused carpels, usually with a single locule, in Emmotum in two or three locules. Metteniusa has four reduced carpels forming a zygomorphic ovary. Styles are long or short, topped with punctate, capitate, lobed or oblique stigmas. Fruits are drupes, sometimes asymmetrical, ribbed or samara-like. Distribution: A pantropical family. Phylogeny and evolution: Morphologically, Metteniusa fits well with Cardiopteridaceae (Aquifoliales), although the f lowers have similarities to members of Cornales. The

EUDICOTS

combination of entire leaves without stipules and pentalacunar nodes is unusual, and this resulted in the placement of Metteniusa in its own family. Molecular studies have placed Metteniusa with several former members of Icacinaceae, most notably Apodytaceae and Emmotaceae, and these are now combined to form a single family. Genera and species: Metteniusaceae includes 10 genera and c. 57 species: Apodytes (c. 6), Calatola (7), Dendrobangia (3), Emmotum (c. 10), Metteniusa (7), Oecopetalum (3), Ottoschultzia (3), Platea (5), Poraqueiba (3) and Rhaphiostylis (c. 10).

Uses: The nuts of Metteniusa edulis are edible after cooking in salt water. Fruits of Poraqueiba sericea are eaten locally in South America, and its seeds can be pressed to produce an oil used for frying food. Several species, such as white pear (Apodytes dimidiata), produce good timber. Etymology: Metteniusa is named for German botanist Georg Heinrich Mettenius (1823–1866), who was a botany scholar in Freiburg and Leipzig. He was an authority in pteridology and studied the marine algae of Heligoland.

GARRYALES Families 386 and 387 comprise the order Garryales, which are sister to the asterid I clade (lamiids). They are estimated to have diverged c. 114 million years ago.

386. EUCOMMIACEAE Chinese-rubbertree family

These are deciduous, unisexual trees with the wood and bark containing latex. They have alternate, spirally arranged, petiolate leaves without stipules. Leaf blades are simple with pinnate venation, serrate margins with glandtipped teeth, obliquely rounded at the base and a narrow, acuminate tip. Leaves contain latex, which can be seen when the leaves are pulled apart, the veins remaining attached by strands. Inflorescences are axillary clusters (male), or flowers are solitary (female) at the base of the current-year branchlets. Flowers are wind-pollinated, without a perianth and unisexual. Male flowers have five to 12 stamens with short, linear filaments and 514

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basifixed, pendent anthers that open by lengthwise slits. The connective is somewhat prolonged above the anthers. Female flowers have a stalked ovary composed of two fused carpels forming a single locule. It is laterally compressed, winged, bilobed at the apex and has two, reflexed and spreading stigmas on the top. Fruits are long-elliptic, compressed samaras with a wing around the margin, gradually narrowing at the base into the stipe and bilobed at the tip. Distribution: The family is restricted to China, where native stands are near threatened. It was previously thought to be extinct, but it has been cultivated in China for millennia and is frequently naturalised there, making it difficult to discern natural and naturalised populations. Phylogeny and evolution: Because of its reduced, wind-pollinated flowers, the taxonomic position of Eucommiaceae has been controversial. They have been variously associated with Hamamelidales, Magnoliales and Urticales or placed in their own order, Eucommiales. Molecular and chemical

studies have suggested that they are closest to Garryaceae in Garryales. Aucubin in the bark is indicative of a relationship to Aucuba. Fossils of Eucommia were found in 10–35-million-year-old brown coal deposits in North America, Europe, central Africa, Anatolia, Central Asia, East Asia and Siberia, indicating that the family had a much wider distribution in the past, their current restriction to central China being relictual. Eucommia is locally naturalised in Ohio, but does not pose a threat of becoming an invasive there. Genera and species: Eucommia ulmoides is the only species in this family. Uses: The sap produces a highly valued latex, used for insulating electrical cables, lining pipelines and dental fillings. The timber is used to make furniture and for fuel, and in Chinese medicine, the bark (duzhong), which contains aucubin, is used to treat arthritis. Etymology: Eucommia is composed of the Greek words ευ (eu), good, and κόμμι (kommi), gum.

GARRYALES

EUDICOTS

Eucommia ulmoides, Royal Botanic Gardens, Kew, UK [386]

Eucommia ulmoides, female flowers, Royal Botanic Gardens, Kew, UK [386] Eucommia ulmoides, male flowers, Royal Botanic Gardens, Kew, UK [386]

Garrya californica, female in fruit, Aucuba japonica, male, private San Francisco Botanical Garden, garden, Kingston upon Thames, California, USA [387] Surrey, UK [387]

387. GARRYACEAE Tasselbush family

This is a family of unisexual evergreen trees and shrubs. They have opposite, leathery petiolate leaves without stipules but with petioles united at the base across the stem. Blades are entire, somewhat pinnatifid or dissected, the venation is pinnate with secondary cross-veins, and margins are entire or broadly toothed. Inflorescences are bracteate, pendulous, silky hairy axillary catkins (Garrya) or cymes grouped in terminal or axillary thyrses (Aucuba). Flowers are unisexual and actinomorphic. Male flowers have four free sepals and four (Aucuba) or no (Garrya) petals. The four stamens are free, in one whorl alternating with the sepals (Garrya) or the petals (Aucuba). Anthers are

Aucuba japonica, female, Royal Botanic Gardens, Kew, UK [387]

basifixed or dorsifixed and open inward by lengthwise slits. Female flowers are sessile with two, four or no sepals, lacking petals (Garrya) or with four free petals (Aucuba). The inferior ovary is composed of one, two or three fused carpels forming a single locule, topped with one to three, free styles with dry stigmas. A thick disk surrounds the style. The fruit is a berry crowned by persistent styles. Distribution: This is a disjunct family, with Garrya found in western North America (Washington, Oregon, California and Mexico), Mesoamerica and the Antilles and Aucuba only in East Asia, from the eastern Himalayas, northern Vietnam, China and Taiwan to Honshu (Japan). Phylogeny and evolution: The placement of this family has been difficult in the past, and they were mostly associated with Cornales. Aucuba was in fact often included in Cornaceae. Chemical studies have shown that Aucuba, Garrya and Eucommia are closely related and form the order Garryales, putatively sister to the lamiid (asterid I) clade.

Garrya elliptica, male, Royal Botanic Gardens, Kew, UK [387]

Genera and species: Garryaceae include only two genera and c. 19 species: Aucuba (4) and Garrya (c. 15) Uses: A much cultivated plant, Japanese laurel (Aucuba japonica) is used in Shinto temples as offerings. Originally only female plants of a virus-infected, yellow-blotched cultivar (‘Variegata’) were introduced into Europe, but the attractive red fruits were not produced. Once a male was introduced and distributed, female plants set seed everywhere. It is highly tolerant to pollution and was popular in UK cities when coal was still burned to heat the houses. Feverbush or silk tassel bush (Garrya elliptica) is also frequently grown in mild gardens in Europe and North America. Garrya wrightii produces small amounts of latex that can be used as rubber. Etymology: Garrya is named by botanical collector David Douglas in honour of Nicholas Garry (1782–1856), secretary of the Hudson Bay Company. He named this genus after his friend who assisted him greatly while collecting plants in the Pacific Northwest. Plants of the World

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GENTIANALES

EUDICOTS

GENTIANALES Families 388 to 392 make up the order Gentianales, a group that evolved c. 71–108 million years ago. Dialypetalanthus had been suggested to be sister to the rest of the order, but it has been shown by molecular studies to be a member of Rubiaceae. Gentianales can be recognised by their wood with vestured pits, opposite leaves that are connected across the stem and nuclear endosperm formation.

388. RUBIACEAE Coffee family

This diverse family includes trees, shrubs, vines, epiphytes and annual and perennial herbs. Their leaves are simple, opposite or apparently whorled, with stipules between the petioles or between the petiole and the stem axis, sometimes fused to and sheathing the stem or becoming leaf-like and forming a false whorl with the true leaves. Bracts subtending inf lorescences are sometimes large and brightly coloured. Flowers are usually in axillary bracteate cymes that are compounded into panicles, fascicles or aggregated into heads, sometimes the flowers Coffea arabica, Oulu Botanical Garden, Finland (AP) [388]

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solitary. Flowers are actinomorphic (rarely zygomorphic), usually bisexual, sometimes unisexual, often heterostylous. The four or five fused sepals usually surround the top of the ovary, the lobes varying from nearly absent to large and often brightly coloured. The four or five (rarely three or eight to ten) petals are fused into a tube with free lobes, rarely bilabiate. Stamens are as many as petals (many more in Dialypetalanthus) and alternate with these, the filaments fused to the corolla tube. Anthers are free from each other or connivent or fused in a ring around the style. They are dorsifixed and open inward by lengthwise slits or (occasionally) by apical pores, sometimes bearing a connective appendix. The inferior ovary (superior in Gaertnera and Pagamea) is composed of usually two (rarely up to nine) fused carpels forming two (rarely one or up to nine) locules. A single terminal style is simple, or the two (to five) styles are free or partially fused, often variable in length. Fruits are fleshy or dry berries or drupes, sometimes schizocarpic capsules or indehiscent nutlets.

Distribution: Rubiaceae are a nearly globally distributed family that is only absent from the Arctic and Antarctic and permanently dry areas in the Sahara and Gobi Deserts. Rubiaceae are dominant in many parts of the world especially in the understory of tropical rainforests, where they can be incredibly diverse.

Asperula taurina, Kwekerij de Border, Delden, the Netherlands [388]

Coprosma robusta, Royal Botanic Gardens, Kew, UK [388]

Phylogeny and evolution: Traditionally Rubiaceae were subdivided into two subfamilies, depending on the numbers of ovules per ovary. This distinction was found to be artificial, and the subfamilies were recircumscribed resulting in three subfamilies that have been reorganised after new data became available. Molecular studies have found three clades (corresponding to Cinchonoideae, Ixoroideae and Rubioideae) with a poor resolution and support. It seems that a clade composed of Acranthera, Coptosapelta and Luculia may be sister either to Rubioideae (and then could be included in it) or to the entire family (and then a new subfamily will be needed that will be difficult

GENTIANALES

EUDICOTS

Gardenia taitensis, Tahiti, French Polynesia

Luculia gratissima, Royal Botanic Garden, Melbourne, Australia [388]

Phuopsis stylosa, Royal Botanic Gardens, Kew, UK [388]

to define morphologically because the genera are so divergent from each other). A four- or two-subfamily system can be accepted, and both have been recently proposed; experts on the family cannot agree, so their taxonomy is not yet crystallised. Molecular studies have also resulted in numerous genetic changes. Galium-like fossils that are 55 million years old are known. Rubiaceae are thought to have an origin in the Old World tropics, from where they migrated through the boreal zones (during a period when the region was tropical) into the New World via an Atlantic land bridge, although origins of some genera are suggested to be in South America c. 78 million years ago after which they further dispersed over long distances. Both scenarios are possible and not mutually exclusive.

(100), Arcytophyllum (18), Argocoffeopsis (10), Argostemma (164), Asperula (193), Astiella (1), Atractocarpus (29), Atractogyne (2), Augusta (4), Aulacocalyx (11), Badusa (3), Balmea (1), Bathysa (10), Batopedina (3), Belonophora (5), Benkara (19), Benzonia (1), Berghesia (1), Bertiera (56), Bikkia (10), Blepharidium (1), Bobea (4), Boholia (1), Bothriospora (1), Botryarrhena (2), Bouvardia (51), Brachytome (8), Bradea (5), Bremeria (18), Brenania (2), Breonadia (1), Breonia (20), Bruxanelia (1), Bullockia (6), Bungarimba (4), Burchellia (1), Burttdavya (1), Byrsophyllum (2), Callipeltis (3), Calochone (2), Calycophyllum (11), Calycosia (8), Calycosiphonia (3), Canephora (5), Canthium (100), Capirona (1), Carajasia (1), Carapichea (6), Carpacoce (7), Carphalea (3), Carterella (1), Casasia (10), Catesbaea (18), Catunaregam (12), Cephalanthus (6), Ceratopyxis (1), Ceriscoides (11), Ceuthocarpus (1), Chaetostachydium (3), Chalepophyllum (1), Chamaepentas (6), Chapelieria (1), Chassalia (114), Chimarrhis (15), Chiococca (23), Chione (1), Chomelia (78), Ciliosemina (2), Cinchona (23), Cinchonopsis (1), Cladoceras (1), Clarkella (1), Coccochondra (4), Coccocypselum (20), Coddia (1), Coelopyrena (1), Coelospermum (11), Coffea (124), Coleactina (1), Colletoecema (3), Condaminea (5), Conostomium (5), Coprosma (110), Coptophyllum (8), Coptosapelta (16), Coptosperma (19), Cordiera (11), Cordylostigma (9), Corynanthe (3), Coryphothamnus (1), Cosmibuena (4), Cosmocalyx (1), Coussarea (120), Coutaportia

(3), Coutarea (6), Cowiea (2), Craterispermum (18), Cremaspora (2), Cremocarpon (9), Crobylanthe (1), Crossopteryx (1), Crucianella (30), Cruciata (8), Cruckshanksia (7), Crusea (15), Cubanola (2), Cuviera (21), Cyanoneuron (5), Cyclophyllum (37), Damnacanthus (12), Danais (31), Debia (4), Deccania (1), Declieuxia (29), Dendrosipanea (2), Denscantia (4), Dentella (8), Deppea (40), Diacrodon (1), Dialypetalanthus (1), Dibrachionostylus (1), Dichilanthe (2), Dictyandra (2), Didymaea (5), Didymochlamys (2), Didymopogon (1), Didymosalpinx (5), Diodella (11), Diodia (22), Dioecrescis (1), Dioicodendron (1), Diplospora (23), Dirichletia (5), Discospermum (7), Diyaminauclea (1), Dolianthus (13), Dolichodelphys (1), Dolicholobium (28), Dolichometra (1), Dolichopentas (4), Donnelyanthus (1), Duidania (1), Dunnia (1), Duperrea (1), Duroia (38), Durringtonia (1), Eizia (1), Elaeagia (25), Eleuthranthes (1), Emmenopterys (2), Emmeorhiza (1), Empogona (28), Eosanthe (1), Eriosemopsis (1), Erithalis (8), Ernodea (8), Eteriscius (1), Euclinia (3), Everistia (1), Exallage (18), Exostema (34), Fadogia (40), Fadogiella (3), Faramea (207), Ferdinandusa (24), Feretia (3), Fergusonia (1), Fernelia (4), Flagenium (6), Flexanthera (1), Fosbergia (4), Gaertnera (70), Galianthe (48), Galiniera (2), Galium (622), Gallienia (1), Galopina (4), Ganguelia (1), Gardenia (133), Gardeniopsis (1), Genipa (3), Gentingia (1), Geophila (29), Gillespiea (1), Gleasonia (5), Glionnetia (1), Globulostylis (2), Glossostipula (3), Gomphocalyx (1), Gonzalagunia

[388]

Genera and species: Rubiaceae include c. 593 genera and c. 13,620 species (subfamilies are not used here, pending proposed subfamily classification): Acranthera (39), Acrobotrys (1), Acrosynanthus (6), Acunaeanthus (1), Adenorandia (1), Adina (7), Afrocanthium (17), Agathisanthemum (4), Agouticarpa (7), Aidia (55), Aidiopsis (1), Airosperma (6), Alberta (1), Aleisanthia (2), Aleisanthiopsis (2), Alibertia (24), Alleizettella (2), Alseis (18), Amaioua (9), Amaracarpus (30), Amphiasma (7), Amphidasya (13), Antherostele (4), Anthorrhiza (9), Anthosper mopsis (1), Anthospermum (39), Antirhea (38), Aoranthe (5), Aphaenandra (1), Aphanocarpus (1), Apomuria (12), Appunia (14), Arachnothryx

Plants of the World

517

GENTIANALES

Portlandia proctorii, Fairchild Tropical Botanical Garden, Florida, USA [388]

EUDICOTS

Pseudomussaenda flava, Singapore [388]

(37), Greenea (6), Greeniopsis (6), Guettarda (151), Guihaiothamnus (1), Gynochthodes (93), Gyrostipula (3), Habroneuron (1), Hamelia (17), Hedstromia (1), Hedyotis (174), Hedythyrsus (3), Heinsenia (1), Heinsia (5), Hekistocarpa (1), Henriquezia (3), Heterophyllaea (3), Hexasepalum (1), Hillia (24), Himalrandia (2), Hindsia (11), Hintonia (3), Hippotis (12), Hodgkinsonia (2), Hoffmannia (112), Holstianthus (1), Homollea (3), Homolliella (1), Houstonia (24), Hutchinsonia (2), Hydnophytum (94), Hydrophylax (1), Hymenocoleus (12), Hymenodict yon (23), Hyperacanthus (11), Hypobathrum (31), Hyptianthera (1), Involucrella (2), Isertia (14), Isidorea (17), Ixora (544), Jackiopsis (1), Janotia (1), Joosia (12), Jovetia (1), Kadua (30), Kailarsenia (6), Kajewskiella (2), Keenania (5), Keetia (32), Kelloggia (2), Kerianthera (2), Khasiaclunea (1), Klossia (1), Knoxia (12), Kochummenia (2), Kohautia (27), Kraussia (4), Kutchubaea (13), Ladenbergia (35), Lamprothamnus (1), Landiopsis (1), Larsenaikia (3), Lasianthus (242), Lathraeocarpa (2), Lecananthus (3), Lecariocalyx (1), Lelya (1), Lemyrea (4), Lepidostoma (1), Leptactina (27), Leptodermis (46), Leptomischus (7), Leptopetalum (5), Leptoscela (1), Leptostigma (7), Lerchea (10), Leucocodon (1), Leucolophus (3), Limnosipanea (3), Luculia (4). Lucya (1), Ludekia (2), Macbrideina (1), Machaonia (31), Macrocnemum (8), Macrosphyra (3), Maguireocharis (1), Maguireothamnus (2), Malanea (39), Manettia (125), Manostachya 518

Christenhusz, Fay & Chase

(3), Mantalania (3), Margaritopsis (49), Martensianthus (5), Maschalodesme (2), Massularia (1), Mastixiodendron (7), Mazaea (2), Melanopsidium (1), Mericarpaea (1), Merumea (2), Mexotis (4), Meyna (9), Micrasepalum (2), Microphysa (1), Mitchella (2), Mitracarpus (62), Mitragyna (10), Mitrasacmopsis (1), Mitriostigma (5), Molopanthera (1), Monosalpinx (1), Morelia (1), Morierina (2), Morinda (44), Morindopsis (1), Motleyia (1), Mouretia (4), Multidentia (10), Mussaenda (186), Mussaendopsis (3), Mycetia (45), Myrioneuron (8), Myrmecodia (27), Myrmeconauclea (4), Myrmephytum (5), Nargedia (1), Nauclea (12), Neanotis (31), Neblinathamnus (2), Nematostylis (1), Nenax (9), Neobertiera (1), Neoblakea (2), Neohymenopogon (3), Neolamarckia (2), Meomussaenda (2), Neonauclea (68), Nernstia (1), Nertera (9), Nesohedyotis (1), Neurocalyx (5), Nichallea (1), Nodocarpaea (1), Normandia (1), Nostolachma (6), Notopleura (102), Ochreinauclea (2), Octotropis (1), Oldenlandia (216), Oldenlandiopsis (1), Oligocodon (1), Omiltemia (2), Opercularia (17), Ophiorrhiza (317), Oreopolus (1), Osa (1), Otiophora (18), Otomeria (8), Ottoschmidtia (1), Oxyanthus (35), Oxyceros (12), Pachystylus (2), Paederia (33), Pagamea (25), Pagameopsis (2), Palicourea (335), Paracarphalea (3), Paracephaelis (4), Parachimarrhis (1), Paracorynanthe (2), Paragenipa (1), Paraknoxia (1), Parapentas (3), Patima (2), Pauridiantha (50), Pausinystalia (5), Pavetta (359), Payera (10), Pentagonia (36),

Psychotria urbaniana, Guadeloupe [388]

Pentaloncha (2), Pentanisia (19), Pentanopsis (2), Pentas (16), Pentodon (2), Peponidium (47), Perakanthus (1), Perama (14), Peripeplus (1), Pertusadina (4), Petitiocodon (1), Phellocalyx (1), Phialanthus (22), Phialiphora (2), Phuopsis (1), Phyllacanthus (1), Phyllis (2), Phyllocrater (1), Phyllomelia (1), Phyllopentas (14), Phylohydrax (2), Picardaea (1), Pimentelia (1), Pinarophyllon (2), Pinckneya (1), Pitardella (3), Pittoniotis (3), Placocarpa (1), Planaltina (3), Platycarpum (12), Plectroniella (1), Pleiocoryne (1), Plocama (34), Plocaniophyllon (1), Pogonopus (3), Polysphaeria (22), Polyura (1), Pomax (1), Porterandia (22), Portlandia (6), Posoqueria (22), Pouchetia (4), Praravinia (49), Preussiodora (1), Prismatomeris (15), Psathura (8), Pseudaida (1), Pseudodiplospora (1), Pseudohamelia (1), Pseudomantalania (1), Pseudomiltemia (2), Pseudomussaenda (6), Pseudonesohedyotis (1), Pseudopyxis (3), Psychotria (1,858), Psydrax (80), Psyllocarpus (9), Pteridocalyx (2), Pubistylus (1), Puffia (1), Pygmaeothamnus (2), Pyragra (2), Pyrostria (60), Rachicallis (1), Ramosmania (2), Randia (103), Raritebe (1), Razafimandimbisonia (5), Remijia (44), Rennellia (8), Retiniphyllum (20), Rhadinopus (2), Rhaphidura (1), Rhipidantha (1), Rhodopentas (2), Richardia (16), Riodocea (1), Riqueuria (1), Robbrechtia (2), Robynsia (1), Rogiera (15), Roigella (1), Ronabea (3), Rondeletia (157), Rosenbergiodendron (4), Rothmannia (41), Rubia (83), Rubovietnamia (2), Rudgea (130), Rustia (17), Rutidea (21),

GENTIANALES

EUDICOTS

Rytigynia (82), Sabicea (149), Sacosperma (2), Saldinia (22), Salzmannia (1), Saprosma (49), Sarcocephalus (2), Schismatoclada (19), Schizenterospermum (4), Schizocalyx (9), Schizocolea (2), Schizomussaenda (1), Schmidtottia (15), Schradera (53), Schumanniophyton (3), Schwendenera (1), Scolosanthus (28), Scyphiphora (1), Scyphostachys (2), Sericanthe (21), Serissa (1), Shaferocharis (3), Sherardia (1), Sherbournia (13), Siemensia (1), Simira (40), Sinoadina (1), Sipanea (19), Sipaneopsis (7), Siphonandrium (1), Sommera (10), Spathichlamys (1), Spermacoce (287), Spermadictyon (1), Sphinctanthus (8), Spiradiclis (38), Squamellaria (4), Stachyarrhena (13), Stachyococcus (1), Staelia (21), Standleya (4), Steenisia (5), Stenaria (6), Stenosepala (1), Stenostomum (48), Stenotis (7), Stephanococcus (1), Stevensia (11), Steyermarkia (1), Stichianthus (1), Stilpnophyllum (4), Streblosa (25), Streblosiopsis (1), Strumpfia (1), Stylosiphonia (1), Suberanthus (7), Synaptantha (2), Syringantha (1), Tamilnadia (1), Tammsia (1), Tamridaea (1), Tarenna (191), Tarennoidea (2), Temnocalyx (1), Temnopteryx (1), Tennantia (1), Tessiera (2), Thamnoldenlandia (1), Thecagonum (4), Theligonum (4), Thiollierea (12), Thogsennia (1), Timonius (182), Tinadendron (2), Tobagoa (1), Tocoyena (19), Tortuella (1), Trailliaedoxa (1), Triainolepis (13), Tricalysia (78), Trichostachys (14), Triflorensia (3), Trigonopyren (9), Uncaria (40),

Urophyllum (119), Valantia (7), Vangueria (57), Vangueriella (18), Vangueriopsis (4), Vidalasia (5), Villaria (5), Virectaria (8), Wandersong (2), Warszewiczia (8), Wendlandia (82), Wit t mack anthu s (1), Xanthophytum (32), Xantonnea (2), Xantonneopsis (1) and Zuccarinia (1). Uses: This family is the source of two famous drinks, coffee and Indian tonic. Coffee is the most heavily traded commodity after fossil fuels. The seeds of several species of Coffea can be roasted to make coffee. The most commonly used are arabica coffee (C. arabica), robusta coffee (C. canephora) and abeokuta coffee (C. liberica). Arabica coffee is cultivated throughout the tropics at higher elevations or in the shade of trees and has a mild aroma. The species is nearly extinct in its native habitat in Ethiopia. Robusta coffee can be grown in warmer climates where arabica will not thrive, and it is less susceptable to diseases. Robusta coffee is more bitter than arabica, and thus it is often used in Italian coffee mixes and to produce instant coffee. Coffe­a liberica seeds are also bitter and often added to robusta blends. Coffea arabica is usually much preferred over the others. Coffee beans were originally only chewed, but the earliest record of consumption of coffee as a beverage date from 15th century Yemen, where it was brewed in a monastery in Mocha. From there, it soon spread across the Middle

East and was soon smuggled to India where plantations were started in the 17th century. From there it spread to Europe, Indonesia and the Americas. Currently, Brazil is the largest producer of coffee, followed by Vietnam, Indonesia and Colombia. It has purported beneficial effects, although negative effects of coffee consumption have also been reported. In moderation, coffee is at least harmless, if not improving dexterity and mental health. Quinine is an antimalarial alkaloid found in high concentrations in species of Cinchona, especially yellow bark (C. calisaya and cultivars), and red bark (C. pubescens), and Georgia bark (Pinckneya bracteata). The bark was originally harvested from trees in the wild in South America and exported to colonies around the world, but plantations were soon established. It is the main flavour component of tonic water, which was widely consumed to combat malaria, but preferably after gin, sugar and lime were added to reduce the bitterness of the drink. Originally popular in British India, gin and tonic is now consumed worldwide. Another important alkaloid is emetine, which is used against amoebic dysentery and harvested from the ipecacuanha plant (Carapichea ipecacuanha). Noni (Morinda citrifolia) has an edible fruit that is eaten in salads, and the young leaves are used in Thai cuisine, even though it is sometimes said to be mildly toxic. It contains xeronine, which is used medicinally

Rothmannia annae, Helsinki Botanical Garden, Finland [388]

Rubia tinctorum, Ruissalo Botanical Garden, Turku, Finland [388]

Ramosmania rodriguesii, Royal Botanic Gardens, Kew, UK [388]

Plants of the World

519

GENTIANALES

520

EUDICOTS

for stomach cramps, but is also used as an insecticide. The bark yields a red dye, whereas the roots can be ground to produce a yellow dye, both commonly used in batik dying techniques. Cat’s claw (Uncaria tomentosa) bark is used medicinally as an anti-inflammatory, but is no longer grown on a large scale. Gambier (U. gambir) has been used as an alternative for chewing with betel nuts (Areca catechu, Arecaceae). It was previously used for tanning and grown in plantations for this purpose in the 18th century, but it is now rarely grown. Other stimulants are the proven aphrodisiac yohimbine (Pausinystalia johimbe) and Psychotria viridis, which is mixed with other hallucinogens in South America to make the drug ayahuasca. Woodruff (Galium odoratum) contains coumarin and can be used as a spice in moderation. It is frequently used to flavour syrupy drinks and some Dutch and German beers and was previously used as a strewing herb. Some species produce edible fruit that is locally eaten, especially black guava (Guettarda argentea, Neotropics), the applelike Guinea peach (Sarcocephalus latifolius,

Africa) and Spanish tamarind (Vangueria madagascariensis, Africa). Apart from noni, several other species produce good natural dyes, for example madder (Rubia tinctorum), dyer’s woodruff (Asperula tinctoria) and Indian madder (Oldenlandia umbellata), the crushed roots of which yield a red dye when mixed with alkali. Genip (Genipa americana) fruits were used to dye cloth and skin by native Americans. Many species are trees, the wood of which is of local importance. A number of genera include valuable ornamental plants, including Alberta, Asperula, Bouvardia, Cephalanthus, Coprosma, Emmenopterys, Galium, Gardenia, Hillia, Hoffmannia, Houstonia, Ixora, Luculia, Manettia, Mitchella, Mussaenda, Myrmecodia, Osa, Pavetta, Pentas, Phuopsis, Pseudomussaenda, Psychotria, Rondeletia, Rothmannia, Rudgea, Serissa, Tarenna and Wendlandia; many others are of local ornamental interest. Etymology: Rubia is derived from the Latin ruber, red. The roots of some species of madder (mainly R. tinctorum) have been used since ancient times to produce a red dye.

This family includes autotrophic (green) and mycoheterotrophic (non-green), glabrous, annual and perennial herbs, shrubs, trees and vines. They usually have opposite leaves (rarely whorled in Swertia and seldom alternate), which are simple and lack stipules, but at the bases are often connected by a line across the node. Blades usually have entire margins, and venation is pinnate, palmate or palmate-parallel with transverse cross-venation; blades are sometimes reduced to scales in achlorophyllous species. Inflorescences are terminal or axillary simple or compound dichasia, with or without bracts and bracteoles. Flowers are usually bisexual

Eustoma grandiflora, Sherman Gardens, Corona del Mar, California, USA [389]

Lisianthius axillaris, Helsinki Botanical Garden, Finland [389]

Blackstonia perfoliata,Seven Sisters, East Sussex, UK [389]

Christenhusz, Fay & Chase

389. GENTIANACEAE Gentian family

GENTIANALES

EUDICOTS

Centaurium venustum, Palm Canyon, Palm Springs, California, USA [389]

Swertia perennis, private garden, Kingston upon Thames, Surrey, UK [389]

Fagraea fragrans, Singapore [389]

Halenia serpyllifolia, Ecuador [389]

and actinomorphic (or the calyx slightly zygomorphic). The four or five (to 12) sepals are fused, usually regular, sometimes bilabiate. Petals are as many as sepals and fused into a usually showy, tubular, campanulate or funnelshaped, regular corolla that often has nectar scales or nectar pits inside. Stamens are as many as and alternate with the corolla lobes, with filaments fused to the corolla and free from each other or (rarely) fused into a ring inside the corolla tube. Anthers are dorsifixed and versatile or rarely basifixed and opening inwardly by lengthwise slits or pores, pores are rarely apical. The superior ovary is composed of two carpels, forming one (rarely two) locules with a single style topping the ovary and a simple or bilobed stigma or the stigma sessile on the ovary. Fruits are usually dehiscent capsules opening with two valves, rarely berries. Seeds are numerous and dust-like.

Distribution: This is a widespread family, found from the far Arctic to the Antarctic islands and throughout the tropics (except for the great deserts of the world) with the greatest diversity in montane and temperate areas. Phylogeny and evolution: The crown age of Gentianaceae is estimated to be c. 50–66 million years. There is Eocene fossil pollen attributed to Gentianaceae, but this is of uncertain affinity. Five clades are recognised in Gentianaceae, which are treated at the tribal level, although relationships between these tribes are not well understood. The genera Anthocleista, Fagraea and Potalia were formerly placed in Loganiaceae but belong here on the basis of their chemistry and molecular analyses. Similarly, the chemistry of Emblingia, previously placed here, suggests a relationship with Brassicales, where it is now placed (as Emblingiaceae).

Irlbachia frigida, Guadeloupe [389]

Obolaria virginica, North Carolina, USA [389]

Saccifolium has strange pouch-like leaves, leading to suggestions that it might be carnivorous, but there is no convincing evidence that these species demonstrate digestion and absorption. The pouches are downward-facing, so it seems highly unlikely that they could be effective trapping structures. This illustrates how little is known about many genera of Gentianaceae, and basic anatomical, chemical and developmental studies are needed to understand the evolution of diversity in this family. It is likely that future studies will result in some generic recircumscriptions. The mycoheterotrophic genera (Bartonia, Cotylanthera, Obolaria, Voyria and Voyriella) are not most closely related to each other; for instance, Obolaria is in Swertiinae and Voyriella in Saccifolieae, whereas Voyria is distant from both of these. Therefore Plants of the World

521

GENTIANALES

Gentiana clusii, Royal Botanic Gardens, Kew, UK [389]

Gentiana lutea, Helsinki Botanical Garden, Finland [389]

Sabatia kennedyana, New York Botanical Garden, USA [389]

mycoheterotrophy has evolved several times independently in Gentianaceae.

Roraimaea (2), Sabatia (20), Saccifolium (1), Schenkia (5), Schinziella (1), Schultesia (15), Sebaea (c. 70), Senaea (2), Sinogentiana (2), Sinoswertia (1), Sipapoantha (2), Swertia (134), Symbolanthus (32), Symphyllophyton (1), Tachia (13), Tachiadenus (11), Tapeinostemon (7), Tetrapollinia (1), Tripterospermum (28), Urogentias (1), Veratrilla (2), Voyria (18), Voyriella (1), Xestea (1), Yanomamua (1), Zeltnera (25), Zonanthus (1) and Zygostigma (1).

Etymology: Gentiana is named for King Gentius of Illyria, who is said to have discovered the medicinal properties of G. lutea in the 2nd century BC.

Genera and species: Gentianaceae include c. 101 genera and c. 1,690 species: Adenolisianthus (1), Anthocleista (14), Aripuana (1), Bartonia (4), Bisgoeppertia (2), Blackstonia (4), Calolisianthus (6), Canscora (7), Canscorinella (2), Celiantha (3), Centaurium (20), Chelonanthus (7), Chironia (15), Chorisepalum (5), Cicendia (2), Comastoma (15), Congolanthus (1), Cotylanthera (4), Coutoubea (5), Cracosna (3), Crawfurdia (20), Curtia (6), Deianira (5), Djaloniella (1), Duplipetala (2), Enicostema (3), Eustoma (3), Exaculum (1), Exacum (65), Exochaenium (32), Fagraea (c. 70), Faroa (19), Frasera (15), Geniostemon (5), Gentiana (334), Gentianella (275), Gentianopsis (24), Gentianothamnus (1), Gynandra (5), Halenia (39), Helia (2), Hockinia (1), Hoppea (2), Irlbachia (9), Ixanthus (1), Jaeschkea (3), Karina (1), Klackenbergia (2), Kuepferia (12), Lagenanthus (1), Lagenias (1), Latouchea (1), Lehmanniella (2), Limahlania (1), Lisianthius (30), Lomatogonium (24), Macrocarpaea (100), Megacodon (2), Metagentiana (14), Microrphium (1), Neblinantha (2), Neurotheca (3), Obolaria (1), Oreonesion (1), Ornichia (3), Orphium (2), Phyllocyclus (5), Potalia (9), Prepusa (5), Pterygocalyx (1), Purdieanthus (1), Pycnosphaera (1), Rogersonanthus (2), 522

EUDICOTS

Christenhusz, Fay & Chase

Uses: The scented f lowers of Fagraea ceilanica (perfume tree) and F. fragrans (kingwood) have been used in the perfume industry. Yellow gentian (Gentiana lutea) produces gentian root, which is used as a bitter flavouring in drinks and liqueurs such as Campari and Cinzano vermouth. Its leaves are similar to those of the poisonous monocot Veratrum (Melanthiaceae), and confusion has resulted in fatal consequences in the past. It is important to know your taxonomy! Eustoma grandiflorum is the ‘lisianthus’ of the cut-flower industry. The wood of Fagraea is rot resistant and has been used to carve tikis (wooden statues) in the Pacific; this genus produces good timber, which is used for construction throughout tropical Asia. Several genera contain valued ornamental plants, especially Centaurium, Eustoma, Exacum, Fagraea, Gentiana, Orphium and Sabatia.

390. LOGANIACEAE Indian-pink family

Loganiaceae are annual and perennial herbs, shrubs, trees and lianas with clear sap. When lianas (e.g. Strychnos), they have axillary curved thorns that hook into supports and stems that twine. Plants rarely have leaves that are highly reduced, but if so then the plant photosynthesises in its green stems (some Logania). Leaves are opposite or whorled, petiolate or sessile and usually fused at the base to each other by a stipular sheath, or the stipules free, forming a line across the stem node, or the stipules developed into leaf-like structures and then the leaves appearing whorled. Blades are simple, the margins entire and venation pinnate, sometimes

GENTIANALES

EUDICOTS

palmate or reduced to a single midvein. Inflorescences are terminal or axillary, rarely leaf-opposed, bracteate cymes or panicles, occasionally racemes or umbels, sometimes flowers solitary. Flowers are subtended by bracteoles (a fused involucre in Phyllangium), bisexual or unisexual and actinomorphic or sometimes some of the corolla lobes larger (Usteria). The four or five sepals are free or fused into a cup-shaped, lobed calyx. The four or five petals are fused with a tubularcampanulate or rotate corolla. The four or five stamens (one in Usteria) are inserted between the lobes halfway up the corolla tube. The dorsifixed, sometimes basifixed, anthers open by lengthwise slits. The superior ovary is composed of two carpels (three in some Geniostoma), each forming a locule. The single style has a single, entire or lobed, usually club-shaped stigma. Mitrasacme has two free styles, each with a capitate stigma. Fruits are dehiscent capsules or

schizocarps that split into two separate nutlets, rarely drupes.

Geniostoma borbonicum, Réunion [390]

Geniostoma borbonicum in fruit, Réunion [390]

Distribution: A pantropical family, Loganiaceae have a few temperate representatives in southeastern North America, northeastern Asia (China, Korea and Japan) and in Australasia (Tasmania, Western Australia and New Zealand). Phylogeny and evolution: This family used to be larger before the advent of molecular systematics. For instance, it included Anthocleista, Fagraea and Potalia (now Gentianaceae), Gelsemium and Mostuea (now Gelsemiaceae), Buddleja (now Scrophulariaceae), Retzia (now Stilbaceae), Desfontainia (now Columelliaceae) and Plocosperma (now Plocospermataceae). The resulting concept of Loganiaceae is sometimes further divided into Antoniaceae, Geniostomataceae, Mitreolaceae, Spigeliaceae

Strychnos henningsii, Lake Sibaya, KwaZulu Natal, South Africa (CD) [390]

and Strychnaceae. Together these form a clade, and there is no advantage in using such narrow family circumscriptions. They all share colporate pollen without lateral extensions. Gelsemiaceae are morphologically similar, but they do not share this pollen type and usually have fewer ovules per locule. Genera and species: Loganiaceae include 15 genera and c. 390 species: Antonia (1), Bonyunia (4), Gardneria (5), Geniostoma (24), Logania (c. 22), Mitrasacme (54), Mitreola (6), Neuburgia (c. 11), Norrisia (2), Orianthera (13), Phyllangium (5), Schizacme (4), Spigelia (c. 50), Strychnos (190) and Usteria (1). Uses: The members of this family are deadly poisonous (containing the alkaloid strychnine), and Spigelia and especially Strychnos have been used as a poison in pest control and crimes. Several species of Strychnos have edible apple-like fruits Phyllangium paradoxum, Mt Benia, Western Australia [390]

Spigelia marilandica, New York Botanical Garden, USA [390]

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(e.g. S. madagascariensis, S. ignatii and S. spinosa), although the seeds are high in strychnine and can be used to make rat poison; however, for that purpose S. nux-vomica is more commonly employed. Strychnos toxifera is used as curare in South America (see also Menispermaceae). Indian pink or Maryland pinkroot (Spigelia marilandica), native to southeastern North America, is occasionally grown as a garden ornamental. Etymology: Logania is named in memory of Irish natural historian and Mayor of Philadelphia, James (Jacobus) Logan (1674– 1751). He came to the Pennsylvania Colony in 1699 as William Penn’s secretary. Of Quaker heritage himself, he opposed Quaker pacifism in Pennsylvania, became wealthy through the fur trade and experimented especially with the pollination of plant seeds (in maize). He was a tutor of John Bartram and Benjamin Franklin.

391. GELSEMIACEAE Yellow-jessamine family

Pteleocarpa lamponga, Singapore (WA) [391]

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This is a family of shrubs, trees and woody vines with clear sap. If vining, they climb with twining stems. Leaves are opposite or whorled and petiolate, and sometimes have stipules reduced to a line across the stem node, rarely fused to form a sheath joining the opposing leaf bases. Blades have pinnate venation, and margins are entire or slightly dentate or wavy. Inflorescences are axillary or terminal cymes often organised into panicles, sometimes with a pair of fused involucral bracts (Mostuea). Flowers are bisexual, actinomorphic (or at least mostly so), often heterostylous and fragrant. The five (rarely four) sepals are nearly free or fused into a short basal tube, persistent in fruit. The five (rarely four) petals are fused into a funnel-shaped or cup-shaped, more or less regular corolla. The (four or) five stamens are free from each other and inserted near the base of the corolla tube between the petals. Anthers are free or connivent, dorsifixed and open by lengthwise slits. The superior ovary is composed of two carpels, each forming a locule. It is topped with two entirely or partially fused styles and two- or four-forked or four-lobed stigmas. Fruits are dehiscent capsules or samaras with the wing surrounding the indehiscent capsule similar to an Ulmus fruit (Pteleocarpa). Seeds are winged (Gelsemium) or not.

southeastern North America along the Gulf of Mexico south to the Yucatán Peninsula and in the Guianas. It is widespread in tropical Africa, Madagascar, Southeast Asia, Sumatra and Borneo.

Distribution: This is a mostly Palaeotropical family, but found in the Americas from

Etymology: Gelsemium is the Latinised form of gelsomino, the Italian word for jasmine.

Mostuea brunonis, Montagne d’Ambre National Park, Madagascar (CD) [391]

Phylogeny and evolution: Gelsemium and Mostuea were originally placed in Loganiaceae, but were found to form a separate clade sister to core Loganiaceae from which they differ in ovule number and pollen type. They were recently elevated to family status. Pteleocarpa lamponga was placed in Icacinaceae or Boraginaceae, in which it was always an odd element and was thus sometimes suggested to form its own family. Pteleocarpa is now known to be sister to the remainder of Gelsemiaceae, with which it shares the bicarpellate ovaries with two ovules per locule and the pentamerous flowers with a fused, regular perianth. Genera and species: This family comprises three genera and 11 species: Gelsemium (2), Mostuea (8) and Pteleocarpa (1). Uses: Yellow jessamine (Gelsemium sempervirens) is highly poisonous and has been used in crimes, but it is occasionally grown as an ornamental vine in North America because of its scented flowers.

Mostuea batesii, Mondah Forest, Gabon (CD) [391]

Gelsemium sempervirens, cultivated in Irvine, California, USA [391]

GENTIANALES

EUDICOTS

392. APOCYNACEAE Periwinkle family

This is a diverse family including annual and perennial herbs, shrubs, vines, lianas, trees, mangroves, epiphytes and leaf less succulents, sometimes cactus-like with photosynthetic stems. When climbing, they twine anti-clockwise or attach themselves by forming roots along the stem. Sap is clear or latex-like, and leaves are usually well developed, but they can be transformed into spines or much reduced to scales, sometimes pitcher-shaped (Dischidia major). Leaves are simple, opposite or whorled, rarely alternate, and with or without stipules. Leaf blades have pinnate to palmate venation and entire margins. Inf lorescences are axillary cymes, racemes, umbels or terminal panicles, or flowers are solitary. Flowers are actinomorphic and bisexual, often with a sweet or foetid fragrance. The five sepals are fused at the base, the lobes longer than the tube. The five petals are fused in a short or Amsonia tabernaemontana, private garden, Hengelo, the Netherlands [392]

Distribution: This is a widespread family found from the Arctic to the tropics and from wet rainforests to dry deserts. They are most diverse in tropical and warm-temperate regions with an impressive radiation in southern Africa.

Phylogeny and evolution: Apocynaceae have been estimated to have a crown age of c. 54 million years, although much younger ages (e.g. 21 million years) have also been suggested. Traditionally Asclepiadaceae were separate from Apocynaceae, but no clear differences in morphological characters separated the two concepts. Molecular studies have shown that members of the two families are interdigitated. Subfamily Rauvolfioideae are paraphyletic, forming a grade relative to the other subfamilies. Generic circumscription is particularly difficult in this family because it has often been based on minute floral characteristics. This attention to fine details has resulted in many larger genera (such as Asclepias, Ceropegia, Hoya and Marsdenia) becoming para- or polyphyletic if segregates are accepted. However, reorganisation is difficult because new characters will have to be found that can be used to identify these larger genera. For instance, several genera are embedded in Caralluma, which in turn is embedded in Ceropegia, the latter badly polyphyletic. Hoya and Dischidia are embedded in Marsdenia. Both problems will have to be addressed taxonomically and nomenclaturally in the future. Asclepias is also grossly polyphyletic and could include a large number of segregate genera. Needless to say, the list of genera below is only tentative.

Alyxia buxifolia, Western Australia [392]

Apocynum androsaemifolium, Michigan, USA [392]

long tube, sometimes with appendages that are simple scales or form a corona. Stamens and pistils are free or fused together to form a gynostegium or stamens and pistils free. When a gynostegium is present, the stamens are fused to each other around the style forming a nectariferous corona, the anthers strongly modified and individually attached to the style-head, basifixed, and the appendages have horn-like wings. When stamens are free from the pistil, they are inserted in the corolla tube, alternating with the petals, the anthers basifixed, sometimes prolonged at the tip. Pollen is shed free or aggregated in pollinia (somewhat similar to Orchidaceae, although in detail different). The superior ovary is composed of two carpels, each forming a cell and topped with a partially joined style and fused stigma, splitting when fruit develops. Fruits are formed of two (or one by abortion) separate follicles that open along the suture, sometimes fleshy and berry-like. Seeds in follicular (dry) fruits are usually conspicuously hairy with a tuft of long silky hairs on the top and usually with a thin wing around the seed.

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Catharanthus roseus, Helsinki Botanical Garden, Finland [392]

Plumeria rosea, planted in Los Angeles, California, USA [392]

Nerium oleander, Salies de Béarn, Provence, France [392]

Periploca graeca, private garden, Kingston upon Thames, Surrey, UK [392]

Vinca minor, Hampton Court Palace Garden, Surrey, UK [392]

Adenium obesum, Aurangabad, India [392]

Wrightia antidysenterica, Singapore Botanical Garden [392]

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Secamone volubilis, Réunion [392]

Ceropegia distincta subsp. haygarthii, Ruissalo Botanical Garden, Turku, Finland [392]

Calotropis gigantea, Singapore Botanical Garden [392]

Hoya macgillivrayi, Mary Selby Botanical Garden, Florida, USA [392]

Christenhusz, Fay & Chase

Orbea variegata, private garden, Kingston upon Thames, Surrey, UK [392]

GENTIANALES

EUDICOTS

Genera and species: Apocynaceae include c. 322 genera and c. 4,300 species in five subfamilies: Rauvolfioideae (79 genera) – Acokanthera (5), Allamanda (c. 15), Alstonia (c. 40), Alyxia (106), Ambelania (3), Amsonia (19), Ancylobothrys (7), Anechites (1), Aspidosperma (70), Bousigonia (2), Callichilia (7), Calocrater (1), Cameraria (2), Carissa (7), Carvalhoa (1), Catharanthus (8), Cerbera (2), Cerberiopsis (2), Chamaeclitandra (1), Chilocarpus (13), Clitandra (1), Condylocarpon (7), Couma (6), Craspidospermum (1), Crioceras (1), Cyclocotyla (1), Cylindropsis (1), Dictyophleba (5), Diplorhynchus (1), Dyera (2), Geissospermum (5), Gonioma (1), Hancornia (4), Haplophyton (1), Himatanthus (13), Hunteria (12), Kamettia (1), Kopsia (22), Lacmella (c. 20), Landolphia (56), Laxoplumeria (3), Lepinia (4), Lepiniopsis (2), Leuconotis (5), Macoubea (3), Melodinus (19), Microplumeria (1), Molongum (3), Mortoniella (1), Mucoa (2), Neocouma (2), Ochrosia (c. 40), Orthopichonia (6), Pacouria (2), Parahancornia (6), Petchia (8), Picralima (1), Plectaneia (3), Pleiocarpa (5), Plumeria (8), Pteralyxia (2), Pycnobotrya (1), Rauvolfia (c. 80), Rhigospira (1), Saba (3), Schizozygia (1), Skytanthus (2), Spongiosperma (6), Stephanostegia (2), Strempeliopsis (2), Tabernaemontana (110), Tabernanthe (2), Thevetia (8), Tonduzia (7), Vahadenia (2), Vallesia (5), Vinca (5), Voacanga (12) and Willughbeia (16); Apocynoideae (77 genera) – Adenium (5), Aganonerion (1), Aganosma (8), Alafia (23), Allomarkgrafia (9), Allowoodsonia (1), Amalocalyx (3), Angadenia (2), Anodendron (17), Apocynum (9), Artia (4), Asketanthera (4), Baharuia (1), Bahiella (2), Baissea (18), Beaumontia (9), Carruthersia (3), Chonemorpha (10), Cleghornia (4), Cycladenia (1), Dewevrella (1), Echites (4), Ecua (1), Elytropus (1), Epigynum (5), Eucorymbia (1), Farquharia (1), Fernaldia (3), Forsteronia (c. 40), Funtumia (2), Galactophora (6), Holarrhena (4), Hylaea (2), Ichnocarpus (12), Isonema (3), Ixodonerium (1), Kibatalia (15), Laubertia (3), Macropharynx (5), Malouetia (c. 19), Malouetiella (2), Mandevilla (c. 130), Mascarenhasia (8), Mesechites (12), Motandra

(3), Neobracea (4), Nerium (1), Odontadenia (20), Oncinotis (7), Pachypodium (25), Papuechites (1), Parameria (3), Parepigyum (1), Parsonsia (82), Peltastes (6), Pentalinon (2), Pinochia (4), Pleioceras (5), Pottsia (3), Prestonia (c. 55), Rhabdadenia (4), Secondatia (4), Sindechites (1), Spirolobium (1), Stephanostema (1, extinct in wild), Stipecoma (1), Streptoechites (1), Strophanthus (38), Temnadenia (3), Thenardia (3), Thoreauea (3), Tintinnabularia (3), Trachelospermum (10), Urceola (16), Vallariopsis (1), Vallaris (3) and Wrightia (23); Periplocoideae (30 genera) – Atherandra (2), Baroniella (8), Baseonema (1), Buckollia (2), Camptocarpus (9), Cryptolepis (c. 15), Cryptostegia (2), Decalepis (4), Ectadium (3), Epistemma (3), Finlaysonia (3), Gymnanthera (2), Hemidesmus (1), Ischnolepis (1), Kappia (1), Maclaudia (1), Mondia (2), Myriopteron (1), Parquetina (3), Pentopetia (21), Periploca (11), Petopentia (1), Raphionacme (34), Sacleuxia (2), Sarcorrhiza (1), Schlechterella (2), Streptocaulon (9), Tacazzea (4), Telectadium (3) and Zygostelma (1); Secamonoideae (6 genera) – Calyptranthera (2), Pervillaea (4), Secamone (c. 80), Secamonopsis (2) and Trichosandra (1); Asclepiadoideae (c. 130 genera) – Adelostemma (3), Aidomene (1), Anatropanthus (1), Anisopus (2), Anisotoma (2), Anomalluma (2), Apteranthes (6), Araujia (10), Asclepias (c. 125), Astephanus (2), Asterostemma (1), Australluma (2), Barjonia (6), Baynesia (1), Blepharodon (c. 45), Boucerosia (6), Brachystelma (122), Calciphila (2), Calotropis (3), Campestigma (1), Caralluma (28), Cathetostemma (1), Caudanthera (4), Ceropegia (160), Cibirhiza (2), Conomitra (1), Cosmostigma (3), Cynanchum (c. 225), Desmidorchis (12), Diplolepis (6), Dischidia (80), Ditassa (c. 55), Dolichopetalum (1), Dregea (c. 12), Duvalia (18), Duvaliandra (1), Echidnopsis (32), Edithcolea (1), Emicocarpus (1), Emplectanthus (2), Eustegia (1), Fischeria (16), Fockea (6), Funastrum (17), Glossonema (5), Gongronema (15), Gonolobus (150), Graphistemma (1), Gunnessia (1), Hemipogon (10), Heterostemma (12), Heynella (1), Holostemma (1), Hoodia (14), Hoya (>200), Huernia (67), Hypolobus (1), Jobinia (3),

Kanahia (2), Larryleachia (5), Lavrania (1), Leptadenia (4), Lygisma (3), Macroscepis (7), Mahawoa (1, extinct), Marsdenia (c. 200), Matelea (186), Merrillanthus (1), Metaplexis (6), Microloma (10), Minaria (19), Morrenia (9), Monolluma (5), Monsanima (1), Nautonia (1), Neoschumannia (1), Nephradenia (10), Notechidnopsis (2), Odontanthera (1), Odontostelma (1), Oncinema (1), Ophionella (2), Orbea (56), Oreosparte (1), Orthanthera (4), Orthosia (20), Oxypetalum (125), Oxystelma (2), Pectinaria (3), Pentarrhinum (2), Pentasachme (4), Pentascyphus (1), Pentastelma (1), Pentatropis (2), Peplonia (6), Pergularia (2), Petalostelma (7), Pherotrichis (2), Piaranthus (8), Pseudolithos (6), Pycnorhachis (1), Quaqua (29), Raphistemma (2), Rhyssolobium (1), Rhyssostelma (1), Rhytidocaulon (10), Richtersveldia (1), Riocreuxia (8), Sarcolobus (14), Sarcostemmma (35, often included in Cynanchum), Schubertia (6), Seshagiria (1), Sichuania (1), Sisyranthus (12), Socotrella (1), Solenostemma (1), Stapelia (47), Stapelianthus (9), Stelmagonum (2), Stenomeria (3), Stigmatorhynchus (3), Tassadia (17), Tavaresia (3), Telosma (10), Treutlera (1), Tromotriche (11), Vincetoxicum (c. 70), Whitesloanea (1) and Widgrenia (1). Uses: Several species have edible fruits, although care has to be taken because the seeds are often poisonous. Best-known edible fruits are produced by Carissa species, such as amatungulu (C. bispinosa) and Natal plum (C. macrocarpa), which are frequently sold in local markets. Fruits of karanga (C. carandas) are pickled in Africa, and in Costa Rica guayato (Gonolobus edulis) is eaten fresh. Hancornia speciosa fruit is used for marmalade and ice-cream. Several others have edible fruits, e.g. Adenium ellenbeckii, Clitandra cymulosa, Couma guianensis, Lacmella, Leptadenia pyrotechnica, Macoubea, Marsdenia australis (alunqua), Morrenia odorata, Parsonsia edulis, Saba comorensis and Urceola esculenta (kyetpaung). Seeds of Ochrosia elliptica are edible, and so are the flowers of Fernaldia species and Periploca aphylla. Wattakaka (Dregea volubilis) leaves are eaten with

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Asclepias incarnata, Matthaei Botanical Garden, Ann Arbor, Michigan, USA [392]

Asclepias incarnata, fruit releasing the seeds, Ann Arbor, Michigan, USA [392]

Monarch caterpillar on Asclepias tuberosa, St Petersburg, Florida, USA [392]

curry in India, D. abyssinica leaves are eaten in Uganda and leaves and flowers of Telosma cordata are used in Thai cuisine. Several species make edible roots or tubers, such as Brachystelma glabriflorum, Ceropegia affinis, C. bulbosa, Cibirhiza dhofarensis, Marsdenia hamiltonii, M. castillonii, M. flavescens, M. viridiflora and Raphionacme arabica. Roots of Finlaysonia khasiana are dried and burnt as incense in Laos for ceremonial purposes. Aganosma leaves are used to brew a tea. Several species produce a blue dye similar to indigo. Commercially, Java indigo (Marsdenia tinctoria), kyetpaung (Urceola esculenta) and Wrightia tinctoria were important for this purpose. Fibre is harvested from many species, especially Indian hemp (the bark of Apocynum cannabinum) and kendyr fibre (A. venetum) for ropes, sails etc. Other genera that have members that yield fibre are Araujia, Asclepias, Calotropis, Chonemorpha, Parameria and Pottsia. The silky seeds (Apocynum, Asclepias, Melaplexis, Periploca, etc.) are sometimes used as ‘vegetable’ silk, dental floss and filling for pillows. The many latex-producing species often make good rubber, and some have been used commercially for this purpose; examples include leche caspi (Couma macrocarpa), rubber vine (Cryptostegia grandif lora), Funtumia elastica, magabeira rubber (Hancornia speciosa), Lacmella, Parameria,

Petopentia, bitinga rubber (Raphionacme utilis) and Urceola. Many species are highly poisonous, including Acokanthera oppositifolia, Nerium oleander and Pachypodium lealii. Strophanthus kombe and S. gratus contain cardiac glycosides that have been applied to poison arrows and unfortunately have contributed to accidental poisonings and assassinations. The great variety of alkoloids in this family has also resulted in a good number of medicinal plants: Madagascar periwinkle (Catharanthus roseus) has been used in the treatment of leukaemia (vinblastine), and the bark of Picralima nitida is used against malaria in Cameroon. Iboga (Tabernanthe iboga) is a hallucinogen traditionally used in tropical Africa during initiation rituals. The active compound, ibogaine, is used in opiate addiction treatments, being more effective than methadone. Leaves of Lacmella lactescens are used as a coca substitute in northwestern South America. Xhoba (Hoodia gordonii) yields an appetite suppressant used commercially in weight-loss products. Dita bark (Alstonia scholaris) wood was used as writing slate in Malaysia. The chalk writing could be wiped off with Tetracera leaves (Dilleniaceae). The wood was formerly used to make coffins and the bark medicinally in child birth. A number of genera (e.g. Alstonia, Aspidosperma, Dyera, Funtumia,

Gonioma, Kibatalia and Wrightia) produce fine timber, used for various purposes, including carving, construction and furniture. Despite many being highly poisonous, many have highly scented, colourful or interesting-looking flowers and are therefore popular ornamentals, the following genera especially are commonly encountered in gardens: Acokanthera, Adenium (A. obesum, desert rose), Allamanda (A. cathartica, golden trumpet vine), Amsonia, Apocynum, Araujia, Asclepias (milkweed), Beaumontia, Brachystelma, Calotropis, Caralluma, Carissa, Catharanthus, Cerbera, Ceropegia (chain-of-hearts, C. linearis subsp. woodii), Chonemorpha, Cryptostegia (C. grandiflora, rubber vine), Cynanchum, Dischidia, Holarrhena, Hoya (waxflower), Mandevilla, Marsdenia (M. floribunda, stephanotis or Madagascar jasmine), Melodinus, Nerium (N. oleander, oleander), Oxypetalum (O. caeruleum, tweedia), Pachypodium (P. lamerei, Madagascar palm), Parsonsia, Periploca (P. graeca, silk vine), Plumeria (e.g. P. rubra, frangipani), Rauvolfia, Saba, Stapelia (S. hirsuta, carrion plant), Strophanthus, Tabernaemontana, Thevetia (T. peruviana or Cascabela thevetia, yellow oleander), Trachelospermum, Vinca (periwinkle) and Wrightia. A great diversity of succulent stapelioids (e.g. Caralluma, Duvalia, Echidnopsis, Edithcolea, Fockea, Hoodia, Huernia, Larryleachia, Orbea, Pectinaria,

Christenhusz, Fay & Chase

BORAGINALES

EUDICOTS

Piaranthus, Quaqua, Sarcostemmma, Stapelia and Tavaresia) are grown in specialist succulent collections for their peculiar habits and flowers. Monarch food: Asclepias species are the main food source for the caterpillars of the monarch butterfly in North America. As a result, the caterpillars accumulate glycosides and thus the larval and adults become poisonous to predators. Monarchs are migratory insects with the longest distance travelled, although they manage their migration in a period of several generations

during the year. They hibernate in forests of Abies religiosa (Pinaceae) in central Mexico, a region where Asclepias oddly is not abundant. Carnivory: The Malayan urn vine (Dischidia major) is an epiphyte and has two types of leaves, normal succulent circular ones and hollow, pouch-like leaves often called pitchers or ascidia. These have an opening at the top near their attachment point in which rainwater accumulates. Organic debris and insects also fall into the pit, and these decay, possibly

with the help of secreted fluids. The plant produces roots that grow into the chamber of the pitcher; in doing so the plant utilises the accumulated nutrients and water, which are otherwise not always available to an epiphyte. Even though it is more of a detritivore than a carnivore (it does not actively attract insects), it does benefit from insects that accidentally die in its pitchers. Etymology: Apocynum is composed of the Greek words από (apo), from, and κύων (kyon), a dog.

BORAGINALES Boraginales have been found to be sister to several orders including Lamiales in which they were included in the past. The sole family is Boraginaceae, treated here in the broadest sense. Some authors have proposed the division of this clade in up to 11 families, but APG IV did not see an advantage of such a treatment and maintained them as a single family. Boraginaceae were in some systems considered closely related to Lamiaceae based on the gynobasic style, but this character is clearly a parallelism, and there is no exclusive relationship between these families.

393. BORAGINACEAE Forget-me-not family

Boraginaceae are a family of annual and perennial herbs, shrubs, trees and vines. Plants are generally autotrophic and green, but some species are achlorophyllous, perennial, herbaceous parasites with scalelike, succulent, colourless leaves (Lennoa, Pholisma). Leaves are simple or pinnately divided, alternate or opposite in the lower part and alterate above, petiolate or sessile, the base sometimes sheathing or forming a wing along the stem, without stipules. Leaves are usually present along the stem but sometimes are all in a basal rosette. Leaf blades are

entire, lobed, crenate or dentate, sometimes pinnately dissected, and venation is pinnate or with several main veins at the base (pinnatepalmate), rarely palmate. The surface is often rough and hispidly hairy, usually with cystoliths (‘crystal sand’) at the base of these hairs, the hairs rarely stinging (Wigandia). Inflorescences are terminal, axillary or leafopposed, coiled cymes, sometimes several aggregated together in spikes, corymbs, panicles or heads, or the flowers are solitary. Flowers are bisexual (rarely unisexual) and generally actinomorphic, sometimes zygomorphic. The five or ten (rarely four or up to 13) sepals are free or basally fused, sometimes with basal appendages (Myosotis). The five (rarely four to ten or 11–13) petals are fused into a tube with the mouth often (partially) closed by a corona of scales in the throat protecting the nectar, sometimes in up to four whorls with a short fused tube (Hoplestigma). The corolla is actinomorphic, rotate or zygomorphic and bilabiate (Echium). Stamens are as many as petals (usually five, 20–35 in Hoplestigma) and fused at the

base to the corolla tube in between the petal lobes, free from each other or cohering at the anthers. Anthers are basifixed to dorsifixed and open by lengthwise slits, with or without connective appendages. The superior ovary is composed of usually two, sometimes four or five, carpels, forming two or four (rarely eight or ten) locules, usually via false septa. Styles are single or two or four, partially fused. The style emerges from a depression in the middle of the ovary at the base of the four locules (gynobasic) or at the top of an acute-tipped ovary. They have a stigma that is simple, two- or four-lobed, dry and papillate. Fruits are schizocarpic nutlets, capsules that may be dehiscent or not, or fleshy berries or drupes, usually with a persistent calyx. Distribution: This family has a nearly global distribution, occurring from the Arctic to the tropics and to Sub-Antarctic South America and New Zealand. They are most diverse in warm-temperate and semi-arid regions, such as southern North America (southern California, Texas and New Mexico), central Mexico, the Plants of the World

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EUDICOTS

Nama hispidum, New Mexico, USA (DZ) [393] Trichodesma zeylanicum subsp. grandiflorum, South Australia [393]

Borago officinalis, private allotment, Kingston upon Thames, Surrey, UK [393]

Ehretia dicksonii, Royal Botanic Gardens, Kew, UK [393]

Codon royenii, near Gannabos, northern Cape, South Africa (CD) [393]

Halgania cyanea, South Australia [393]

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Cynoglossum grande, Muir Woods, California, USA [393]

Lithospermum carolinianum, Old Mission Peninsular, Michigan, USA [393]

Symphytum ottomanum, Pindus Mountains, Greece [393]

BORAGINALES

EUDICOTS

Hydrophyllum virginianum, Royal Botanic Gardens, Kew, UK [393]

Heliotropium curassavicum var. oculatum, Santa Barbara, California, USA [393]

Pholisma arenarium, Kern County, California, USA (CD) [393]

Andes, the Mediterranean, South Africa and Australia.

Auxemma (2), Borago (3), Bothriospermum (5), Bourreria (c. 50), Brachybotrys (1), Brunnera (3), Caccinia (5), Cerinthe (10), Chionocharis (1), Choriantha (1), Codon (2), Coldenia (1), Cordia (c. 300), Craniospermum (6), Cryptantha (c. 100), Cynoglossopsis (2), Cynoglossum (c. 50), Cynoglottis (2), Cystostemon (16), Dasynotus (1), Decalepidanthus (1), Draperia (1), Echiochilon (15), Echiostachys (3), Echium (c. 60), Ehretia (c. 50), Ellisia (1), Embadium (3), Emmenanthe (1), Eriodictyon (9), Eritrichium (c. 30), Eucrypta (2), Gastrocotyle (1), Glandora (6), Gymnomyosotis (1), Gyrocaryum (1), Hackelia (45), Halacsya (1), Halgania (18), Harpagonella (1), Heliotropium (c. 400), Hesperochiron (2), Hoplestigma (2), Howellanthus (1), Hydrophyllum (8), Ivanjohnstonia (1), Lacaitaea (1), Lappula (40), Lasiarrhenum (3), Lasiocaryum (7), Lennoa (2), Lepechiniella (9), Lepidocordia (2), Lindelophia (11), Lithodora (7), Lithospermum (c. 55), Lobostemon (28), Macromeria (11), Maharanga (10), Mairetis (1), Melanortocarya (1), Memoremea (1), Menais (1), Mertensia (45), Microcaryum (4), Microula (30), Mimophytum (1), Moltkia (3), Moltkiopsis (1), Moritzia (5), Myosotidium (5), Myosotis (c. 100), Nama (45), Neatostema (1), Nemophila (17), Nesocaryum (1), Nihon (5), Nomosa (1), Nonea (41+1 extinct), Ogastemma (1), Omphalodes

(24), Omphalolappula (1), Oncaglossum (1), Onosma (150), Onosmodium (7), Oxyosmyles (1), Paracar yum (9), Pardoglossum (6), Patagonula (2), Pectocaryum (10), Pentaglottis (1), Phacelia (c. 100), Pholisma (3), Pholistoma (3), Plagiobotrys (c. 70), Pontechium (1), Pseudomertensia (8), Psilolaemus (1), Pulmonaria (19), Rindera (c. 25), Rochefortia (13), Rochelia (20), Romanzoffia (6), Saccellium (3), Sinojohnstonia (3), Solenanthus (c. 17), Stenosolenium (1), Stephanocaryum (2), Suchtelenia (2), Symphytum (35), Thaumatocaryon (4), Thyrocarpus (3), Tianschaniella (2), Tiquilia (27), Tournefortia (115), Trachelanthus (3), Trachystemon (2), Tricardia (1), Trichodesma (35), Trigonocaryum (1), Trigonotis (50), Turricula (1), Wellstedia (6) and Wigandia (3).

Phylogeny and evolution: Diversification of woody Boraginaceae may have taken place in the Mid Cretaceous in South America, although most age estimates are closer to 65 million years (Late Cretaceous), which seems more likely. Well-supported clades are found in Boraginaceae, which are best treated at the tribal level, although some authors prefer to treat them as families, probably to maintain Hydrophyllaceae, a family previously accepted (because of having capsular fruit, ebracteolate flowers, and attenuate style and pendulous ovules), but Hydrophyllaceae are deeply embedded in Boraginaceae with which they share many general characteristics. The African genera Wellstedia and Codon are sister to the rest of the Boraginaceae. Heliotropium and Tournefortia if broadly defined are both polyphyletic. Fossil Ehretia are known from Eocene Europe, where this genus does not currently occur. Genera and species: Boraginaceae include c. 135 genera and c. 2,535 species: Actinocarya (2), Afrotysonia (3), Alkanna (c. 30), Amblynotus (1), Amsinckia (15), Anchusa (c. 33), Anchusella (2), Ancistrocarya (1), Anoplocaryum (5), Antiotrema (1), Antiphytum (10), Arnebia (25), Asperugo (1),

Uses: Some species have edible fruit, although these are mostly eaten locally. Examples are cypre (Cordia alliodora), manjack (C. collococca), ziricote (C. dodecandra), kou (C. subcordata) and some heliotropes (Tournefortia species). Sand food (Pholisma sonorae), a parasitic plant, makes a swollen tuber that is usually buried in the sand. Only a circle of pale lilac or purple flowers emerges from the top, appearing to come directly out of the sand. The root is starchy and was consumed by Native Americans in Arizona. Borage (Borago officinalis) is occasionally Plants of the World

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BORAGINALES

532

EUDICOTS

Cordia sebestena, Playa Maya, Yucatán, Mexico [393]

Echium vulgare, Helsinki Botanical Garden, Finland [393]

Myosotis alpestris, Helsinki Botanical Garden, Finland [393]

grown as a pot herb for its edible flowers that taste like cucumber. It is commercially grown for its valuable oil (starflower oil) that is rich in gamma linoleic acids used for treating menstrual pain. It is also used as a flavouring in drink mixes such as claret cup. Camel bush (Trichodesma zeylanicum), introduced as food for camels and naturalised locally, is a potential oil-seed crop. Oysterplant (Mertensia maritima) is entirely edible, its hairless leaves are eaten as a vegetable with a slightly salty, mushroomy flavour. It is finding its way now into specialist cuisines and Michelin-star restaurants. The rhizomes are a staple diet for the Inuit. Young shoots of comfrey (Symphytum officinale) can be eaten like asparagus. Comfrey contains allantoin, which promotes healing, and it was therefore formerly used on wounds and mixed in plaster to heal broken bones. Scented flowers of huanita (Bourreria huanita) are used in Mexico to flavour tobacco and drinks, and Heliotropium flowers are cultivated for the perfume industry. Alkaloids are plentiful in Boraginaceae, some of which can be used medicinally or made into tea-like medicinal infusions such as yerba santa tea (Eriodictyon californicum) and Fukien tea (Ehretia microphylla). Ehretia aquatica contains psychoactive alkaloids that are used as a drug in religious ceremonies. Many species were (or still are) used to produce colourful dyes. Best known are

alkanet (Alkanna tinctoria, red dye), madwort (Asperugo procumbens, orange dye), golden drops (Onosma echioides, red dye), yellow puccoon (Lithospermum canescens, red dye), hairy puccoon (L. carolinense, red dye), and Indian paint (L. incisum, blue dye). The last three were often used as face paint by Native Americans, whereas Alkanna was sometimes used to colour poor-quality port. Ogastemma pusillum produces a red dye that is used for cosmetics in North Africa. A few species become large trees and produce valuable timber (Cordia, Ehretia, Patagonula) or firewood (Echiochilon). Leaves of some Cordia species are so rough and full of silica bodies that they can be used as sand paper. Species of Echium and Phacelia are occasionally sown as bee fodder for the production of honey. Many species are popular ornamental plants; examples include members of Amsinckia, Anchusa, Brunnera (B. macrophylla, Siberian bugloss), Cerinthe (C. glabra; C. major, honeywort), Cordia (C. sebestena, geiger tree), Cryptantha, Cynoglossum (hound’s tongue), Echium (viper’s bugloss), Ehretia, Eritrichium, Heliotropium (e.g. H. arborescens, heliotrope), Hesperochiron, Lindelophia, Lithodora, Lithospermnum (stoneseed), Lobostemon, Mertensia (M. maritima, oysterplant; M. virginica), Moltkia, Myosotidium hortensia (Chatham Island forget-me-not), Myosotis (e.g. M. sylvestris, garden forget-me-not), Nemophila

(N. menziesii, baby-blue-eyes), Omphalodes (e.g. O. verna, blue-eyed Mary), Onosma (O. frutescens, golden drop), Paracaryum, Phacelia, Pulmonaria (lungwort), Romanzoffia, Symphytum (comfrey), Trachystemon and Wigandia. Forget-me-nots were popularised in Samuel T. Coleridge’s poem The Keepsake (c. 1800). Pentaglottis sempervirens escapes from cultivation and can become a noxious weed disliked by gardeners because of its irritating hairs. Similarly Echium plantagineum (Paterson’s curse), a Mediterranean native, has invaded large swathes of Australia, reducing productivity for cattle and suppressing native vegetation.

Christenhusz, Fay & Chase

Parasites: The parasitic genera Lennoa and Pholisma, previously recognised as the separate family Lennoaceae, are placed in the otherwise non-parasitic family Boraginaceae. They are morphologically closely related to Ehretia but lack chlorophyll. They have congested inflorescences, pentato decamerous f lowers and scale-like, spirally arranged leaves. They can be found from California and Arizona to northern Colombia, where they typically parasitise roots of Clematis (Ranunculaceae), Euphorbia (Euphorbiaceae) and shrubby Asteraceae. Etymology: Borago is composed of the Latin burra, a furry garment, and -ago, like, in reference to the hoary leaves of borage.

VAHLIALES

EUDICOTS

VAHLIALES This is an unplaced order, which is likely to be close to Boraginales or Solanales, but seems unlikely to be included in either of them. Vahlia resembles some members of Saxifragales in which it was previously placed.

394. VAHLIACEAE Flindersbush family

This family consists of annual and biennial, bisexual herbs that are sometimes glandular. Leaves are simple, opposite, sessile, without stipules, and have an entire margin with obscure, pinnate venation. Inflorescences are terminal or axillary cymes, or the flowers are in pairs. Flowers are bisexual and actinomorphic, with five, free sepals and five free, ovate-spathulate petals. The five stamens alternate with the petals and are free from them and each other. Anthers are

dorsifixed, opening inwardly by lengthwise slits. The inferior ovary is composed of two or three carpels fused to form a single locule. The ovary is topped by two or three, thick, divaricating styles, each topped by a capitate stigma. Fruits are dry, dehiscent capsules, opening at the tip by two or three valves exposing numerous, appendaged seeds. Distribution: The family is widespread across periodically dry Sub-Saharan Africa, Madagascar, Sri Lanka and on the Indian subcontinent.

asterids, probably near Solanales, although its precise position remains uncertain. The older name Bistella has been rejected against Vahlia. Genera and species: The sole genus in Vahliaceae is Vahlia with eight species. Etymology: Vahlia is named for the Norwegian-born Danish botanist and zoologist Martin Vahl (1749–1804).

Vahlia capensis, Royal Botanic Gardens, Kew, UK [394]

Phylogeny and evolution: An Upper Cretaceous fossil from Sweden (Scandianthus) has been associated with Vahlia, but its characters do not fully agree, and this fossil may be better placed in Cornales. Vahlia was previously included in Saxifragaceae, with which it has some similarities, particularly in ovary characteristics, but molecular evidence has placed it in the

Vahlia capensis, habit, South Africa (NH) [394]

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SOLANALES

EUDICOTS

SOLANALES Families 395 to 399 comprise Solanales, an order that diversified c. 76–78 million years ago, with some analyses indicating that it may already have evolved c. 100 million years ago. A key character in this order is the accrescent calyx in fruit. The order includes five families, with Solanaceae and Convolvulaceae together sister to Montiniaceae, Sphenocleaceae and Hydroleaceae.

395. CONVOLVULACEAE Bindweed family

Convolvulaceae are a family of mostly bisexual and some unisexual (Hillebrandtia), annual and perennial climbing and trailing herbs, shrubs, lianas and occasionally trees. Sometimes plants are self-supporting and not climbing, or in Cuscuta, the plants are nearly leafless, rootless and parasitic, with roots (haustoria) that form coils that pierce the stems of the host plant. Climbers twine anti-clockwise around supports. Leaves are simple or compound, alternate or spiralling around the stem, lacking stipules. Blades are cordate, hastate, sagittate or pinnatifid or palmatifid-divided, sometimes finely so, sometimes only shallowly lobed. Margins are entire or lobed, and venation is pinnate or palmate or a combination of these. Inflorescences are axillary cymes or umbels, or flowers are solitary, usually with bracts and bracteoles, sometimes aggregated along the stem in clusters (Cuscuta) or epiphyllous (Neuropeltis). Flowers are bisexual or unisexual and actinomorphic (zygomorphic in Humbertia). The (three to) five sepals are free or rarely fused and persistent in fruit. The (three to) five petals are fused into a valvate or contorted, plicate, funnel-, tube- or cup-shaped corolla. The five (or ten in Cuscuta) stamens are inserted on the corolla tube near the base or midway, opposite the sepal lobes, in Cuscuta often alternating with fringed scales. Anthers 534

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are basifixed to dorsifixed and open inwardly by lengthwise slits. The superior ovary is composed of usually two (rarely up to five) carpels, forming (one) two (or five) locules. It is topped with one, two (to five) free or partially joined styles, topped by dry stigmas. A circular nectar disk is usually present around the style. Fruits are aggregated nutlets or berries, sometimes loculicidal, circumscissile or irregularly opening capsules. Seeds are sometimes conspicuously hairy. Distribution: The family has a global distribution but is more diverse in warm temperate and tropical areas; it is absent from the polar regions and the great deserts. Phylogeny and evolution: Six subfamilies are sometimes recognised: Humbertioideae, Eryciboideae, Cuscutoideae, Convolvuloideae, Dichondroideae and the Cardiochlamys clade. The first four form a grade leading to a clade uniting the last two. All except the cardiochlamydoids have at some stage been accepted at the family level, but they are all closely related and share characters in seeds, pollen, flowers, vining habit and palmate leaves etc. The family most likely originated in the Old World tropics, probably in Madagascar or tropical Asia. The crown group of Convolvulaceae has been estimated to have diversified c. 68 million years ago. Some generic circumscriptions need to be revised, e.g. Calystegia (only differentiated by its enlarged bracteoles) is embedded in Convolvulus, and Merremia is polyphyletic. Genera and species: Convolvulaceae include c. 57 genera and c. 1,660 species: Aniseia (4), Argyreia (130), Astripomoea (12), Blinkworthia (2), Bonamia (57), Calycobolus (28), Camonea (5), Cardiochlamys (2), Convolvulus (254), Cordisepalum (2), Cressa

(4), Cuscuta (201), Daustinia (1), Davenportia (1), Decalobanthus (1), Dichondra (15), Dicranostyles (15), Dinetus (8), Dipteropeltis (3), Distimake (35), Duperreya (3), Erycibe (72), Evolvulus (98), Falkia (3), Hewittia (2), Hildebrandtia (14), Humbertia (1), Hyalocystis (2), Ipomoea (c. 425), Iseia (1), Itzaea (1), Jacquemontia (65), Keraunea (1), Lepistemon (6), Lepistemonopsis (1), Lysiostyles (1), Maripa (20), Merremia (18), Metaporana (6), Nephrophyllum (1), Neuropeltis (13), Neuropeltopsis (1), Odonellia (2), Operculina (15), Paralepistemon (1), Polymeria (11), Porana (2), Poranopsis (3), Rapona (1), Remirema (1), Rivea (3), Seddera (31), Stictocardia (13), Stylisma (6), Tetralocularia (1), Tridynamia (4), Turbina (14), Wilsonia (3) and Xenostegia (5). Uses: Sweet potato (Ipomoea batatas) is a cultigen originating in Central America but that has been spread to the Pacific and New Zealand (it was a staple food of the Maori) before European exploration of Polynesia. It was then spread across the globe by Spanish settlers. It is probably derived from wild species such as I. trifida and I. leucantha, which both have edible tubers. It has been crossed with I. trifida to increase nematode resistance, and it is now a major staple crop around the tropics and beyond. It is also used to produce bioplastics and alcohol. The epithet batatas derives from a Malay word for yam (Dioscorea) and has in the early years of discovery been applied by Europeans to a variety of tropical root crops. The common name of Solanum tuberosum (Solanaceae), potato, is derived from the Malay ‘batat’. Yala (I. costata) has edible tubers formerly eaten by the Aboriginals of Australia, and the whitestar potato (I. lacunosa) and wild potato vine (I. pandurata) were eaten by natives in North America. The shoots of kankong or ong tsoi

SOLANALES

EUDICOTS

(I. aquatica) are commonly eaten as a vegetable (water spinach) in Asia. Rhizomes and young leaves of common bindweed (Convolvulus sepium) are eaten in China and India. Hallucinogenic Rivea corymbosa was smoked by the Aztecs. Seeds of Mexican morning glory (I. tricolor) and ololiuqui (Turbina corymbosa) also have hallucinogenic properties. The roots of rhodium wood (Convolvulus scoparius) and Canary Island guadil (C. floridus) yield an essential oil (lignum rhodium) used medicinally. An extract from the seeds of Cuscuta chinensis (a parasite of soybean plants) can be used to treat acne and dandruff. The timber of Humbertia madagascariensis is hard and scented like sandalwood.

There are many species grown for their ornamental value, either as perennials or annuals, trellis vines or bedding plants etc. Examples are elephant climber (Argyreia nervosa), silverbush (Convolvulus cneorum), blue rock-bindweed (C. sabatius), pony’s foot (Dichondra micrantha), kidneyweed (D. repens), moonflower (Ipomoea alba), tuberous morning glory (I. batatas), redstar (I. coccinea), blue morning glory (I. indica), firevine (I. lobata), ivy morning glory (I. nil), common morning glory (I. purpurea), cypress vine (I. quamoclit), cardinal climber (I. sloteri), Mexican morning glory (I. tricolor), beach moonf lower (I. violacea), yellow morning glory (Merremia tuberosa), bridal bouquet (Poranopsis paniculata), African

Convolvulus althaeoides, near Ronda, Cuscuta epithymum, flower detail, Perth, Western Spain [395] Australia [395]

Dichondra repens, flower detail, Sicily, Italy [395]

morning glory (Stictocardia beraviensis) and Christmas vine (Turbina corymbosa). Introduced to Japan in the 16th century by the Portuguese, Ipomoea nil has become popular in Japan, especially during the Edo period, when elaborate selections and cultivars with highly divided and flamed coronas were developed. They are commonly depicted on Japanese screens, and exhibitions of Ipomoea at which these unusual cultivars are grown in the traditional way remain popular today. A number of Convolvulaceae are troublesome garden weeds, especially Cuscuta spp., Convolvulus arvensis and C. sepium. Etymology: Convolvulus is derived from the Latin convolva, to twine around.

Cuscuta victoriana, near Coober Erycibe tomentosa in fruit, Pedy, South Australia [395] Singapore (WA) [395]

Ipomoea batatas, Helsinki Botanical Ipomoea aquatica, Singapore Botanical Garden, Finland [395] Garden [395]

Evolvulus alsinoides, Taita Hills, Kenya [395]

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SOLANALES

396. SOLANACEAE Nightshade family

These are a family of annual and perennial herbs, shrubs, trees and climbers, rarely epiphytes. Stems are sometimes prickly or thorny, and they are often covered with simple, branched, stellate or glandular hairs. Leaves are simple or pinnately compound, alternate, solitary, paired or clustered, sometimes in a basal rosette, petiolate or not and lacking stipules. Leaf blades have entire, toothed, lobed or dissected margins and pinnate venation. Inf lorescences are terminal, often overtopped by continuing axes and then appearing axillary, frequently appearing randomly from the stem or leaf opposed. They are usually racemes, panicles, umbels or clusters, rarely true cymes, or flowers are solitary. Bracts are sometimes present. Flowers are mostly bisexual and generally actinomorphic, sometimes weakly to strongly zygomorphic. The five (rarely four to nine) fused sepals are present along with a similar number of petals, the latter fused into a cup-, funnel- or tube-shaped corolla, often deeply lobed or the margins fringed or dissected. Stamens are as many as petal lobes (usually five), alternating with them and fused to the inside of the petal-tube. They are all similar or one to several stamens reduced in size. Anthers are dorsifixed and open by lengthwise slits or apical pores. The superior ovary is composed of two to five carpels, each forming a locule topped by a single style and capitate stigma. Fruits are berries or capsules surrounded by the persistent, often enlarged sepals. Distribution: Solanaceae have a global distribution but are absent in the Arctic, Antarctic and the great deserts of the Sahara

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EUDICOTS

and Arabia. They are widespread in temperate and tropical regions but most diverse in the western Neotropics. Phylogeny and evolution: Solanaceae have been estimated to have diverged c. 30–58 million years ago, with the crown group, Solanoideae including the bulk of the species, evolving c. 21 million years ago, with Solanum itself being perhaps 15.5 million years old. Fossils attributed to Solanaceae are all dubious. Solanaceae originated in the Americas with a good number of dispersal events (up to nine) to the Old World. There are eight clades in Solanaceae, sometimes recognised at the subfamily level, but the relationships among them are still somewhat uncertain. The eight subfamilies are: Schizanthoideae, Goetzeoideae, Duckeodendron+Reyesia (as yet not formally named), Browallioideae, Schwenckioideae, Petunioideae, Nicotianoideae and Solanoideae. Some generic reorganisation has recently been taking place. Atrichodendron, formerly placed in Solanaceae, does not belong here, but the specimen on which it was based is too poorly preserved to say to which family it should be assigned. Genera and species: Solanaceae include c. 100 genera and c. 2,600 species: Acnistus (1), Alkekengi (1), Anisodus (4), Anthocercis (11), Anthotroche (4), Athenaea (7), Atropa (3), Atropanthe (1), Aureliana (8), Benthamiella (12), Bouchetia (2), Brachistus (3), Browallia (9), Brugmansia (10), Brunfelsia (47), Calibrachoa (27), Calliphysalis (1), Capsicophysalis (1), Capsicum (36), Cestrum (c. 123), Chamaesaracha (7), Coeloneurum (1), Combera (2), Crenidium (1), Cuatresia (20), Cyphanthera (7), Darcyanthus (1), Datura (11), Deprea (9), Discopodium (2), Duboisia (4), Duckeodendron (1), Dunalia (5), Dyssochroma (3), Eriolarynx (3), Espadaea (1), Exodeconus (6), Fabiana (15), Goetzea (2), Grammosolen (3), Henoonia (1), Heteranthia (1), Hunzikeria (3), Hyoscyamus (23), Iochroma (24), Jaborosa (23), Jaltomata (57), Juanulloa (11), Larnax (30), Latua (1), Leptoglossis (7), Leucophysalis (2), Lycianthes (c. 100),

Lycium (91), Mandragora (3), Margaranthus (1), Markea (19), Melananthus (5), Mellissia (1), Merinthopodium (3), Metternichia (1), Nectouxia (1), Nicandra (1), Nicotiana (>76), Nierembergia (16), Nolana (91), Nothocestrum (4), Oryctes (1), Pantacantha (1), Petunia (18), Physaliastrum (9), Physalis (>75), Physochlaina (8), Plowmania (1), Protoschwenckia (1), Przewalskia (1), Quincula (1), Reyesia (4), Salpichroa (c. 15), Salpiglossis (2), Saracha (2), Schizanthus (12), Schraderanthus (1), Schultesianthus (5), Schwenckia (20), Sclerophylax (13), Scopolia (2), Sessea (22), Solandra (10), Solanum (c. 1,300), Streptosolen (1), Symonanthus (2), Trianaea (6), Tsoala (1), Tubocapsicum (1), Tzeltalia (3), Vassobia (2), Vestia (1), Withania (c. 20) and Witheringia (14). Uses: Economically, the most important crop of Solanaceae is the potato (Solanum tuberosum), a cultigen derived c. 8,000 years ago from wild species (probably S. brevicaule) in the Andes and from which the alkaloids and bitterness have been bred out. In 1537, it became known to the Spanish, who exported it to many parts of the world (grown in Ireland as early as 1566), and it soon became a staple food. It provides most nutrients needed for a humanbeing to thrive. Potatos are boiled, mashed or fried (chips, crisps) or used in pancakes, casseroles, salads, bread, or sometimes mashed with other vegetables. Potato starch can be used as a clear, tasteless food binder and is one of the main sources of commercial starch for industry, for instance to make wallpaper glue and biodegradable plastics. It is also distilled into alcoholic drinks such as Irish poitín, vodka and schnapps. Tomatoes (Solanum lycopersicum) are second in importance to potatoes in this family. These were originally weeds of the maize and bean fields of the Aztecs in Mexico, but when discovered and introduced into Europe soon became popular for sauces (e.g. ketchup), soups and juice. They are also eaten fresh in salads. Yellow forms were already cultivated in Italy in 1544 (and then called pomi d’oro, golden apples). Lycopene in the fruits helps reduce the likelihood of developing various

SOLANALES

EUDICOTS

Anthocercis viscosa, Albany, Western Australia [396]

Capsicum frutescens, Emsflower, Germany [396]

Exodeconus miersii, Galápagos Islands [396]

Atropa belladonna, private garden, Kingston upon Thames, Surrey, UK [396]

Brugmansia sanguinea, Ecuador [396]

Benthamiella patagonica, Royal Horticultural Society Garden, Wisley, UK [396]

Cestrum aurantiacum, private garden, Datura stramonium, Ecuador [396] Kingston upon Thames, Surrey, UK [396]

Espadaea amoena, Fairchild Tropical Botanical Garden, Florida, USA (CD) [396]

Nicotiana benthamiana, Judbara-Gregory, Nicotiana tabacum, Helsinki Northern Territory, Australia [396] Botanical Garden, Finland [396]

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SOLANALES

EUDICOTS

Jaborosa sativa, private garden, Kingston upon Thames, Surrey, UK [396] Hyoscyamus aureus, Baalbek, Lebanon [396]

538

Mandragora autumnalis, Royal Botanic Gardens, Kew, UK [396]

Lycium europaeum, Royal Botanic Gardens, Kew, UK [396]

Goetzea elegans, Fairchild Tropical Botanical Garden, Florida, USA (SZ) [396]

Iochroma fuchsioides, San Francisco Botanical Garden, California, USA [396]

cancers, especially prostate cancer. The seeds are rich in oils and can be pressed into cooking oil or used for soap. The leaves are high in solanine and make a good insecticide. Eggplant or aubergine (Solanum melongena) is originally from Africa but was first domesticated in Southeast Asia. It is an important vegetable in many cuisines, especially Thai food and Greek moussaka. However, when grown during drought, aubergines may accumulate alkaloids and can cause nightmares. Bitter berry (S. aethiopicum) has small fruits that are bitter, but its fruits and leaves are important vegetables in Africa. Similarly, young branches and leaves of glossy nightshade (S. americanum) and some cultivars of black nightshade (S. nigrum) are cooked and

eaten like spinach. Borodina or cannibal’s tomato (S. viride ‘Anthropophagorum’) was domesticated in ancient Fiji, where it was used to wrap human flesh for baking. A sauce of the fruit (‘cannibal chutney’) was supposedly eaten with the meat. Several species of Physalis also produce edible berries, especially common are Cape gooseberry (P. peruviana), tomatillo (P. philadelphica) and strawberry tomato (P. pruinosa). Chinese lantern (P. alkekengi) has fruits that are edible as well, but it is mainly grown as a cut flower or everlasting flower. Species of Capsicum (chili peppers) have been cultivated in the Americas for over 6,000 years and were introduced to other parts of the world during the early 16th century when they soon became part

of local cuisine. It is difficult to imagine Asian cuisine without chilies and Hungarian goulash without paprika. Most cultivated forms are derived from Capsicum annuum, which has five main groups: the bell peppers or paprika and pimento (Grossum Group), which are usually not hot, whereas the others, cherry peppers (Cerasiforme Group), red cone peppers (Fasciculatum Group), cone peppers (Conoides Group) and cayenne and chili peppers (Longum Group) with upright or pendent pungent, spicy fruits, are mostly used as condiment. Spontaneous (wild?) forms of C. annuum are called bird peppers and usually classified as var. glabriusculum. Other species of Capsicum grown as a condiment are the hot chilies often referred to as C. frutescens, which are mostly used for hot sauces. This

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SOLANALES

EUDICOTS

Petunia exserta, Royal Botanic Gardens, Kew, UK [396]

Nolana paradoxa, Ruissalo Botanical Garden, Turku, Finland [396]

Streptosolen jamesonii, Adelaide Botanic Garden, South Australia [396]

Physalis alkekengi, private garden, Hengelo, the Netherlands [396]

Schizanthus hookeri, La Laguna Maule, Chile (CD) [396]

Scopolia carniolica, Helsinki Botanical Garden, Finland [396]

includes cultivars such as C. frutescens ‘Piri Piri’ and ‘Tabasco’. The hottest peppers are the cultivars ‘Naga Jolokia’ or ghost pepper (800,000–1,001,304 Scoville Hotness Units, SHU), ‘Trinidad Scorpion Butch’ (1,463,700 SHU), ‘Trinidad Moruga Scorpion’ (2,009,231 SHU), and ‘Carolina Reaper’ (1,569,383– 2,200.000 SHU), but these are mostly used for pepper spray. Locoto pepper (C. pubescens) is used for salsas and as a local spice in Latin America, especially Bolivia. Sweet or bell peppers are a major vegetable, cultivated around the world. Capsaicinoids, the compounds that make chillies hot, have evolved to protect fruits from attack by fusarium fungi. Goji berry (Lycium chinense) is high in vitamin C and promoted as a health food. This

and other species of boxthorn (L. barbarum and L. ferocissimum) can become invasive. They are also planted to stabilise sand. Minor fruits locally eaten fresh or for juice are tree tomato or tamarillo (Solanum betaceum), sunberry (S. ×burbankii), chambala (S. cajanumense), bush tomato (S. cinereum), Indian nightshade (S. indicum), naranjilla (S. quitoense), red nightshade (S. nigrum ‘red makoi’), huckleberry (S. scabrum), cocona (S. sessiliflorum), susumber (S. torvum), tlanochtle (Lycianthes moziniana), pepino (L. (Solanum) mucronatum), pale wolfberry (Lycium pallidum) and many more. Leaves of tobacco, Nicotiana, are chewed or smoked as a stimulant, but this is incredibly bad for your health, the tar constricting blood vessels causing heart attacks, hydrochloric

acid causing gastric ulcers, carbon monoxide reducing oxygen uptake by the blood and nicotine causing methylation of DNA, resulting in tumours and cancer. Nicotine is also highly addictive, and tobacco is produced and consumed worldwide, particularly in China, the Middle East, tropical Africa, the USA and tropical America (e.g. Cuban cigars). The most commonly grown cultivars are forms of N. tabacum, a natural allotetraploid hybrid of N. sylvestris and N. tomentosiformis. In pre-Columbian times, Aztec tobacco (N. rustica), another allotetraploid species of hybrid origin (N. paniculata × N. undulata) was more commonly smoked in North America. Other species are smoked locally (in Latin America) or chewed (especially by Australian Aboriginal peoples). Plants of the World

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SOLANALES

540

EUDICOTS

Schwenckia americana, Tobati, Paraguay (CD) [396]

Solanum tuberosum, private allotment, Kingston upon Thames, Surrey, UK [396]

Solandra grandiflora, cultivated in Sidon, Lebanon [396]

Solanum betaceum, Royal Botanic Gardens, Kew, UK [396]

Apart from nicotine, Solanaceae are well known for their complex secondary compounds, especially tropane alkaloids. All parts of deadly nightshade (Atropa belladonna) are poisonous, even though the berries have a pleasant taste, which may result in accidental poisonings. It is high in hyoscyamine and lower in atropine. The latter alkaloid has an effect on the nerves and was formerly used by women to enlarge their pupils and make them look more beautiful (hence belladonna), but often with partial blindness as a result. Today, deadly nightshade is used in eye operations, cough medicine and nerve gas antidote. Brugmansia, Datura, Duboisia, Hyoscyamus, Mandragora, Scopolia and Withania are high in scopolamine, atropine and hyoscyamine, which may lead to intense hallucinations and intoxication. Their use is dangerous and often goes wrong, but Brugmansia species have been employed by native American shamans for centuries. Datura ceratocaula and D. innoxia were sacred herbs to the Aztecs and other Native Americans. Thorn-apple (D.

stramonium), now widely naturalised around the temperate zones, was used by North American tribes in transition rites for boys entering adulthood. The leaves and seeds are strongly hallucinogenic and deliriant, but the plant has been cultivated as a medicine for asthma and Parkinson’s disease (stramonin). In coastal Brazil, inkberry (Cestrum laevigatum) is also used in shamanic rites by the Craós tribe. The effect is somewhat like that of cannabis, but it is not to be taken by the non-initiated. Herbal mixtures including henbane (Hyoscyamus albus, H. muticus) and mandrake (Mandragora officinarum), and later also Datura stramonium, were ground and mixed with grease in wicca rituals in pre-Mediaeval Europe. This ‘witch’s cream’ was spread on places where the skin is thin (such as arm pits) giving the sensation of flying (perhaps the origin of the witch on a broomstick). Henbane and other Hyoscyamus species have been applied as a pain killer since Neolithic times and were smoked in England as a recreational, hypnotic and hallucinogenic

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Solanum phoxocarpum, Mt Kenya, Kenya [396]

drug until the early 20th century. Chilean Latua pubiflora, also called ‘sorcerer’s tree’ or ‘palo de los brujos’, contains high concentrations of many alkaloids and was formerly used as a punishment, causing permanent madness in victims. Nicotinoids, solanines and atropines are effective systemic insecticides for plants, but they also kill the bees that pollinate treated plants. Solasodine, mostly commercially obtained from Solanum viarum, is a precursor in the production of complex steroids such as those used in contraceptive pills. Solanaceae have many species with large and attractive flowers that are commonly grown as garden ornamentals, houseplants and bedding plants. Examples of commonly grown ornamentals are bush violet (Browallia speciosa), angel’s trumpets (especially Brugmansia ×candida), yesterday, today and tomorrow (Brunfelsia spp.), million bells or dwarf petunias (Calibrachoa ×hybrida), orange jessamine (Cestrum aurantiacum), day-blooming jessamine (C. diurnum), lady-ofthe-night (C. noctiflorum), green poison-berry

SOLANALES

EUDICOTS

(C. parqui), pichi (Fabiana imbricata), violet tubeflower (Iochroma cyaneum), blue potato bush (Lycianthes rantonnetii), shoo-fly plant (Nicandra physalodes), jasmine tobacco (Nicotiana alata), shrub tobacco (N. glauca), Langsdorff’s tobacco (N. langsdorffii), two-coloured tobacco (N. mutabilis), flowering tobacco (N. sylvestris), ornamental tobacco (N. ×sanderae), white cup (Nierembergia rivularis), cupflower (N. scoparia), Chilean bell flowers (Nolana spp.), petunia (Petunia ×atkinsiana, a hybrid between P. axillaris and P. integrifolia), Chinese lantern (Physalis alkekengi), cock’s-eggs (Salpichroa origanifolia), painted tongue (Salpiglossis sinuata), poor man’s orchid (Schizanthus pinnatus), chalice vine (Solandra maxima), winter cherry (Solanum capsicastrum, S. pseudocapsicum), potato vine (S. laxum) and macaw bush (S. mammosum), to name a few. Killer tomatoes: Various members of Solanaceae have hairs that catch small insects. Darwin already reported that tobacco (Nicotiana tabacum) is covered with sticky hairs but, in his experiments with infusions of meat and ammonium carbonate, he produced no convincing evidence of absorption. However, later studies found that species of Petunia have proteolytic enzymes, but were unable to absorb the nutrients through their leaves. Potato (Solanum tuberosum) and species of wild tomatoes also secrete proteinases from their leaves. Even though Kaliphora madagascariensis, Antsirabe, Madagascar (CD) [397]

they may not be able to absorb nutrients directly, the nutrients do run down into the soil, where the plants can absorb them. Solanaceae are suitable plants to include in studies of the various degrees of carnivory found in the plant kingdom. From the seed of a hanged man: Mandrake (Mandragora officinarum) has tuberous roots that resemble a human form, and it has therefore been long associated with myths. It grows large orange-yellow fruits that may have been the golden apples of Aphrodite (see also Cydonia, Rosaceae, and Citrus, Rutaceae) or the ‘apples of Sodom’ (a name also given to some Solanum species). The foetid smell of these plants was associated with fertility, and it was often referred to as ‘dudaim’, or love-plant, for instance in the Bible, where it is also associated with fertility (Genesis 30:14). However, other magical properties were also attributed to mandrake. When the roots are dug up, the plants screams (so it was difficult for Harry Potter and his fellow students to transplant the baby plants into new pots). Many beneficial properties have been attributed to these roots; they were dressed in specially made clothes and treasured as amulets. It was often the ‘last resort’ medicine in Mediaeval times. But where to find plants of mandrake? It cannot be collected anywhere; the best mandrake can be found on a hill near the gallows because it was said to sprout from the semen of a hanged man.

Montinia caryophyllacea, female, South Africa (FF) [397]

When does extinct really mean extinct? Mellissia begoniifolia (St. Helena boxwood) is endemic to the island of St Helena in the South Atlantic Ocean. After not being seen since the 19th century, it was rediscovered in 1998, a remarkable situation on a small, wellstudied island. Molecular studies show that it is closely related to Withania. It is currently being propagated for reintroduction at the Royal Botanic Gardens, Kew. Etymology: Solanum is the classical Latin name for nightshade, possibly from ‘solare’, to sooth or comfort, for its medicinal use.

397. MONTINIACEAE Wild-clovebush family

These are bisexual shrubs and small trees with simple, alternate to opposite leaves without stipules. Blades have entire margins and pinnate-arching venation. Male inflorescences are few-flowered, corymbose terminal or axillary cymes, and female flowers are solitary in the leaf axils. The

Montinia caryophyllacea, male, Kogelberg Nature Reserve South Africa (CD) [397]

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541

SOLANALES unisexual flowers are actinomorphic. Male flowers have three to five fused sepals with entire margins and three to five free petals. The three to five stamens alternate with the petals, the filaments thick and short and anthers relatively large, dorsifixed and opening outwardly by lengthwise slits. Female f lowers have four or five fused sepals and the same number of free petals. The inferior ovary is composed of two fused carpels, each forming a locule. A nectar disk is fleshy and angled, surrounding a single short style with two large stigmas. Fruits are dehiscent or indehiscent capsules.

EUDICOTS

Uses: The pungent scent of the crushed leaves of Grevea and Montinia caryophyllacea are used to soothe colds and headaches. They have a peppery, clove-like taste. Etymology: Montinia is named for Swedish botanist and physician Lars Jonasson Montin (1723–1785). He was a disciple of Linnaeus and studied snow grouse in Lapland and plants in Halland, Sweden.

398. SPHENOCLEACEAE Gooseweed family

Genera and species: Montiniaceae include three genera and five species: Grevea (3), Kaliphora madagascariensis and Montinia caryophyllacea. Sphenoclea zeylanica, Colombia (MF) [398]

These are annual, succulent herbs with alternate, simple leaves and no stipules. Leaf blades have pinnate venation and entire margins. Inflorescences are bracteate, terminal or leafopposed, dense spikes. Flowers are sessile, bisexual, actinomorphic and subtended by two bracteoles. The five sepals and five petals are fused into a calyx and a tubular corolla, split to the middle and forming five lobes. The five stamens are inserted in the upper part of the corolla tube and alternate with the petals. Filaments are short, making the anthers appear nearly sessile. Anthers are basifixed, opening by slits outwardly. The inferior ovary is composed of two fused carpels, each forming a locule, the ovary is topped by a single style and a faintly bilobed, capitate stigma. Fruits are circumscissile capsules with numerous oblong seeds surrounded by the persistent calyx. Distribution: This family is widely distributed in the Old World tropics, throughout Sub-Saharan Africa, Madagascar, and from India and China throughout tropical Asia to northern Australia. They have been introduced in the Neotropics. They grow in permanently wet habitats.

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Genera and species: The sole genus in this family is Sphenoclea with two species, S. pongatium and S. zeylanica. Uses: Gooseweed (Sphenoclea zeylanica) is a common weed of rice paddies and suppresses the growth of the crop. In Asia, young shoots of gooseweed are sometimes cooked like spinach.

Distribution: Montiniaceae are restricted to Sub-Saharan Africa and northern Madagascar. Phylogeny and evolution: The crown group of Montiniaceae is estimated to be c. 42 million years old. The family has been hard to place on morphological grounds, but the genera were generally included in a broad concept of Saxifragaceae because of their two carpels. Kaliphora was sometimes placed in its own family, but molecular studies placed it with Montiniaceae in Solanales.

Phylogeny and evolution: Sphenocleaceae are only superficially similar to Campanulaceae, with which they used to be placed. DNA sequence analyses place them securely in Solanales in a clade with Montiniaceae and Hydroleaceae. The main differences from Campanulaceae are that they lack latex and have a short, glabrous style and circumscissile fruit dehiscence.

Etymology: Sphenoclea is composed of the Greek words σφήνος (sfenos), wedge, and κλείω (kleio), to shut or enclose, in reference to the capsule.

399. HYDROLEACEAE Fiddleleaf family

Hydroleaceae include annual and perennial herbs and shrubs, usually growing in wet habitats. Stems have spines at their nodes. Leaves are simple, alternate and lack stipules, and blades have a pinnate venation with entire margins. Inflorescences are usually corymbs on lateral shoots or terminal, leafy panicles or axillary fascicles, or flowers are solitary. Flowers are bisexual and actinomorphic. The five sepals are only basally fused and persistent, enlarging in fruit. The five petals are fused into a broadly campanulate corolla, with the five, free stamens inserted in the tube alternating with the petal lobes. Anthers are dorsifixed and open by longitudinal slits. A

LAMIALES

EUDICOTS

nectar disk is absent. The superior ovary is composed of two (or up to five) carpels, each forming a locule. The two free styles (rarely three to five) are glabrous, glandular or hairy, with funnel-shaped stigmas. Fruits are capsules that open septicidally, loculicidally or irregularly with numerous tiny seeds.

Uses: The leafy shoots of Hydrolea zeylanica are used as a potherb in Indonesia. Etymology: Hydrolea is derived from Greek ύδωρ (hydor), water and ελαιά (elaia), olive, in reference to the watery habitat and leaves that resemble those of an olive tree.

Distribution: This is a pantropical family, extending into the warm temperate zones of eastern North America, southeastern South America and temperate East Asia. Phylogeny and evolution: Previously included in Hydrophyllaceae (now Boraginaceae), they were found to be distantly related in molecular studies. They differ mainly in their spiny habit, its axile (not parietal) placentation and characters in the embryo, but they are easily mistaken for Boraginaceae. Genera and species: The single genus of this family is Hydrolea with c. 11 species.

Hydrolea spinosa, Colombia (MF) [399]

Hydrolea spinosa, Santa Catarina, Brazil [399]

LAMIALES Families 400 to 422 belong to Lamiales, an order in which there are some well-established families, mostly originally based on characters of ovule placentation, fruit type and position of the style on the ovary. Molecular evidence has shown these families to be closely related, with Plocospermataceae, Carlemanniaceae, Oleaceae and Tetrachondraceae forming successive sister clades to the rest. Among the rest, family circumscriptions have been in great flux since the dawn of molecular studies. It was found that a number of genera have been misplaced on the basis of their morphology. Families such as Verbenaceae and Scrophulariaceae were redefined to exclude a number of genera and include some genera formerly placed elsewhere (e.g. Buddleja, formerly in Loganiaceae, is now placed in Scrophulariaceae). The biggest changes were in membership of the formerly large Scrophulariaceae, which were placed all over the Lamiales tree, redefining Gesneriaceae, Plantaginaceae and Orobanchaceae. Redefinition and subsequent splitting to maintain established names is the cause of many new, usually small families such as Mazaceae, Paulowniaceae, Schlegeliaceae and Thomandersiaceae. Lamiaceae, traditionally defined by a gynobasic style, now includes several genera without this, formerly placed in Verbenaceae (e.g. Vitex). Acanthaceae, formerly defined by their explosively schizocarpic fruit, now include Thunbergia, Nelsonia and Avicennia, stretching the circumscription of that family beyond their former delimitation and leaving this enlarged Acanthaceae without clear synapomorphies. Therefore, many families accepted below are difficult to define morphologically. The descriptions we provide are general and, although descriptive, provide little help in diagnosing the families. Accepting larger entities has been proposed and accepted here in the case of Gesneriaceae (and where we diverge from APG IV), but we otherwise maintain the status quo of many smaller families pending further studies. Lamiales have been estimated to have evolved c. 97 million years ago, and the order represents 12.3% of the diversity of eudicots. They are predominantly herbaceous plants with opposite leaves and five-lobed zygomorphic, often two-lipped flowers with the stamens fused to the corolla tube. Chemically, all members of the order share the presence of verbascosides, secondary compounds that have antimicrobial and anti-inflammatory properties in humans.

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543

LAMIALES

EUDICOTS

Plocosperma buxifolium, Nicaragua (OO) [400]

Carlemannia tetragona, Yunnan, China (BY) [401]

Silvianthus tonkinensis, Yunnan, China (YN) [401]

400. PLOCOSPERMATACEAE

with parietal placentation. A terminal elongate style forks dichotomously two times at the tip into four slender branches with apical stigmas. Fruits are bivalved capsules with one to four seeds that each bear a apical tuft of hairs.

401. CARLEMANNIACEAE

Staghorn-shrub family

Fragrant-princess family

Distribution: This family is restricted to Mesoamerica (southern Mexico, Guatemala, Nicaragua and Costa Rica), where it grows in dry forests.

These are terrestrial unisexual shrubs and small trees. They have simple, opposite (nearly opposite or whorled) leaves that are shortly petiolate and lack stipules. Blades are ovate with an entire to minutely toothed, often slightly recurved margin, an often emarginate tip and pinnate-arcuate venation with several rows of areoles parallel to the midrib. Inflorescences are axillary racemes composed of one, two or up to seven flowers subtended by a pair of leaves. Functionally unisexual flowers are slightly zygomorphic. The five (or six) sepals are basally fused, and this tube is much shorter than the lobes. The five (or six) petals are fused to form a large, funnel- or bell-shaped corolla with five lobes. Stamens are as many as and free from the petals, alternating with the corolla lobes. Anthers are basifixed and open by lengthwise slits. Stamens are staminodial in female flowers with a nectar disk surrounding the ovary. Male flowers have a pistillode. The superior ovary is stalked and composed of two (rarely one) fused carpels that form a single locule 544

Christenhusz, Fay & Chase

Phylogeny and evolution: Plocosperma was difficult to place when it was first discovered. Because of its branched style it was first placed near Gelsemium (then Loganiaceae, Gentianales), but the tuft of hairs on the seeds made some botanists suggest a relationship to Apocynaceae. As Lithophytum (a synonym of Plocosperma), it has also been placed among Solanaceae. Molecular studies have shown that the genus has no affinity with Gentianales or Solanales, but it was instead sister to Lamiales, an order in which it is now included. It shares with the other members of the order chemical compounds such as cornoside, which are otherwise only found in Gesneriaceae and Tetrachondraceae. Genera and species: Plocosperma buxifolium is the only species in this family. Etymology: Plocosperma is composed of the Greek words πλόκος ( plokos), a lock of hair, and σπέρμα (sperma), seed, an obvious reference to this characteristic of the only species.

Carlemanniaceae include perennial herbs and shrubs with simple, opposite, petiolate, more or less asymmetrical leaves without stipules, but with a tissue connecting the petioles across the node. Blades are pinnately veined with dentate or crenate-serrate margins. Inflorescences are terminal or axillary cymes or corymbs. Flowers are bisexual and slightly zygomorphic. The four or five sepals are fused into a tube with more or less unequal lobes that is adnate to the ovary. The four or five petals are fused into a tubular, funnelor bell-shaped slightly unequal corolla. The two stamens with short filaments are inserted in the middle of the corolla tube; they have dorsifixed, linear anthers that open by lengthwise slits and are connivent around the style. A conical or cylindrical disk surrounds the style on the top of the ovary. The inferior ovary is composed of two carpels, each forming a locule with axile to basal placentation. Fruits are biloculate, dry or fleshy dehiscent capsules with a persistent calyx and many (30–100) seeds.

LAMIALES

EUDICOTS

Distribution: This family is tropical and occurs from the eastern Himalayas, to northern Burma, Yunnan, northern Thailand, Laos and Vietnam, and northern Sumatra. Phylogeny and evolution: Carlemannia was originally placed in Rubiaceae and Silvianthus in Caprifoliaceae, but molecular studies have shown that these plants are closely related to each other and to other members of Lamiales. They share the two anthers with Oleaceae and appear superficially similar, but they differ from Oleaceae in their inferior ovary and the tissue between the leaves. The family is poorly known.

402. OLEACEAE Olive family

Etymology: Carlemannia is named for British botanist Charles Morgan Lemann (1806– 1852), who collected plants in Gibraltar and Madeira. George Bentham found a specimen of this genus from India in Lemann’s herbarium and named it for him.

Oleaceae are woody vines, shrubs and trees with opposite leaves (rarely alternate in some Jasminum). They are petiolate and lack stipules or tissue across the node. Leaf blades are simple, trifoliolate or unevenly pinnate, the venation is pinnate (rarely palmate) and leaf margins are entire or toothed. Inflorescences are terminal or axillary thyrses, panicles, racemes or cymes, but sometimes the flowers are solitary. Flowers are usually bisexual (rarely unisexual) and actinomorphic. The usually four sepals (rarely absent in some Fraxinus or up to 15 in some Jasminum) are basally fused. The four petals (rarely absent in some Fraxinus or up to 12 in some

Fraxinus velutina var. tourneyi, Royal Botanic Gardens, Kew, UK [402]

Abeliophyllum distichum, Royal Botanic Gardens, Kew, UK [402]

Genera and species: There are two genera and five species in this family: Carlemannia (3) and Silvianthus (2).

Jasminum) are fused into a tubular corolla (rarely free as in Fontanesia). Two stamens are fused to the corolla and alternate with the petal lobes (when four). Anthers are basifixed or dorsifixed and often bear osmophores, opening by lengthwise slits. A nectary is often present as a ring around the ovary. A superior ovary is composed of two carpels, fused to form two locules with a median septum and axile, apical or basal placentation. The ovary is topped by a single, simple style (sometimes absent or short) with a bilobed, club-shaped, dry stigma. Fruits include loculicidal capsules, samaras, schizocarps, berries and drupes. Distribution: This family can be found nearly worldwide with equal diversity in temperate and tropical areas. They are absent from northern regions in North America and Asia and the great deserts of Africa, Central Asia and Australia. Phylogeny and evolution: Fossil evidence of Fraxinus-like winged seeds from Eocene deposits have been dated to 44 million years old. The diversification of Oleaceae occurred during the Tertiary. Some generic realignment

Forsythia ovata, Helsinki Botanical Garden, Finland [402]

Fontanesia fortunei, Royal Botanic Gardens, Kew, UK [402]

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LAMIALES

Uses: Economically, the most important species is the olive (Olea europaea); the species forms a complex of subspecies along the Mediterranean, western Asia and Africa. Olives have been cultivated since ancient times for their valuable oily fruits. This was first documented in the eastern Mediterranean, with commercial cultivation by the Minoans on Crete around 3000 BC, giving the people there great wealth and allowing them to develop their civilization. Wild olives had already been collected during Neolithic times in Asia Minor and Greece. From there, better selections were rapidly distributed across

the region and became important for many local cultures. Olive oil was highly valued and was, for instance, imported from Assyria and Greece into Egypt and later also further afield. Currently, Spain produces nearly half of all olive oil, although it is cultivated in all countries with a mediterranean climate (California, Chile, South Africa, Australia, China etc.). Extra-virgin and virgin olive oil are pressed mechanically. Refined olive oil has undergone chemical treatment to remove the flavour and acid content of the natural oil, making it more homogeneous. Extra-virgin oil is of higher quality, has a fruitier flavour and is used mostly for salads, and virgin oil is used mostly for cooking, whereas refined oil is used in processed foods or for industrial applications. Fruits of olives are bitter and astringent when just picked, and to make them edible they first have to be cured and sometimes fermented. To cure them, they are soaked in lye or salt, and then they are either pickled, eaten fresh or fermented. Green olives have been picked unripe and have a yellowish colour, and ripe olives are reddish brown, purplish or black; however, bright green or dark black olives from the supermarket may have been dyed. An olive grove is valuable because trees yield fruit for thousands of years and pass down the generations. Unfortunately,

old olive trees transplant reasonably well, and ancient plants have been taken from groves to be used in landscaping elsewhere in Europe, where they do not always survive the climate. Stretchberry (Forestiera pubescens) is sometimes also cultivated for its similarly edible fruit. Timber of ash (Fraxinus excelsior) is elastic and has been used for cartwheels, tool handles, tennis racquets, polo mallets, baseball bats, hockey sticks and billiard cues. Many mythical and medicinal properties have also been attributed to this tree in the past. Other species of Fraxinus are also planted occasionally for timber. Timber from ironwood, Nestegis apetala and N. sandwichensis, found in New Zealand and Hawaii, respectively, is hard and highly valued. Australian Notelaea also yields valuable hard wood. Leaves of Chinese ash (Fraxinus chinensis) were used to feed scale insects (e.g. Ceroplastes ceriferus, Ericerus pela) that excrete a waxy substance used to polish paper and jade and for coating pills and previously candles. Jasmine is grown for its scented flowers that can be used for perfume and to flavour tea. The most common species for this purpose are Jasminum grandiflorum,

Olea europaea, Gibraltar [402]

Olea europaea, Lebanon [402]

Ligustrum vulgare, Canbury Gardens, Kingston upon Thames, Surrey, UK [402]

will be necessary because several genera (Chionanthus, Olea, Osmanthus) were not found to be monophyletic. Genera and species: Oleaceae include 24 genera and 790 species: Abeliophyllum (1), Chionanthus (c. 60), Comoranthus (3), Dimetra (1), Fontanesia (1), Forestiera (c. 16), Forsythia (9), Fraxinus (c. 42), Haenianthus (2), Hesperelaea (1), Jasminum (c. 450), Ligustrum (c. 40), Myxopyrum (4), Nestegis (5), Noronhia (44), Notelaea (11), Nyctanthes (2), Olea (c. 33), Osmanthus (32), Phillyrea (2), Picconia (1), Priogymnanthus (2), Schrebera (8) and Syringa (c. 20).

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EUDICOTS

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LAMIALES

EUDICOTS

Jasminum officinale, Jardin des Plantes, Paris, France [402]

J. odoratissimum, J. officinale and J. sambac. The last is commonly used in tropical Asia for scented flower arrangements and in religious offerings, although locally in India Nyctanthes arbor-tristis (harsinger tree) takes this place, as it is a sacred tree there. The flowers also yield an orange dye that is sometimes used as an alternative to saffron. Members of the genera Abeliophyllum (white forsythia), Chionanthus (fringe tree), Forsythia (forsythia), Fraxinus (ash), Jasminum (jasmine), Ligustrum (privet), Nyctanthes (harsinger tree), Olea (olive), Osmanthus (sweet olive), Phillyrea (false olive) and Syringa (lilac) are frequently cultivated in subtropical and temperate gardens. Abeliophyllum distichum is threatened in its native habitat in central Korea, but it is a good example of how a plant can be saved from global extinction through horticulture. Species of privet (Ligustrum vulgare, L. lucidum) are commonly grown in gardens for hedging, due to their tolerance of pollution. Flowering branches of Forsythia and Syringa are sometimes used as cut flowers. Etymology: Olea is the Latin name for olive, derived from the Greek έλαιο (elaio), oil.

Osmanthus serrulatus, Royal Botanic Gardens, Kew, UK [402]

Holy olive: An olive branch has often been a symbol of peace, fertility, abundance and victory. Olive branches were often ritually offered to rulers and gods as a symbol of purification and glory. They were used to crown brides, virgins and victors of games and wars alike, and their symbolism in western culture is derived from ancient Greek tradition. Olives were widespread across the Levant and a major food crop for the population, so it is not surprising that the olive plays an important role in the Torah, Bible and Koran alike. One of the hills outside Jerusalem is called the Mount of Olives because of the groves that were found there in earlier times. It has been a Jewish cemetery for three millennia, and it is the site from where Jesus Christ is said to have ascended to heaven, making it a place for Christian pilgrimage since early Christian times. Famous of course is the olive branch that a dove (a symbol of the holy spirit) brought back to Noah to show that the water had retreated enough so that is was safe to leave the Ark. A white dove with an olive branch is now a universal symbol of peace. In modern culture, the bald eagle on the Great Seal of the USA holds a bundle of arrows in one claw and an olive branch in the other, denoting the power of peace and war vested in the Congress.

Syringa reflexa, Ruissalo Botanical Garden, Turku, Finland [402]

403. TETRACHONDRACEAE Rustweed family

This is a family of perennial, aquatic and partially aquatic herbs with creeping succulent stems. Their sessile leaves are opposite and fused across the node with an interstipular sheath. Blades are linear to ovate and succulent, aromatic and glanddotted, with a minutely toothed margin and an obscure, principally pinnate venation. Inflorescences are axillary, leafy cymes, or flowers are solitary. Flowers are bisexual and actinomorphic. The four sepals are basally fused and persistent. The four petals are fused into a short tube; in Polypremum, with a ring of hairs in the throat of the tube. The four stamens alternate with the petals, the filaments are fused to the corolla tube and anthers are dorsifixed and open inwardly by lengthwise Plants of the World

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Polypremum procumbens, North Palm Beach, Florida, USA (BP) [403]

Peltanthera floribunda in fruit, Puntarenas, Costa Rica (CD) [404]

Aeschynanthus sikkimensis, Royal Botanic Gardens, Kew, UK [404]

slits. The half-inferior (Polypremum) or superior (Tetrachondra) ovary is composed of two carpels with basal placentation. The ovary apex is slightly sunk or highly constricted with an apical (Polypremum) or gynobasic (Tetrachondra) style and a pointed stigma. Fruits are bivalved capsules or schizocarps with four nutlets.

identical, and thus it is assumed that dispersal to New Zealand from South America is probably recent. Polypremum diverged from Tetrachondra c. 46 million years ago.

distichous or spirally arranged. Blades are petiolate or not, lack stipules, the margins fused across the node or not, sometimes the leaf pair fused across the stem at their base, with pinnate or palmate venation and entire, crenate, serrate or dentate margins. Leaves and stems are typically somewhat succulent or f leshy and covered with dense hairs, sometimes glandular, some without hairs and herbaceous leaves, or in various combinations of these. Inflorescences are leafy or bracteate, indeterminate thyrses with axillary paired cymes that are sometimes reduced to single flowers. Flowers are bisexual and usually zygomorphic. The (four or) five sepals are free or fused, equal or unequal, sometimes bi- or trilabiate. The (four or) five petals are fused to an often bilabiate corolla that is variable in shape and size, tubular, curved, funnel-, bellsac- or cup-shaped or flat and open (rarely actinomorphic). Sac-shaped or pouched flowers can bear oil glands (Calceolaria). The two, four or five stamens are all fertile or some sterile (staminodes). Filaments are free from each other but fused to the corolla tube, often two long and two short (didynamous) when there are four stamens. Anthers are variable in shape, basifixed or dorsifixed, opening by lengthwise slits or (basal or) apical pores, often cohering at their tips or for their entire length. A nectar disk is fused to the base of the ovary or free from it, forming a ring, half-moon-shaped or composed of separate free nectar glands,

Distribution: Polypremum is found from southeastern North America (USA and Mexico), Mesoamerica and the Caribbean to northern South America. It has been accidentally introduced to several Pacific islands (Guam, Hawaii, Palau and Wallis), and there is a specimen from Paraguay. Tetrachondra occurs disjunctly in southern South America (Patagonia, Tierra del Fuego) and New Zealand (Stewart Island, Otago and Southland). They generally grow in permanently to periodically wet habitats. Phylogeny and evolution: These diminutive plants have reduced flowers and have thus been difficult to place taxonomically in the past. Tetrachondra was originally placed in Crassulaceae, whereas Polypremum was considered to be a member of Loganiaceae or Rubiaceae. Their four-lobed ovaries with a gynobasic style have suggested a relationship with Lamiaceae or Boraginaceae, but we now know that this character has evolved independently several times. DNA sequences of the two species of Tetrachondra are nearly 548

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Genera and species: This is a small family with two genera and three species: Polypremum procumbens, Tetrachondra hamiltonii and T. patagonica. Etymology: Tetrachondra is composed of the Greek words τετρα (tetra), four, and χόνδρος (chondros), cartilage.

404. GESNERIACEAE Gloxinia family

This family includes perennial (rarely annual) herbs, shrubs, treelets and climbers. They are epiphytic, epilithic or terrestrial and often have underground stems, tubers, rootstocks, scaly rhizomes or simple-branching aboveground stems. Leaves are simple (rarely lobed or pinnately dissected), opposite or alternate, sometimes whorled, nearly

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sometimes absent. The superior, half-inferior or inferior ovary is composed of two carpels forming a single locule, topped by a usually well-developed style and a capitate, bilobed or variously split stigma. Ovule placentation is usually parietal (semi-parietal in Sanango). Fruits are dehiscent or indehiscent, fleshy or dry capsules, opening septicidally by two or loculidically by four valves, sometimes nutlets or berries.

Distribution: Gesneriaceae have a pantropical distribution, but they are especially diverse in tropical mountainous areas. They extend into temperate East Asia, southern Chile, New Zealand and have two isolated genera in the Pyrenees and the Balkan mountains of Europe. Phylogeny and evolution: Gesneriaceae have often been divided into two families, the predominantly Neotropical Gesneriaceae and

the mostly Palaeotropical Didymocarpaceae, which were united into a single family in the late 19th century. The two families alternatively were retained at subfamilial level and were mainly separated on the position of the ovary. Molecular studies have now shown that several genera previously placed in Scrophulariaceae (a widely polyphyletic family in the traditional sense) were found close to Gesneriaceae, and thus the family could be expanded to include

Calceolaria pinifolia, Royal Botanic Gardens, Kew, UK [404]

Calceolaria uniflora, Royal Botanic Gardens, Kew, UK [404]

Primulina tamiana, Royal Botanic Gardens, Kew, UK [404]

Streptocarpus prolixus, Helsinki Botanical Garden, Finland [404]

Jovellana violacea, Glasgow Botanic Gardens, UK [404]

Haberlea ferdinandi-coburgii, Royal Botanic Gardens, Kew, UK [404]

Streptocarpus teitensis, Mbololo Hill, Taita Hills, Kenya [404] Petrocosmea minor, Royal Botanic Gardens, Kew, UK [404]

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Rhytidophyllum exsertum, Royal Botanic Gardens, Kew, UK [404]

EUDICOTS

Achimenes erecta, National Botanic Gardens Alloplectus cristatus, Guadeloupe [404] of Ireland, Glasnevin [404]

Niphaea oblonga, Royal Botanic Gardens, Kew, UK [404]

Koellikeria erinoides, Royal Botanic Gardens, Kew, UK [404]

Calceolaria, Jovellana, Peltanthera and Sanango. Only the last, found to be sister to both traditional subfamilies, was included by APG IV, but there has been a tendency to accept Calceolariaceae as a separate family, which unfortunately was followed by APG IV, leaving Peltanthera as an unplaced genus. We opt for a reduction of families here because there are few distinguishing characters and the families are better treated at subfamilial levels. Calceolariaceae for instance have tetramerous f lowers, but this character is also found in some South American Gesnerioideae. Sanango was originally included in Loganiaceae, but it has now been 550

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Sinningia longiflora, private collection, Kingston upon Thames, Surrey, UK [404]

found to be sister to Gesneriaceae (of APG III) in molecular studies. Its ovaries have parietal placentation in the upper part of the ovary (as in Gesnerioideae and Didymocarpoideae), but have axile placentation in the lower part of the ovary (as in Calceolarioideae and Peltantheroideae), which shows that this character is not suitable for distinguishing between families. These families are better united, as treated here. This is the only case among the angiosperms in which this book differs from APG IV. Crown Gesnerioideae may have diversified c. 43–45 million years ago. The clade including Coronanthera is disjunct

Gesneria pedicellaris × G. pedunculosa, Royal Botanic Gardens, Edinburgh, UK [404]

Sarmienta repens, Royal Botanic Gardens, Kew, UK [404]

between southern Chile, Australasia and northeastern Asia, and was therefore once suggested to be of Gondwanan origin, but they evolved a mere 9.5 million years ago and thus must have achieved their current distribution through long-distance dispersal. Generic delimitation was poor before the advent of molecular studies, but as relationships have been clarified, many genera have been recircumscribed in recent times. For instance, Gloxinia is now placed in a separate tribe from Sinningia; they are not closely related. The peloric forms of Sinningia speciosa were formerly placed in Gloxinia, hence the confusion of the common name.

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Genera and species: Gesneriaceae include 175 genera and c. 3,810 species in five subfamilies: ‘Peltantheroideae’ – Peltanthera (1); Calceolarioideae – Calceolaria (c. 265) and Jovellana (6); Sanangoöideae – Sanango racemosum; Didymocarpoideae – Acanthonema (2), Aeschynanthus (c. 160), Agalmyla (>90), Allocheilos (2), Allostigma (1), Anna (3), Beccarinda (c. 8), Billolivia (5), Boea (10), Boeica (7), Briggsiopsis (1), Calcareoboea (1), Cathayanthe (1), Championia (1), Chayamaritia (2), Codonoboea (81), Conandron (1), Corallodiscus (5), Cyrtandra (c. 600), Cyrtandromoea (15), Damrongia (10), Deinostigma (1), Didissandra (8), Didymocarpus (c. 70), Didymostigma (2), Dolicholoma (1), Dorcoceras (4), Emarhendia (1), Epithema (>20), Glabrella (2), Gyrocheilos (4), Gyrogyne (1), Haberlea (2), Hemiboea (25), Henckelia (c. 230), Hexatheca (4), Jerdonia (1), Kaisupeea (3), Lagorosolen (2), Leptoboea (3), Liebigia (14), Loxonia (3), Loxostigma (>7), Lysionotus (c. 30), Metapetrocosmea (1), Microchirita (25), Middletonia (4), Monophyllaea (>30), Nodonema (1), Opithandra (10), Orchadocarpa (1), Oreocharis (100), Ornithoboea (16), Paraboea (c. 90), Petrocodon (1), Petrocosmea (27), Phylloboea (1), Platystemma (1), Primulina (c. 160), Pseudochirita (1), Ramonda (4), Raphiocarpus (11), Rhabdothamnopsis (1), Rhynchoglossum (c. 10), Rhynchotechum (c. 14), Ridleyandra (>20), Senyumia (1), Sepikea (1), Somrania (3), Spelaeanthus (1), Stauranthera (5), Streptocarpus (c. 176), Tengia (1), Tetraphyllum (6), Trachystigma Bacopa monnieri, Helsinki Botanical Garden, Finland [405]

(1), Tribounia (2), Trisepalum (13) and Whytockia (8); Gesnerioideae – Achimenes (c. 24), Alloplectus (>75), Alsobia (2), Anetanthus (2), Anodiscus (1), Asteranthera (1), Bellonia (2), Besleria (>200), Capanea (2 or 11), Christopheria (1), Chrysothemis (7), Cobananthus (1), Codonanthe (>20), Codonanthopsis (5), Columnea (>270), Coronanthera (11), Corytoplectus (15), Cremersia (1), Cremosperma (c. 27), Cremospermopsis (2), Depanthus (2), Diastema (>20), Drymonia (>140), Episcia (9), Eucodonia (2), Fieldia (1), Gasteranthus (35), Gesneria (>60), Gloxinia (15), Goyazia (2), Heppiella (5), Hygea (1), Koellikeria (1), Kohleria (17), Lampadaria (1), Lembocarpus (1), Lenbrassia (1), Lesia (1), Mitraria (1), Monopyle (>20), Moussonia (11), Napeanthus (>20), Nautilocalyx (>70), Negria (1), Nematanthus (30), Neomortonia (3), Niphaea (2–5), Oerstedina (3), Pachycaulos (1), Pagothyra (1), Paliavana (6), Paradrymonia (>70), Pearcea (17), Pheidonocarpa (1), Phinaea (>10), Reldia (5), Resia (2), Rhabdothamnus (1), Rhoogeton (4), Rhytidiphyllum (c. 20), Rufodorsia (4), Sarmienta (1), Shuaria (1), Sinningia (>60), Smithiantha (7), Solenophora (16), Titanotrichum (1), Tylopsacas (1) and Vanhouttea (8); Unplaced – Coptocheile (1).

ionanthus) and Cape primroses (Streptocarpus ×hybridus, a complex hybrid derived from S. rexii). Other genera often grown for their beautiful flowers include Achimenes, Aeschynanthus, Columnea, Episcia, Haberlea, Jovellana, Kohleria, Nematanthus, Petrocosmea, Primulina, Ramonda and Smithiantha. Etymology: Gesneria commemorates Swiss naturalist and philosopher Konrad Geßner (Conradus Gesnerius, 1516–1565), who is known as the father of modern zoology, and among many botanical works he was the first to describe a cultivated tulip in western Europe. He also maintained an interesting garden. For a while, he was professor of Greek at the academy of Lausanne, after which he received his doctorate and became a physician in Zurich. He attempted to assemble a universal library, containing all books published at the time, from which he created a subject-based catalogue, one of the first databases.

405. PLANTAGINACEAE Speedwell family

Uses: Several species are popular houseplant and garden ornamentals. Best known are slipperworts (Calceolaria ×herbeohybrida, complex hybrids involving C. cana, C. corymbosa and C. crenatiflora), florist’s gloxinias (peloric forms of Sinningia speciosa), African violets or saintpaulias (Streptocarpus Linaria aeruginea, Ronda, Spain [405]

Callitriche stagnalis, Stott Park, Lake District, UK [405]

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Intuitive taxonomy - Charles Edwin Bessey (1845–1915) Bessey’s goal was to arrange flowering plants in a scheme that ref lected evolutionary relationships. His being one of the first of such schemes, Bessey’s scheme had an enormous influence on later

Photograph of Charles Edwin Bessey from The Cornhusker, the Book of the Upper Classes of the University of Nebraska, vol. 1. 1907. Lincoln, University of Nebraska

classifications by, for instance, Cronquist, Takhtajan and Thorne. He had an intuitive approach, which worked surprisingly well in some cases, but in others failed to discern more obvious relationships (e.g. he placed

Photograph of Veronica wyomingensis (Besseya wyomingensis) in the rock garden of the Royal Botanic Gardens, Kew, UK

This variable family includes annual, biennial and perennials herbs, shrubs, (herbaceous) vines and small trees. They can be terrestrial or submerged aquatics, rarely epiphytes, carnivores or resurrection plants. Leaves are simple or compound, opposite or alternate and spirally arranged, or whorled, without stipules. Leaf blades have pinnate or palmate, rarely parallel venation, sometimes reduced to a single vein, and margins are serrate, crenate or entire, the blades sometimes bearing extrafloral nectaries. Inflorescences are terminal or axillary cymes or racemes, spikes or heads, or flowers solitary in leaf axils, often bracteate, the bracts scale- or leaflike, sometimes forming an involucre. Flowers are bisexual and zygomorphic or secondarily actinomorphic in Sibthorpia, Aragoa and Plantago. The (two to) four or five (to eight) sepals are usually fused (free in Trapella, absent in Callitriche). The (two to) four or five (to eight) petals (absent in Callitriche, Hippuris and some Veronica) are fused into a cup-, bell-, funnel-shaped or two-lipped 552

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Cactales with Myrtales rather than Caryophyllales). Intuition is still commonly used by taxonomists (especially those in favour of recognising paraphyletic groups, a process that relies on intuition to determine when a group is modified enough to warrant taxonomic recognition), sometimes more successfully than others, and it has been opposed to the application of a more objective system of classification. The genus Besseya was named in his honour, but molecular phylogenetics has shown that it is part of Veronica, in which genus it now resides. In spite of the limitations of his approach, Bessey’s scheme was a great step forward in the understanding of plant evolutionary relationships.

corolla (usually with two upper and three lower lobes), sometimes with a nectar spur or pouch. Stamens are usually two longer and two shorter (didynamous), but a great variety in stamen numbers exists, from one (Callitriche, Hippuris), two (e.g. Veronica, Gratiola, Trapella), rarely five to eight. Filaments are usually fused to the corolla tube, alternating with the petals, rarely free. Anthers are dorsifixed and open by lateral slits or (in Campylanthus, Globularia and Poskea) by short apical slits on fused thecae. The connective in Trapella is large and peltate. Staminodia are one, two or absent. A nectar disk is circular and surrounds the ovary or absent. The usually superior ovary (inferior in Hippuris and Trapella) is composed of two connate carpels (rarely three, in Hippuris one), each forming a locule, or in Callitriche tetralocular via secondary septa. The ovary is topped by two, partially free or entirely fused styles. The stigma is capitate, bilobed or two-lipped. In Callitriche, the two styles are completely free from each other

Chart to show relationships of the orders, with the relationships indicated by position and the area being approximately proportional to the numbes of species in the orders (From C. E. Bessey, 1915, The phylogenetic taxonomy of flowering plants. Annals of the Missouri Botanical Garden 2: 109–164).

and have stigmatic areas along their entire length. Ovule placentation is usually axile, sometimes apical, rarely intrusively parietal, basal or apical. Fruits are capsules that open septicidally, loculicidally, poricidally or circumscissily, rarely a berry, nut, schizocarp or drupe. Distribution: Plantaginaceae can be found nearly worldwide, extending into arid regions, the Arctic and Antarctic, temperate and tropical forests, grasslands, mountains etc. Phylogeny and evolution: It is not easy to distinguish between Plantaginaceae and other families in Lamiales. Septicidal capsule dehiscence is unusual among Lamiales, although this is by no means present in all genera of Plantaginaceae (most notably this is not found in the type genus Plantago) and also not unique to them. This lack of morphological characters makes a case for uniting the bulk of Lamiales (including Acanthaceae, Lamiaceae, Verbenaceae, Bignoniaceae, Scrophulariaceae,

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EUDICOTS

Cymbalaria muralis, private garden, Kingston upon Thames, Surrey, UK [405]

Angelonia parviflora, Helsinki Botanical Garden, Finland [405]

Digitalis purpurea, Bix Bottom, UK [405]

Globularia trichosantha, Royal Botanic Gardens, Kew, UK [405]

Veronica chamaedrys, Box Hill, UK [405]

Plantaginaceae, Gesneriaceae etc.) into a single family, although this finds lots of resistance in the user community and is not in line with the traditional, historical taxonomy of the group. Phylogenetic studies initially found only weak support for Plantaginaceae (often called Veronicaceae in early works), but soon stronger support was found for an alliance of Plantaginaceae, including Callitrichaceae, Hippuridaceae, Globulariaceae, Trapellaceae and Scrophulariaceae (e.g. Antirrhinoideae, Chelonoideae, Digitalidoideae, Gratioloideae, Hemiphragmateae, Sibthorpioideae, Veronicoideae). Plantaginaceae is the oldest name and thus has nomenclatural priority. Several

Veronica pinguifolia, Royal Botanic Gardens, Kew, UK [405]

Veronicastrum axillare, Royal Botanic Gardens, Kew, UK [405]

genera are poorly defined or based on minute differences that may not hold up in future revisionary or phylogenetic studies. Veronica has already been greatly expanded. The Bacopa alliance also appears poorly circumscribed. Gratioloideae are found as sister to the rest of the family, sometimes in an unresolved polytomy. Evidence is too scanty to make a decision on providing a subfamilial classification of Plantaginaceae, and Gratioloideae are therefore not maintained here. Fossil pollen possibly originating from Plantago has been found in Miocene and younger layers in Europe.

Genera and species: Plantaginaceae encompass 99 genera and c. 1,900 species: Acanthorrhinum (1), Achetaria (7), Adenosma (15), Albraunia (3), Anamaria (1), Anarrhinum (8), Angelonia (c. 26), Antirrhinum (c. 20), Aragoa (c. 20), Asarina (1), Bacopa (c. 55), Basistemon (8), Benjaminia (1), Boelckea (1), Braunblanquetia (1), Brookea (4), Callitriche (17), Campylanthus (15), Chaenorhinum (21), Cheilophyllum (8), Chelone (4), Chionophila (2), Collinsia (c. 20), Conobea (7), Cymbalaria (9), Darcya (3), Deinostema (2), Digitalis (c. 25), Dintera (1), Dizygostemon (2), Dopatrium (14), Encopella (1), Ellisiophyllum (1), Epixiphium (1), Erinus (1), Gadoria (1), Galvezia (4), Plants of the World

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LAMIALES Gambelia (4), Globularia (23), Gratiola (c. 25), Hemiphragma (1), Hippuris (2), Holmgrenanthe (1), Holzneria (2), Howelliella (1), Hydrotriche (4), Kashmiria (1), Keckiella (7), Kickxia (9), Lafuentea (1), Lagotis (c. 20), Leucospora (1), Limnophila (37), Linaria (c. 150), Lophospermum (6), Mabrya (5), Maeviella (1), Maurandella (1), Maurandya (2), Mecardonia (c. 10), Melosperma (1), Misopates (7), Mohavea (2), Monopera (2), Monttea (3), Nanorrhinum (10), Neogaerrhinum (2), Neopicrorhiza

(1), Nothochelone (1), Nuttallanthus (4), Otacanthus (7), Ourisia (28), Penstemon (c. 275), Philcoxia (3), Picrorhiza (4), Plantago (c. 270), Poskea (3), Pseudorontium (1), Rhodochiton (3), Russelia (52), Sairocarpus (13), Schistophragma (2), Schizosepala (1), Schweinfurthia (6), Scoparia (c. 20), Scrofella (1), Sibthorpia (5), Sophronanthe (1), Stemodia (c. 60), Tetranema (5), Tetraulacium (1), Tonella (2), Trapella (2), Uroskinnera (c. 5), Veronica (c. 420), Veronicastrum (2), Wulfenia (5) and Wulfeniopsis (2).

Uses: American brooklime (Veronica americana) is sometimes eaten as a salad, and tastes similar to watercress. Medicinally, the most important derivative is digitalin, a cardiac glycoside used as a cardiac stimulant since the 18th century. It is mostly harvested from common foxglove (Digitalis purpurea) and Austrian foxglove (D. lanata). Psyllium (Plantago afra) is added to pasta and cereals to lower cholesterol. In stronger concentrations, it has a laxative effect and is a main component of over-the-counter treatments for constipation.

Sibthorpia peregrina, Helsinki Botanical Garden, Finland [405]

Ourisia coccinea, Royal Botanic Gardens, Kew, UK [405]

Penstemon campanulata, private garden, Kingston upon Thames, Surrey, UK [405]

Plantago afra, Sicily, Italy [405]

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Rhodochiton atrosanguineus, private garden, Kingston upon Thames, Surrey, UK [405]

Hippuris vulgaris, National Botanic Gardens of Ireland, Glasnevin [405]

Tetranema rosea, New York Botanical Garden, USA [405]

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EUDICOTS

Many species are grown as ornamental plants, especially Angelonia (summer snapdragon), Antirrhinum (snapdragon), Asarina procumbens, Bacopa monnieri (water hyssop), Chelone obliqua (turtlehead), Cymbalaria muralis (ivy-leaved toadflax), Digitalis (foxglove), Ellisiophyllum pinnatum, Erinus alpinus (fairy foxglove), Globularia (globe daisy), Gratiola (hedge hyssop), Hemiphragma heterophyllum, Hippuris vulgaris (mare’s tail), Linaria (toadflax), Lophospermum (lophos, asarina), Ourisia coccinea, Penstemon (beard tongue), Rhodochiton atrosanguineus (purple bellvine), Russelia equisetiformis (firecracker plant), Tetranema rosea (Mexican violet), Veronica (speedwell), Veronicastrum (Culver’s root) and Wulfenia, to name a few. Bacopa of horticulture is botanically Sutera cordata (Scrophulariaceae) and is not related to Bacopa in spite of morphological similarity. Carnivory: Philcoxia comprises three rare Brazilian terrestrial species with subterranean stems and peltate leaves with circinate vernation, unusual in Plantaginaceae. The lower side of their leaves is provided with stalked capitate glands that produce phosphatases. They catch and digest nematodes, and nutrient uptake has also been detected. The inclusion of this genus in Plantaginaceae shows that carnivory in combination with circinate leaf vernation evolved several times in Lamiales, as it is also found in Byblidaceae and Lentibulariaceae, and thus these families may not be so different from other Lamiales as previously assumed. Philcoxia was only discovered in 1992 in Minas Gerais (Brazil), but the exact locality was not given. The population was only relocated in 2007. Later, two other species were recognised from Bahia and Goiás (also Brazil), the latter from a herbarium specimen preserved at the New York Botanical Garden. It had been misfiled under Lentibulariaceae and was thus not recognised as something new for decades. Etymology: Plantago is composed of Latin planta, the sole of the foot, and the suffix -ago, like. The plant grows where people have trodden the soil.

406. SCROPHULARIACEAE Figwort family

These are bisexual (rarely functionally unisexual), annual, biennial and perennial herbs, evergreen and deciduous shrubs, rarely trees and vines. Leaves are alternate, opposite or verticillate, simple or pinnately lobed, without stipules or in Buddleja sometimes with interpetiolar stipule-like lobes or leaf-like stipules at the base. Petioles are sometimes fused in pairs at base, often winged. Leaf blades have pinnate or palmate venation, and margins are serrate, crenate or entire, sometimes lobed or the margins inrolled and the leaves ericoid. In species of Androya, Eremophila, Bontia and Myoporum, there are glandular cavities containing oil or resin, and the blade often has translucent or raised dots when secretory cavities are large. Inflorescences are terminal or axillary, compound cymes forming panicles, thyrses, racemes, spikes or heads, the flowers rarely solitary. Flowers are usually zygomorphic (sometimes actinomorphic). The (two to) five sepals are often unequal in size, usually basally fused. The (four or) five petals are fused into a funnel-, bell- or tube-shaped or two-lipped corolla, rarely with one or two spur- or pouch-like appendages. Stamens are usually two longer and two shorter (didynamous), two or four stamens of equal length (Buddleja) or five equal stamens (Capraria, most Verbascum). Filaments are usually fused to the corolla tube, and anthers are dorsifixed, often the thecae confluent apically, placed head-to-head, in a U or parallel to each other and opening by lengthwise or sometimes transverse slits. There are one, two or three staminodia or all stamens fertile. The superior ovary is composed of two carpels (rarely one),

each forming a locule, with axile, apical or central placentation and topped by a single simple style and a capitate (rarely bifid or tongue-shaped) stigma. Male flowers have a pistillode. Fruit is usually a septicidal, septifragal or apically opening capsule, rarely a berry, drupe, nutlet or schizocarp. Distribution: Scrophulariaceae have a nearly global distribution but are absent from the far Arctic and Antarctic and sparse or absent in tropical Africa and the Atacama, Sahara, Arabian, Turkmenian and Gobi Deserts. They grow mainly in subtropical and warmtemperate regions, with their greatest diversity in South Africa and Australia. Phylogeny and evolution: Large-scale investigation of phylogenetic relationships using DNA sequence data have radically altered the circumscription of many families in Lamiales, especially Scrophulariaceae. These studies have indicated that a number of genera traditionally placed here should be excluded, but that the family should now include the former families Buddlejaceae, Selaginaceae and Myoporaceae, previously not even necessarily associated with Lamiales. This family used to include a number of genera that are now placed in Calceolariaceae, Linderniaceae, Phrymaceae, Plantaginaceae and Stilbaceae, placed there on the basis of their phylogenetic relationships. An age of 69 million years has been estimated for Scrophulariaceae. Genera and species: Scrophulariaceae include 59 genera and c. 1,830 species: Alonsoa (11), Androya (1), Antherothamnus (1), Anticharis (14), Aptosimum (20), Barthlottia (1), Bontia (1), Buddleja (c. 90), Camptoloma (3), Capraria (4), Chaenostoma (46), Chenopodiopsis (3), Colpias (1), Cromidon (12), Dermatobotrys (1), Diascia (c. 50), Diclis (10), Dischisma (11), Emorya (2), Eremogeton (1), Eremophila (c. 215), Freylinia (9), Glekia (1), Globulariopsis (7), Glumicalyx (6), Gomphostigma (2), Gosela (1), Hebenstretia (c. 40), Hemimeris (6), Jamesbrittenia (c. 85), Leucophyllum (12), Limosella (15), Lyperia (6), Manulea (74), Manuleopsis (1),

Plants of the World

555

LAMIALES

Alonsoa meridionalis, Ecuador [406]

EUDICOTS

Buddleja globosa, Royal Botanic Gardens, Kew, UK [406]

Diascia integerrima, Beth Chatto Garden, UK [406]

Jamesbrittenia macrantha, Lydenburg, South Africa (CD) [406]

Eremophila oldfieldii, Western Australia [406]

Scrophularia sambucifolia, Ronda, Spain [406]

Verbascum dumulosum, Royal Botanic Gardens, Kew, UK [406]

Myoporum laetum, Sea Ranch, California [406]

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Selago thunbergii, Adelaide Botanic Garden, South Australia [406]

Leucophyllum laevigatum, The Living Desert, Palm Desert, California, USA [406]

Zaluzianskya capensis, Ruissalo Botanical Garden, Turku, Finland [406]

LAMIALES

EUDICOTS

Melanospermum (6), Microdon (3), Myoporum (28), Nathaliella (1), Nemesia (c. 65), Oftia (3), Oreosolen (4), Peliostomum (7), Phygelius (2), Phyllopodium (26), Polycarena (17), Pseudoselago (28), Ranopisoa (1), Reyemia (2), Rhabdotosperma (7), Scrophularia (c. 200), Selago (c. 190), Strobilopsis (1), Sutera (c. 50), Teedia (4), Tetraselago (4), Trieenea (9), Verbascum (c. 360) and Zaluzianskya (57). Uses: Hawaiian Myoporum sandwicense and African Buddleja salviifolia produce good timber. A large number of genera are used as garden ornamentals and bedding plants, particularly Alonsoa warscewiczii (mask flower), Buddleja (butterfly bush), Diascia (twinspur), Eremophila (emu bush), Freylinia (honeybells), Leucophyllum (Texas ranger), Myoporum (mousehole tree), Nemesia (nemesias), Phygelius (Cape figwort), Scrophularia (figwort), Sutera (garden bacopa), Verbascum (mullein) and Zaluzianskya (night phlox). The butterfly bush (Buddleja davidii) is often planted because it more attractive to butterflies than native plants, but in suitable climates in Europe and North America, Australia and New Zealand, the plant can become invasive, germinating in walls and masonry and along railway tracks and roads. As a common weed in gardens, it can form large stands that crowd out native plants. Myoporum, Verbascum and Phygelius can also become problematic invasives.

Etymology: Scrophularia is derived from Latin scrofulae, swellings of the neck glands, referring to its medicinal properties.

407. LINDERNIACEAE Wishbone-flower family

These annual and perennial herbs, sometimes shrubs, often have square stems in crosssection. Chamaegigas and Craterostigma are resurrection plants in vernal pools, and other genera also often grow in wet habitats. Leaves are opposite, usually simple, in Scolophyllum sometimes pinnately compound, without stipules, but sometimes fused pairwise at the base with petioles often winged. Leaf blades are serrate, crenate, lobed or entire, and venation is pinnate or palmate. Inflorescences are terminal or axillary racemes or heads, or flowers are solitary in the leaf axils. Flowers are bisexual and zygomorphic, usually upright (resupinate in Lindernia hypandra). The (four or) five sepals are fused at least basally, sometimes unequal in size and often winged. The four or five petals are fused into

Craterostigma wilmsii, Buffelskloof Private Nature Reserve, South Africa (CD) [407]

a two-lipped to funnel-shaped corolla with glandular hairs on the adaxial side and in Lindernia with lobes covering the anthers. Stamens are four or five, usually four or two fertile and the others sterile. Filaments are free from each other and fused to the corolla tube. Anthers are dorsifixed, side-by-side, headto-head or (in Torenia) fused and opening by longitudinal slits. Staminodia (sterile stamens) are jointed and Z-shaped or long and curved with clavate or spur-like appendages and blue and yellow glandular hairs, sometimes strongly reduced or absent. The superior ovary is composed of two fused carpels, each forming a locule with axile to basal placentation. Styles are single and simple, topped with a bilobed, papillate, wet stigma. Fruits are usually septicidal and septifragal, sometimes poricidal capsules. Distribution: These plants are found throughout the tropics, with a few species extending their range into temperate North America, Europe and East Asia. Their greatest diversity is in tropical Africa and Southeast Asia. Phylogeny and evolution: The sister-group relationship of Linderniaceae is unresolved, but they may be closest to Byblidaceae. They are certainly not closely related to Scrophulariaceae, in which the genera of this family used to be placed. Cubitanthus, formerly in Gesneriaceae or Scrophulariaceae,

Torenia grandiflora, private collection, Kingston upon Thames, Surrey, UK [407]

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LAMIALES

Torenia crustacea, Guatemala (MV) [407]

is sister to Stemodiopsis; together this pair are sister to the rest of the family. It is possible that some unsampled genera, now tentatively placed in Plantaginaceae, also belong to this family. Linderniaceae also likely include Ameroglossum, Hemiarrhenia, Schizotorenia and Scolophyllum, and the positions of Bampsia, Bythophyton, Dintera and Encopella are not yet certain. Genera and species: Linderniaceae include 25 genera and 263 species: Ameroglossum (1), Artanema (4), Bampsia (2), Bonnaya (12), Bythophyton (1), Chamaegigas (1), Craterostigma (25), Crepidorhopalon (30), Cubitanthus (1), Dintera (1), Encopella (1), Hartliella (4), Hemiarrhena (1), Legazpia (1), Lindernia (30), Linderniella (16), Micranthemum (14), Microcarpaea (2), Picria (1), Pierranthus (1), Schizotorenia (2), Scolophyllum (3), Stemodiopsis (6), Torenia (51) and Vandellia (52). Uses: Vandellia micrantha is consumed in Laos, but most members of this family taste bitter. Some are used as ornamental plants, especially wishbone f lowers (Torenia fournieri and T. thouarsii), which are common bedding plants, and the resurrection plant, blue gem (Craterostigma plantagineum), is also sometimes grown. Baby tears (Micranthemum umbrosum) is sold as an aquarium plant. Torenia crustacea 558

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EUDICOTS

Torenia fournieri, Ruissalo Botanical Garden, Finland [407]

is widely naturalised throughout the tropics, especially between pieces of pavement in urban areas. Carnivory: Some species of Vandellia in Australia, e.g. V. cleistandra, are covered in glandular hairs with which they catch many insects. Whether these insects are actively absorbed is not yet known, but the close relationship between this family and carnivorous Byblidaceae, also from Australia, makes this an interesting topic for further research. Etymology: Lindernia is named for Alsatian botanist, author and physician Franz Balthazar von Lindern (1682–1755). He became well known for his work on sexually transmitted diseases (Speculum veneris) and his book on the plants of the Alsace.

408. BYBLIDACEAE Rainbowplant family

These are annual and perennial, shrub-like

herbs, often with a woody rhizome. Leaves are alternate and spirally arranged, linear to filiform with flat or circinate, curved, vernation without stipules or leaf sheaths. Veins are parallel, the lamina strongly reduced so only the midrib remains, the margins entire, with a knob-like swelling at the leaf tip. The blade is covered with mucilage-secreting insect-trapping stalked glandular hairs and sessile enzyme-secreting hairs. Inflorescences are axillary solitary flowers without bracts. Flowers are bisexual and slightly zygomorphic. The five sepals are basally fused and persistent in fruit. The five petals appear nearly free, but are basally fused and serrate to fimbriate at the margin. The five stamens alternate with the petals or are displaced and bent against one side of the flower. Filaments are short, twisted and often slightly fused to the base of the petals. Anthers are basifixed, connivent, cone-shaped at the apex, opening on the inside by apical pores or short slits. The superior ovary is composed of two carpels that are fused to form a locule. The single, simple style is long and topped with a twisted, punctate to capitate stigma. Ovule placentation is apical-axile. Fruits are loculicidal capsules. Distribution: The two perennial species are found in southwestern Western Australia, whereas the remaining species are annuals found across tropical northern Australia

LAMIALES

EUDICOTS

Byblis gigantea, near Perth, Western Australia (KD) [408]

with one species extending into southern New Guinea. Phylogeny and evolution: A fossil seed of Byblis has been found in strata from the Mid Eocene in South Australia. The superficial resemblance of Byblis to Drosophyllum led many to treat these taxa as related. They share a similar habit with linear leaves covered in two types of glands. Because the petals of Byblis appear to be free (as in Drosophyllum) they also seem to resemble each other in floral morphology, but close inspection reveals that the petals of Byblis are fused basally. Phylogenetic studies have indicated placement in Lamiales, although with no strong support for interfamilial relationships within that order. A relationship with Linderniaceae is suggested. Genera and species: The sole genus of this family is Byblis with seven species. Carnivory: Like Drosophyllum (Drosophyllaceae), Byblis has two types of glands. The stalked glands work just like fly-paper and catch insects merely by them becoming stuck in the sticky drops of liquid produced by these glands. The stalked glands do

not move toward the insect as in Drosera (Droseraceae). The sessile glands do produce enzymes to digest insects, but it has not yet been shown that the leaves are capable of taking up the amino acids that are released. Similarly to Roridula (Roridulaceae), Byblis has been reported to have a mutualistic relationship with carnivorous bugs, in this case of the genus Setocoris, that are able to move over the surface of the plants without becoming trapped in the mucilage. They eat the dead insects, and the plant benefits from their subsequent defecation, absorbing the nutrients through their roots. The dynamics of these mutualistic relationships are not yet well understood. It is possible that Byblis represents an early evolutionary stage not yet at full carnivory, perhaps with Vandellia (Linderniaceae) as another intermediate stage. Etymology: Βύβλης (Byblis) is a goddess in Greek mythology, whose story was told by Ovid in Metamorphoses. She was the niece of Apollo and fell in love with Caunus, her twin brother. He rejected her love, and she cried endless streams of glistening tears. The droplets on the leaves of Byblis refer to her tears.

Byblis filifolia, Pilbara, Western Australia [408]

409. STILBACEAE Candlesticks family

These are evergreen trees, shrubs and perennial herbs, sometimes with a woody tuber. Leaves are simple, usually whorled, sometimes opposite, rarely alternate, without stipules. Leaf blades have an entire, rarely slightly serrate, margin that is sometimes inrolled and then the leaf ericoid. Venation is pinnate, or leaves have a single midvein only. Inflorescences are usually terminal spikes or heads or axillary panicles, thyrses, spikes or heads, and rarely flowers are solitary. Flowers are bisexual and usually zygomorphic, and subtended by bracteoles that are as long as the sepals. The four or five (to seven) sepals are fused into a tube-, bell- or two-lipped calyx that is persistent. Petals are of the same number as the sepals (four or five to seven) and Plants of the World

559

LAMIALES

Halleria lucida, Royal Botanic Gardens, Kew, UK [409]

EUDICOTS

Nuxia glomerulata, Lowveld National Botanical Garden, South Africa (CD) [409]

fused into a tubular or funnel-shaped corolla. Stamens are as many as the petals, alternate with the petal lobes and are fused to the corolla tube. Anthers are dorsifixed and open inwardly by lengthwise slits. A single staminode is present when there are four stamens. A nectar disk is either small or absent, but oil-secreting glandular hairs are often present in the flower. The superior ovary is composed of two carpels, fused to each other and forming a locule, but with one locule often sterile or unilocular or partly so. Placentation is usually axile or basal, sometimes axile-peltate or apical. The single, simple or slightly bifid style tops the ovary and is terminated by a capitate, punctate or slightly bilobate stigma. Fruits are a loculicidal (sometimes also septicidal) capsule or a nutlet with a persistent calyx and corolla.

560

Retzia capensis, Kogelberg Nature Reserve, Cape Province, South Africa (CD) [409]

two families were subsequently merged. Nuxia, formerly placed in Loganiaceae, also belongs here, as corroborated by shared chemistry. In addition the former tribe Bowkerieae and Halleria (both previously in Scrophulariaceae) and gesneriad-like Charadrophila were also found to belong to this family, making it somewhat polymorphic and difficult to define morphologically. Genera and species: Stilbaceae embrace 12 genera and c. 40 species: Anastrabe (1), Bowkeria (5), Campylostachys (1), Charadrophila (1), Euthystachys (1), Halleria (4), Ixianthes (1), Kogelbergia (2), Nuxia (c. 15), Retzia (1), Stilbe (7) and Thesmophora (1).

Distribution: This family is confined to Sub-Saharan Africa and Madagascar, extending into southern Arabia and the Mascarenes. The main diversity is in the Western Cape of South Africa.

Uses: An alcoholic drink called betsa-betsa is made in Africa from sugarcane juice that is fermented with the bark of Nuxia verticillata as a flavouring. Bowkeria (shell-flower bush), Halleria lucida (African honeysuckle) and Nuxia floribunda (kite tree) are cultivated as ornamentals.

Phylogeny and evolution: Formerly this family was restricted to Campylostachys, Euthystachys and Stilbe, but a molecular phylogenetic study showed a close relationship between Stilbaceae and Retziaceae, and the

Etymology: Στίλβη (Stilbe, meaning ‘shininess’) was a nymph and daughter of the river god Peneus and the naiad Creusa in Greek mythology. She was the mother of Centaurus, ancestor of the centaurs.

Christenhusz, Fay & Chase

Campylostachys cernua, South Africa (JA) [409]

410. MARTYNIACEAE Unicornplant family

These are viscid-pubescent, glandular, usually strongly foetid, annual and perennial herbs, rarely shrubs, sometimes with large roottubers (Craniolaria). Leaves are simple, opposite to nearly opposite without stipules. Blades are usually basally cordate, the margins entire to sinuate, dentate, denticulate or lobed and the venation palmate or pinnate. Glandular hairs consist of unicellular or multicellular uniseriate stalks and multicellular apical heads that produce sticky mucilage. Inflorescences are terminal racemes, and f lowers are bisexual and zygomorphic, subtended by two bracteoles. The calyx is composed of five free or fused sepals, if united then the calyx split to the base on the lower part. The five petals are fused into a corolla with a cylindrical tube at the base and the throat bell- or funnel-shaped,

LAMIALES

EUDICOTS

with five lobes that are sometimes arranged in two lips (with two upper and three lower lobes). Stamens are usually four, two short and two long (didynamous), or two with the second pair staminodial (sterile). Filaments are attached to the corolla, and the anthers are dorsifixed, with the connivent thecae separated at 180º, opening by lengthwise slits. A superior ovary is composed of two carpels that are fused to form a single locule with two parietal placentas expanded to form false partitions, making the ovary appear tetra- or multilocular. The style is simple and topped by a bilobed stigma. Fruits are drupaceous capsules, dehiscing incompletely along their length, terminated by an incurved beak or rostrum and formed by the sterile upper part of the ovary. The capsule becomes somewhat woody and splits in two valves, making it resemble the tusks of an elephant (hence the name Proboscidea). These peculiarly hooked fruits are distributed by large mammals, with the hooks getting caught in their hooves. Ibicella lutea, Rancho Santa Ana Botanical Garden, California, USA [410]

Distribution: This is a family restricted to the Americas. They grow in arid and semi-arid regions, both temperate and tropical. Phylogeny and evolution: Martyniaceae are morphologically similar to Pedaliaceae with which they have often been combined in the past. However, molecular studies have not shown a clear relationship between these two. Martyniaceae are probably sister to Schlegeliaceae, although a relationship with Verbenaceae is also suggested in some studies. Genera and species: Martyniaceae include five genera with 16 species: Craniolaria (3), Holoregmia (1), Ibicella (3), Martynia (1), Proboscidea (8). Uses: Species of Ibicella and Proboscidea are occasionally grown as ornamental plants. Their fruits are peculiarly shaped and sometimes used in dried f lower arrangements.

Carnivory: Mart ynia, Ibicella and Proboscidea are herbs covered in glands that are similar to those in Byblidaceae and Lentibulariaceae and produce a foul-smelling, sticky exudate. Even though the glandular hairs catch many insects, no uptake of amino acids has been demonstrated, but the presence of proteolytic enzymes (similar to trypsin) has been found. It seems that these plants practise, like some species of Solanaceae, an intermediate form of carnivory, the glands functioning mostly to deter herbivores, but catching insects in the process. The plants may benefit to some extent from the partially digested proteins, which are absorbed by their leaves and roots. Etymology: Martynia is named for English botanist Thomas Martyn (1736– 1825), professor of botany at Cambridge University.

Proboscidea louisiana subsp. fragrans, fruits, Ruissalo Botanical Gardens, Turku, Finland [410]

Martynia annua, fruits, Netherlands Antilles AP) [410]

Proboscidea louisiana subsp. fragrans, Ruissalo Botanical Garden, Turku Finland [410]

Martynia annua, Netherlands Antilles (AP) [410]

Plants of the World

561

LAMIALES

EUDICOTS

Ceratotheca triloba, Helsinki Uncarina grandidieri, Royal Botanic Gardens, Kew, UK [411] Botanical Garden, Finland [411]

411. PEDALIACEAE Sesame family

Pedaliaceae are annual and perennial herbs, sometimes deciduous trees and shrubs, sometimes with swollen succulent trunks, in many species with water-storing tuberous roots. Plants often have a strong scent due to their glandular mucilage-excreting hairs. Leaves are usually opposite (sometimes alternate and spiral in Sesamum), simple or pinnately lobed, rarely palmately compound, without stipules. Petioles are sometimes modified into spines. Venation is usually pinnate, sometimes palmate, and blade margins are serrate, lobed or entire. Inflorescences are axillary cymes reduced to a single flower, sometimes a few flowers together, or in Sesamothamnus, a raceme. In the axils of bracteoles, several aborted floral buds are modified into vascularised nectar 562

Christenhusz, Fay & Chase

glands, and nectaries are also often present on the pedicel. Flowers are bisexual and zygomorphic. The five sepals are unequal in size and more or less fused, persistent in fruit. The five petals are fused to form a two-lipped, funnel-shaped corolla (with two upper and three lower lobes) or an irregularly five-lobed tube, rarely with a spur at the base. Stamens are usually four, two longer and two shorter (didynamous), often with an additional staminode. Filaments are fused to the corolla tube, alternating with the petal lobes. Anthers are dorsifixed or basifixed, often the pairs connivent, opening on the inside by longitudinal slits, the thecae inserted at right angles to the filaments and the connective usually with an apical gland. A circular nectar disk is often asymmetrical. The superior ovary is composed of usually two (four in Josephinia) carpels that are fused to form two locules (or eight in Josephinia), the locules often parted in two by secondary walls. The ovary is topped by a single, simple, filiform style and a bilobed, sensitive stigma. The ovules have axile placentation. Fruits are usually a loculicidal (or loculicidal-septicidal) capsule, sometimes a nut or schizocarp. The style base hardens and has beak-shaped outgrowths from the sterile part of the ovary. Seeds are often winged.

Pedalium murex, Malindi, Kenya [411]

Distribution: Pedaliaceae are found mostly in Sub-Saharan Africa, Madagascar, Pakistan, India, Indonesia (Sunda Islands, Sulawesi), New Guinea and Australia. Phylogeny and evolution: Pedaliaceae and Martyniaceae have often been combined because of their morphological resemblance. They differ in placentation and pollen morphology only. Molecular studies have not placed these families with certainty. Pedaliaceae may be sister to Acanthaceae, although their sister-group relationship is not well-supported, whereas Martyniaceae fall with Schlegeliaceae, also without strong support. Genera and species: Pedaliaceae include 13 genera and c. 74 species: Ceratotheca (5), Dicerocaryum (3), Harpagophytum (2), Holubia (1), Josephinia (4), Linariopsis (3), Pedaliodiscus (1), Pedalium (1), Pterodiscus (13), Rogeria (4), Sesamothamnus (6), Sesamum (c. 20) and Uncarina (11). Uses: Sesame (Sesamum indicum) is one of the oldest oilseed crops (it was already cultivated in Mesopotamia and the Indus Valley c. 3500 BC) and is still one of the ten most important oilseed crops worldwide, with Burma, India

LAMIALES

EUDICOTS

and China being the largest producers. The oil is used in similar ways to olive oil or as a flavouring in salads and hot food. The seeds are roasted and used as a sprinkling on breads, cakes and confectionery, or ground into a paste that forms the main ingredient of desserts called halva or of spreads and sauces called tahini, which are common in the cuisines of the Middle East. The polyunsaturated omega 6 oils are healthy and said to prevent stomach cancer. There are various cultivars, with white, black or brown seeds. Ceratotheca sesamoides is also cultivated for its seeds, which are used similarly to sesame. Pedalium murex is locally eaten as a vegetable and has been used to treat impotence in men because it may increase testosterone levels. It is sometimes also promoted as a supplement for bodybuilders, but scientifically conducted trials on humans are still wanting. The fruits of devil’s claw (Harpagophytum procumbens) has large woody hooks that attach themselves to grazing animals, causing much trouble for sheep and cattle, but the fruits of Harpagophytum and Uncarina also make good mouse-traps. Harpagophytum is also used medicinally. Some species are grown occasionally as ornamental plants, particularly Ceratotheca triloba, Pterodiscus, Sesamothamnus and Uncarina.

412. ACANTHACEAE Bear’s-breeches family

Etymology: Pedalium is the Latinised form of the Greek πηδάλιον (pedalion), a rudder.

These are bisexual or rarely unisexual annual and perennial herbs, shrubs, vines and rarely evergreen trees and mangroves (Avicennia). Most are terrestrial, but there are some epiphytes and aquatics. Leaves are opposite, simple or pinnately lobed or dissected, without stipules. Blades are pinnately veined with serrate, crenate or entire margins. Hairs are sometimes glandular, producing ethereal oils, salt glands are present in Avicennia and extrafloral nectaries are sometimes present on leaf blades (e.g. in Ruellia). Inflorescences are terminal or axillary thyrses, spikes, racemes or false umbels, but sometimes flowers are solitary. Bracts are often large and showy, sometimes with involucres subtending partial inflorescences or the bracts sometimes leaflike. The flowers are bisexual or unisexual and zygomorphic (in Avicennia almost actinomorphic) and sometimes inverted (resupinate). The usually five (rarely three to

Carlowrightia parviflora, San Antonio Botanical Garden, Texas, USA [412]

Pseuderanthemum sinuatum, Helsinki Botanical Garden, Finland [412]

16) sepals are free or fused and sometimes strongly reduced. The five petals are fused and form a corolla that is more or less divided into a five-lobed, two-lipped corolla (two upper and three lower lobes), sometimes strongly curved and scalloped. The two or four free stamens are of the same length or two long and two short (didynamous), three or one then sterile staminode (staminodial). Filaments are fused to the corolla tube. Anthers are dorsifixed and open by lengthwise slits, and the connective sometimes has an appendage. A nectar disk is circular or composed of five separate glands, surrounding the ovary. A superior ovary is composed of two fused carpels, each forming a locule (unilocular in Mendoncia), topped by a single style and a variously shaped stigma (often bifid). Male flowers lack a pistillode. Ovules are usually axile (rarely parietal). Fruits are usually loculicidal, explosive capsules, sometimes a few-seeded drupe. Distribution: Acanthaceae are predominantly tropical with a few representatives in the warm-temperate zones of North America, southern South America, southern Europe, western, central and northeastern Asia to Korea and Japan. Phylogeny and evolution: Diversification of Acanthaceae may have taken place c. 57–81 million years ago, but the crown group has been estimated to have evolved c. 90 million Ruttya fruticosa, Royal Botanic Gardens, Kew, UK [412]

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years ago, depending on the calibration points. Avicennia was found to be embedded in Acanthaceae, as sister to Thunbergioideae. Avicennia is morphologically odd due to its adaptations to the mangrove habitat. Nelsonioideae, previously included in Scrophulariaceae or considered intermediate between the two families, are sister to the rest of Acanthaceae (including Avicennioideae and Thunbergioideae). Thomandersia was previously included in this family, but it is closer to Verbenaceae and now placed in a separate family, Thomandersiaceae. Most modern distribution patterns have been achieved by long-distance dispersal events from the Old World to the New World during the past 20 million years or so. Generic delimitation in Acanthaceae is still poor, and several realignments and recircumscriptions will be necessary to achieve monophyly. Genera and species: Acanthaceae include c. 210 genera, c. 4,000 species, in four subfamilies: Nelsonioideae (7 genera) – Anisopetalum (3), Elytraria (17), Gyno-

Nelsonia canescens, Tanzania (RB) [412]

Avicennia germinans, Guadeloupe [412]

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craterium (1), Nelsonia (1), Ophiorhiziphyllon (5), Saintpauliopsis (1) and Staurogyne (c. 80); Avicennioideae (1 genus) – Avicennia (4–7); Thunbergioideae (5 genera) – Anomacanthus (2), Mendoncia (c. 60), Meyenia (1), Pseudocalyx (7) and Thunbergia (c. 90); Acanthoideae (196 genera) – Acanthopale (c. 15), Acanthopsis (7), Acanthostelma (1), Acanthus (22), Afrofittonia (1), Ambongia (1), Ancistranthus (1), Andrographis (c. 20), Angkalanthus (1), Anisacanthus (8), Anisotes (19), Apassalus (3), Aphanosperma (1), Aphelandra (c. 180), Ascotheca (1), Asystasia (c. 70), Ballochia (3), Barleria (c. 300), Barleriola (6), Benoicanthus (2), Blechum (6), Blepharis (130), Borneacanthus (6), Boutonia (1), Brachystephanus (11), Bravaisia (3), Brillantaisia (12), Brunoniella (6), Calacanthus (1), Calycacanthus (1), Camarotea (1), Carlowrightia (c. 25), Celerina (1), Cephalacanthus (1), Chalarothyrsus (1), Chameranthemum (4), Chileranthemum (2), Chlamydacanthus (4), Chlamydocardia (4), Chlamydostachya (1), Chorisochora (3), Chroestes (4), Cindasia (1), Clinacanthus (2),

Thunbergia grandiflora, Singapore Botanical Garden [412]

Clistax (2), Codonacanthus (2), Conocalyx (1), Cosmianthemum (8), Crabbea (16), Crossandra (55), Crossandrella (2), Cyclacanthus (2), Cynarospermum (1), Cyphacanthus (1), Danguya (1), Dasytropis (1), Diceratotheca (1), Dichazothece (1), Dicladanthera (2), Dicliptera (c. 150), Dischistocalyx (c. 20), Dolichostachys (1), Duosperma (26), Duvernoia (2), Dyschoriste (c. 65), Ecbolium (22), Echinacanthus (4), Encephalosphaera (2), Epiclastopelma (2), Eranthemum (c. 30), Eremomastax (1), Eusiphon (3), Filetia (8), Fittonia (2), Forcipella (5), Geissomeria 15), Glossochilus (2), Graphandra (1), Graptophyllum (c. 10), Gymnostachyum (c. 30), Gypsacanthus (1), Haplanthodes (4), Harpochilus (3), Hemigraphis (c. 30), Henrya (2), Herpetacanthus (10), Heteradelphia (2), Holographis (10), Hoverdenia (1), Hulemacanthus (2), Hygrophila (c. 25), Hypoestes (c. 40), Ichtyostoma (1), Ionacanthus (1), Isoglossa (c. 50), Isotheca (1), Jadunia (2), Jurausia (2), Justicia (c. 600), Kalbreyeriella (3), Kosmosiphon (1), Kudoacanthus (1), Lankesteria (7), Thunbergia mysorensis, Royal Horticultural Society Garden, Wisley, UK [412]

LAMIALES

EUDICOTS

Ruellia brittoniana, Turbaco, Colombia [412]

Aphelandra schiedeana, Botanical Garden Berlin-Dahlem, Germany [412]

Lasiocladus (5), Leandriella (2), Lepidagathis (c. 115), Leptosiphonium (10), Linariantha (1), Louteridium (6), Lychniothyrsus (5), Mackaya (1), Marcania (1), Megalochlamys (3), Megalostoma (1), Megaskepasma (1), Melittacanthus (1), Mellera (5), Metarungia (3), Mexacanthus (1), Mimulopsis (c. 30), Mirandea (6), Monothecium (3), Morsacanthus (1), Neriacanthus (4), Neuracanthus (c. 30), Oplonia (19), Oreacanthus (4), Orophochilus (1), Pachystachys (11), Pararuellia (5), Pelecostemon (1), Perenideboles (1), Pericalypta (1), Peristrophe (c. 15), Petalidium (c. 35), Phaulopsis (22), Phialacanthu s (5), Phlogacanthu s (c. 15), Physacanthus (5), Podorungia (5), Poikilacanthus (6), Polylychnis (2), Populina (2), Pranceacanthus (1), Pseuderanthemum (c. 60), Pseudodicliptera (2), Pseudoruellia (1), Psilanthele (1), Ptyssiglottis (33), Pulchranthus (4), Razisea (5), Rhinacanthus (6), Rhombochlamys (2), Ritonia (3), Ruellia (c. 150), Ruelliopsis (3), Rungia (c. 50), Ruspolia (4), Ruttya (3), Salpixantha (1), Samuelssonia (1), Sanchezia (c. 25), Sapphoa (2), Satanocrater (4), Sautiera (1), Schaueria (8), Schaueriopsis (1), Sclerochiton (19), Sebastiano-schaueria (1), Sinoacanthus (3), Spathacanthus (3), Sphacanthus (2), Sphinctacanthus (1), Spirostigma (1), Stenandrium (43), Stenostephanus (c. 75), Stenothyrsus (1), Streblacanthus

Pachystachys lutea, Singapore Botanical Garden [412]

(7), Streptosiphon (1), Strobilacanthus (1), Strobilanthes (c. 400), Strobilanthopsis (5), Suessenguthia (6), Tessmanniacanthus (1), Tetramerium (28), Thysanostigma (2), Trichanthera (2), Trichaulax (1), Trichocalyx (2), Trichosanchezia (1), Vavara (1), Whitfieldia (c. 10), Xantheranthemum (1), Xerothamnella (2), Yeatesia (3) and Zygoruellia (1). Uses: Wood of Avicennia mangroves is often used for charcoal and fuel. The wood is unique in that the heartwood floats, whereas the sapwood sinks. Many genera make excellent ornamentals for the garden or as house plants, flowering for a long time and often having decoratively marked leaves. Particularly commonly cultivated are members of the genera Acanthus (bear’s breeches), Aphelandra (zebra plants), Asystasia (creeping foxgloves), Barleria (porcupine f lower, bush violets), Brillantaisia, Crossandra (firecracker f lowers), Eranthemum (blue sage), Fittonia (nerve plants), Graptophyllum (caricature plants), Hemigraphis (red ivy), Hypoestes (polka dot plants), Justicia (waterwillows, shrimp plants, etc.), Megaskepasma (Brazilian red-cloak), Pachystachys (lollipop plants), Pseuderanthemum, Ruellia (wild petunia), Ruttya (jammy mouth), Sanchezia, Strobilanthes (Persian shield), Suessenguthia, Thunbergia (T. alata: black-eyed Susan; T.

Asystasia gangetica subsp. micrantha, Singapore [412]

grandiflora: clockvine; T. mysorensis: lady’s slipper vine) and Whitfieldia. Hygrophila species are commonly cultivated as aquarium plants, especially H. difformis (water wisteria) and H. polysperma (Indian swampweed). Corinthian columns: Acanthus leaves have been the inspiration for one of the most common foliage-based ornaments in architecture: the Corinthian capital. They are based on the deeply lobed leaves of Acanthus mollis or A. spinosus, both common species in the eastern Mediterranean, where the design was first used. In Ancient Greek architecture, the acanthus ornament appears extensively in Corinthian and composite capitals atop columns. The oldest known example of a Corinthian column is found in the Temple of Apollo Episcurius in classical Arcadia (now Messenia, Greece), built c. 450–420 BC. Later, the Romans elaborated the acanthus design, which they chose as their most popular design for grand buildings. The motif continued in Byzantine times and then into Mediaeval art, after which it was revived during the Renaissance and became popular again in Neoclassical buildings of the 19th century. Why acanthus was chosen as an architectural motif is not certain, but it has been suggested that it originated as a sculptural version of the palmette, an architectural design used in ancient Egypt and only later coming to Plants of the World

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Justicia californica, Palm Springs, California, USA [412]

Acanthus mollis, Sicily, Italy [412]

Acanthus leaf decoration on corinthian capital Brillantaisia nyanzarum, Royal Botanic Gardens, Kew, UK [412] from Delos, Roman, 2nd or 3rd century AD, British Museum, London, UK [412]

resemble acanthus leaves. However, the Roman writer Vitruvius explained that the design of the Corinthian column was inspired by the sight of an acanthus leaf growing through a basket of toys on the grave of a little girl. It is also said that acanthus symbolises life endurance and resurrection, later associated with the suffering of Christ. Whatever the origin and symbolism, acanthus leaves now crown columns of many buildings worldwide and have become an important visual aspect of architecture, furniture and other items. Etymology: Acanthus is a Latinised form of Greek άκανθα (akantha), a thorn.

413. BIGNONIACEAE Trumpetvine family

These are usually evergreen (rarely deciduous) vines, shrubs, trees and perennial herbs. When climbing, the stem twining or with leaf tendrils. Leaves are usually opposite, 566

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sometimes whorled (rarely alternate), usually pinnately compound, sometimes twice or more times pinnate or palmately compound, occasionally unifoliolate or simple (sometimes bipinnate or more, or palmately compound, or simple/unifoliolate), terminal leaflets present or replaced by a tendril. Margins entire, pinnately or palmately lobed, rarely toothed, and venation is pinnate or palmate. Leaves are petiolate, but lack a stipule or leaf sheath. Extrafloral nectaries are often present on leaf tips and buds. Inflorescences are terminal or axillary dichasia, cincinni, thyrses, raceme, or flowers are solitary. Flowers are zygomorphic (in some cases only weakly so) and bisexual. Sepals are fused to form a bell-shaped, fivelobed calyx, sometimes truncate or bilobed to campanulate, often with nectaries on the outside. The five petals are fused into a bellor funnel-shaped corolla that often has two lips, with two upper and three lower lobes. Four stamens alternate with the petals; two are longer and two shorter (didynamous) or all of the same length. A fifth stamen is absent or usually modified into a staminode, but in Catalpa and Paragonia (now within Tanaecium) there are two fertile stamens and three staminodia. Filaments are fused to the petal tube, and the dorsifixed anthers are often fused or confluent head-to-head opening on the inside by longitudinal slits. A nectar disk is circular, cup-shaped or absent. The superior

ovary is composed of two carpels that are fused so each form a locule (rarely with one or four locules), often bearing nectaries on the upper side. The single, simple style is topped by a capitate or broadly bilobed sensitive stigma. Ovules have axile or parietal placentation. Fruits are loculicidal or septicidal capsules (sometimes large and lignified), sometimes indehiscent, usually with flat, winged seeds. Distribution: This pantropical family has a few representatives in warm temperate regions of North America and East Asia. It has the greatest diversity in tropical South America. Phylogeny and evolution: Fossil winged seeds attributable to Bignoniaceae are reported from the Oligocene of North America and Europe. Fossil seeds from Washington State have been dated to c. 49 million years, and for this reason it seems that the family has a New World origin. Molecular analyses have shown that there have been at least five dispersals to the Old World and one dispersal back to the Americas. Palmate leaves have evolved several times in New World Bignoniaceae. Tabebuia in the traditional sense is paraphyletic to some genera (Amphitecna and Crescentia). Genera and species: This family has c. 82 genera and 870 species: Adenocalymma (79),

LAMIALES

EUDICOTS

Amphilophium (43), Amphitecna (19), Anemopaegma (45), Argylia (13), Astianthus (1), Bignonia (27), Callichlamys (1), Campsidium (1), Campsis (2), Catalpa (10), Catophractes (1), Chilopsis (1), Colea (26), Crescentia (6), Cuspidaria (16), Cybistax (1), Delostoma (4), Deplanchea (8), Digomphia (3), Dinklageodoxa (1), Dolichandra (8), Dolichandrone (10), Eccremocarpus (3), Ekmanianthe (2), Fernandoa (15), Fridericia (69), Godmania (2), Handroanthus (33), Heterophragma (2), Hieris (1), Incarvillea (17), Jacaranda (47), Kigelia (1), Lamiodendron (1), Lundia (14), Manaosella (1), Mansoa (12), Markhamia (5), Martinella (2), Mayodendron (1), Millingtonia (1), Neojobertia (2), Neosepicaea (4), Newbouldia (1), Nyctocalos (3), Ophiocolea (10), Oroxylum (1), Pachyptera (4), Pajanelia

Campsis grandiflora, Royal Botanic Gardens, Kew, UK [413]

(1), Pandorea (9), Paratecoma (1), Parmentiera (10), Pauldopia (1), Perianthomega (1), Perichlaena (1), Phyllarthron (18), Phylloctenium (2), Pleonotoma (17), Podranea (2), Pyrostegia (2), Radermachera (17), Rhigozum (7), Rhodocolea (13), Romeroa (1), Roseodendron (2), Santisukia (2), Sparattosperma (2), Spathodea (1), Sphingiphila (1), Spirotecoma (4), Stereospermum (25), Stizophyllum (3), Tabebuia (74, polyphyletic), Tanaecium (16), Tecoma (9), Tecomanthe (5), Tecomella (1), Tourrettia (1), Tynanthus (15), Xylophragma (7) and Zeyheria (2). Uses: Musical instruments and bowls are often made from calabashes, the fruit of Crescentia cuyete and related species. As semillas de jicaro, the seeds are cooked to

Incarvillea mairei, Yunnan, China [413]

Chilopsis linearis subsp. arcuata, Santa Barbara Botanical Garden, California, USA [413]

produce a drink in Nicaragua. The flowers of the mangrove trumpet tree (Dolichandrone spathacea) are eaten as kaeng in Thai cuisine, and leaves and young pods of ‘midnight horror’ (Oroxylum indicum) are eaten as a vegetable in Southeast Asia, especially in Laos. Chica (Fridericia chica, formerly Arrabidaea) is a cultigen used for dying barkcloth in traditional Amazonian societies. A blue dye can be produced from the leaves of Cybistax antisyphilitica. Some Bignoniaceae are locally used in medicine (e.g. Handroanthus) or as an aphrodisiac (Anemopaegma). Koribo (Tanaecium nocturnum) is a hallucinogen, made into a snuff from the roasted, dry leaves. The snuff, which smells of almonds, is inhaled

Catalpa speciosa, Ruissalo Botanical Garden, Turku, Finland [413]

Colea seychellarum in fruit, Mahé, Seychelles [413]

Crescentia cujete, Guadeloupe [413]

Plants of the World

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LAMIALES

Kigelia africana, Rio de Janeiro Botanical Garden, Brazil [413]

EUDICOTS

Amphilophium buccinatorium, Sicily, Italy [413]

Spathodea campanulata, cultivated in Guadeloupe [413]

during special ceremonies of Amerindian tribes, particularly for healing ceremonies and rites of passage. Tynanthus guatemalensis contains eugenol, smells of cloves and is used as a spice. It was valued as a tea by the Mayans. A few species of Catalpa, Crescentia, Handroanthus, Jacaranda, Markhamia, Millingtonia, Oroxylum, Pajanelia, Roseodendron, Stereospermum, Tabebuia and Zeyheria are locally harvested for timber or firewood. Many species have beautiful flowers and are therefore commonly cultivated in gardens especially in the tropics, but also in temperate zones. Cultivated ornamentals include Amphilophium crucigerum (monkey comb), 568

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Tecoma capensis, Wundanyi, Kenya [413]

Pyrostegia venusta, St Denis, Réunion [413]

Tabebuia rosea, St Petersburg, Florida. USA [413]

Bignonia capreolata (crossvine), Campsidium valdivianum ( pilpilvoqui), Campsis grandiflora (Chinese trumpet-flower), C. radicans (trumpet creeper), Catalpa (Indian bean or cigar-tree), Chilopsis linearis (desert willow), ×Chitalpa tashkentensis (the hybrid between Catalpa bignonioides and Chilopsis linearis), Dolichandra unguis-cati (cat’s claw creeper), Eccremocarpus scaber (Chilean glory flower), Handroanthus (ipê), Incarvillea (hardy gloxinia), Jacaranda mimosifolia (blue jacaranda), Kigelia africana (sausage tree), Mansoa alliacea (garlic vine), Markhamia lutea (Nile trumpet, siala tree), Millingtonia hortensis (tree jasmine), Newbouldia laevis (boundary tree), Pandorea jasminoides (bower plant), P. pandorana (wonga-wonga

vine), Parmentiera (candle tree), Podranea ricasoliana (pink trumpet vine), Pyrostegia venusta (flame vine), Radermachera sinica (Asian bell tree, emerald tree, china doll), Spathodea campanulata (African tulip tree), Tabebuia (trumpet roble), Tecoma capensis (Cape honeysuckle) and T. stans (yellow trumpetbush). Some species can become invasives, and particularly Spathodea campanulata and Tecoma stans can become problematic outside their native range. Etymology: Bignonia is named for French statesman and priest Abbé Jean Paul Bignon (1662–1743), who was librarian for King Louis XIV of France. He was a tutor of Joseph Pitton de Tournefort, who named this vine for him.

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EUDICOTS

These are carnivorous, perennial and annual herbs that grow terrestrially, aquatically and as epiphtically. Roots are present (Pinguicula) or absent, sometimes forming swollen tubers, but stems are often modified into rhizoids (Utricularia), stolons or rhizomes, and they are

rarely developed. Bladder-like (Utricularia) or V-shaped (Genlisea) traps can be found on rhizoids, stolons or leaves. Pinguicula does not have such traps, but its leaves are covered with sessile and stalked glands that act as fly-paper traps. Leaves are alternate, spirally arranged and often form a rosette, simple or variously divided, sometimes peltate or without a lamina and stipules. Leaf blades have entire, serrate or lobed margins, with glands or traps secreting proteolytic enzymes, the venation pinnate. Inflorescences are racemes or spikes that are often reduced to a solitary flower, the peduncle usually erect, sometimes twining. Flowers are zygomorphic and bisexual. The five sepals are fused into a bilabiate calyx with two, four or five lobes. The five petals are fused to form an equally five-lobed or

two-lipped corolla, the lower lip entire or with two or three (to six) lobes, the upper lip entire or with two or three lobes. A nectarsecreting subulate, cylindrical, conical or saccate corollar spur projects backward and is sometimes bent inwards. The two stamens are inserted at the base of corolla tube and have short, curved or straight linear filaments and dorsifixed anthers with lengthwise dehiscence. A nectar disk is absent. The superior ovary is composed of two carpels that are fused to form a single locule, which has free-central or basal placentation. The ovary is topped by a single style and a bilobed stigma, or the stigma is sessile. Fruits are loculicidal, circumscissile, or rarely indehiscent capsules surrounded by the persistent calyx, but sometimes a one-seeded nutlet.

Pinguicula alpina, Yunnan, China [414]

Utricularia warmingii, Colombia (MF) [414]

Utricularia bisquamata, Hampton Court Flower Show, UK [414]

414. LENTIBULARIACEAE Bladderwort family

Pinguicula primuliflora, Carolina Beach, North Carolina, USA [414]

Utricularia gibbosa, Guadeloupe [414]

Utricularia gibbosa, Royal Botanic Gardens, Kew, UK [414]

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LAMIALES Distribution: Utriculariaceae are nearly a cosmopolitan family but are absent from great desert areas because the plants are often strongly dependent on water. Phylogeny and evolution: Genlisea has the smallest genome known for any angiosperm. Among vascular plants, only the genomes of some lycopods of the genus Selaginella (Selaginellaceae) are smaller. Utricularia gibba has lost almost all of its non-coding DNA, but remains functionally the same as related species that have maintained this DNA component. Genome-size variation is substantial in Lentibulariaceae, and mutation rates are among the highest in angiosperms. Previously, there were two additional genera recognised in this family, but Biovularia and Polypompholix, segregated on the basis of a slighly different shape of the suction traps, have been shown in molecular studies to belong to Utricularia, with which they share many characteristics. The crown group of Lentibulariaceae is estimated to be c. 28–42 million years old. Genera and species: The three genera of this family encompass 316 species: Genlisea (21), Pinguicula (c. 80) and Utricularia (c. 215). Uses: Butterwort (Pinguicula vulgaris) has been used for a long time in northern Europe to curdle milk, making a buttermilk-like fermented dairy product. Today, this is still consumed in the Nordic countries ( filmjölk, tätmjølk, viili). Pinguicula produces a bactericide that prevents insects from rotting while being digested. The leaves have been known to have healing power and were used in European traditional medicine to heal sores and clean wounds. Pinguicula also makes a good natural flypaper, occasionally used for pest control in greenhouses. Several species are grown as ornamental plants but usually as curiosities in specialist collections Three ways to catch prey: Lentibulariaceae have evolved three ways to catch their prey. The simplest method of carnivory is displayed by the fly-paper traps of Pinguicula, which

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EUDICOTS

form a rosette of broad leaves that have stalked and sessile glandular hairs on the upper surface of the leaf, secreting mucilage and proteolytic enzymes. Pinguicula catches mostly small insects. Genlisea has peculiarly structured leaves that are split into two and twisted into capillary tubes that are joined at the tip in a V-shape, like eel traps. The spiralling grooves down their length trap soilborne invertebrates and protozoa because the inwardly pointing hairs prevent them from escaping. The only direction they can move is to the apex of the tube where they are digested. The third type are the bladder-shaped suction traps found in Utricularia. These are bladders with a trapdoor at the mouth, which have sensitive hairs that release the trapdoor. They rapidly spring open, and the small animal that released the trigger is sucked into the bladder by the negative pressure on the inside. The trapdoor then closes, and the bladder secretes enzymes and absorbs the products of digestion. The prey may die in part of anoxia. Aquatic species have these bladders on photosynthetic submerged branches, whereas terrestrial species have the bladders on achlorophyllous runners. Aquatic species usually have larger bladders with which they can feed on more substantial prey such as water fleas (Daphnia), nematodes, fish fry, mosquito larvae and even tadpoles. One species, Utricularia humboldtii, is peculiar in that it only inhabits the water tanks (phytotelmata) of Brocchinia species (Bromeliaceae), which are also said to be carnivorous. The flowers are produced on relatively long pedicels held away from the carnivorous leaves, possibly to prevent carnivory of potential pollinators. Even though most species are carnivorous, Utricularia purpurea may have partially lost its appetite for meat. This Australian species can still trap and digest prey in its traps, but does so sparingly. Instead algae, zooplankton and debris are present in the bladders, suggesting that U. purpurea favours mutualism rather than carnivory. Etymology: The derivation of Lentibularia is uncertain. It was first coined by Gesner and probably refers to the lentil-shaped bladders.

Lentibularia is a synonym of Utricularia, utricula being Latin for a small bottle, also referring to the bladder traps.

415. SCHLEGELIACEAE Higuerito family

These are evergreen trees, shrubs and vines with a whitish bark. They are terrestrial or more often epiphytic. Leaves are opposite, simple, without stipules but with winged petioles. Leaf blades have pinnate venation, often bear extraf loral nectaries and are serrate or almost entire. Sometimes the lamina is covered below by small glandular hairs with radially arranged head cells. Inflorescences are axillary cymes, compound in thyrses or racemes (in Exarata). Flowers are bisexual and zygomorphic. The (three to) five sepals are fused into a short tube and in Schlegelia the sepals bear sunken groups of nectariferous glands. The five petals are fused into tubular, two-lipped corollas (the Schlegelia parasitica, Fairchild Botanical Garden, Florida, USA (CD) [415]

LAMIALES

EUDICOTS

upper lip with two, the lower lip with three lobes). Stamens are four, two longer and two shorter (didynamous), alternating with the petals and fused to the corolla tube. Anthers are basifixed and open by longitudinal slits. A sterile stamen (staminodium) is present or not. A nectar disk is sometimes composed from vascularised carpel bundles or absent. The superior ovary is composed of two carpels fused to form one locule with axile placentation. It is topped by a single, simple style and a capitate, bi- or tri-lobed stigma. Fruits are woody berries surrounded by the persistent calyx. Distribution: This Neotropical family is found in the Caribbean, Mesoamerica and northern South America.

Etymology: Schlegelia is named for German zoologist Hermann Schlegel (1804–1884), conservator at the Leiden zoological museum and later director of that institution.

416. THOMANDERSIACEAE West-African-bitterbush family

Genera and species: The family includes four genera and 22 species: Exarata (1), Gibsoniothamnus (8), Schlegelia (12) and Synapsis (1).

A family of evergreen shrubs and small trees (sometimes vines), these have simple, opposite leaves of unequal size (anisophyllous) without stipules. Petioles are swollen at the base and apex, and leaf blades have pinnate venation and entire, serrate or lobed margins, often bearing extrafloral nectaries. Inflorescences are terminal or axillary, raceme-like cymes. Flowers are bisexual and zygomorphic. The five sepals are persistent in fruit and on the outside have vascularised nectaries that are each up to three millimetres wide and surrounded by a swelling. The five petals are fused into a two-lipped corolla (the upper lip with two lobes and the lower

Thomandersia congolana, fruit, Fougère Brulée, Lope Reserve, Gabon (CD) [416]

Thomandersia congolana, Fougère Brulée, Lope Reserve, Gabon (CD) [416]

Phylogeny and evolution: Schlegeliaceae were usually included in Bignoniaceae or Scrophulariaceae, often placed as ‘transitional genera’, but molecular phylogenetic studies place them close to Lentibulariaceae, Martyniaceae and possibly Thomandersiaceae. Support for their placement is, however, weak.

with three). Nectaries are present between the stamens. The four stamens with two longer (didynamous) and a sterile stamen alternate with the petals, the filaments fused to the corolla tube. Anthers are dorsifixed and open outwardly by longitudinal slits. A superior ovary is composed of two carpels fused to each form a locule, with a waist between them, with nectariferous tissue at the base. Placentation is axile with expanded placentae. The single, simple style is topped by a cylindrical or bilobed stigma. Fruits are woody, loculicidal, non-explosive capsules with a persistent and enlarging calyx. Each seed is situated on a flattened, elongate, hookshaped ejaculator (modified funicle), the seed having a large hilum. Distribution: This family is restricted to western and central Africa, from southern Liberia to the Congo and the southern Central African Republic. Phylogeny and evolution: Thomandersia was previously placed in Acanthaceae or Schlegeliaceae. It was separated in its own family on the basis of its different leaf and anther anatomy, a segregation that was later confirmed by molecular studies, although its exact position in Lamiales remains unresolved. Pollen morphology resembles that of Pedaliaceae, but molecular studies place it as a relative of Bignoniaceae, Schlegeliaceae or Verbenaceae, always with weak support. Genera and species: The single genus in this family is Thomandersia with six species. Uses: Thomandersia laurifolia (and probably other species also) has medicinal properties. It is used by local people and gorillas in central Africa to rid themselves of parasites. The leaves taste bitter. Etymology: Thomandersia is named for Scottish physician and botanist Thomas Anderson (1832–1870), who first described this new genus (as Scytanthus, a later homonym of a genus of Apocynaceae). Anderson was director of the Calcutta Botanical Garden.

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EUDICOTS

417. VERBENACEAE Vervain family

These are bisexual (rarely unisexual in some species of Citharexylum), evergreen and deciduous trees, shrubs, vines, perennial and annual herbs. Stem and branches are often square in cross-section, and plants are often aromatic. Leaves are usually opposite, rarely whorled or alternate, simple or pinnately compound, entire or lobed, sometimes scalelike, without stipules, but the petiole bases are fused by thin tissue across the node. Leaf blades have pinnate venation with serrate, crenate, lobate or entire margins.

Lippia rosmarinifolia, Galápagos Islands [416]

Lantana camara, Réunion [416]

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Extrafloral nectaries are present in many species on the petiole and lower side of the leaf blade. Inflorescences are terminal or axillary racemes, spikes, heads or simple cymes, sometimes subtended by an involucre of large, colourful petaloid bracts, but these are usually small and green. Flowers are usually bisexual and zygomorphic. The (four or) five sepals are fused into a cup-shaped, persistent calyx. The (four or) five petals are fused into a funnel-shaped corolla. Stamens are usually four, two longer (didynamous), sometimes with a sterile stamen (staminode) as well (but often absent). Filaments are fused to the corolla tube, alternating with the petal lobes, and anthers are basifixed or dorsifixed and open inwardly by longitudinal slits. The connective sometimes bears a glandular appendage. A nectar disk forms a ring around the ovary. The superior ovary is composed of one or two fused, transverse carpels that each form a locule or are each divided in two by secondary septa forming four locules. One carpel is aborted in many

Duranta erecta, garden in Palm Springs, California, USA [416]

species, and Duranta has four carpels forming eight locules. Placentation is usually basal or axile (rarely parietal), and ovules are attached directly to the margins of the secondary carpellary septa (as in Lamiaceae). The ovary is topped by a single, simple or bilobed style bearing capitate or slightly square stigmas that are often unequally swollen and glandular. Fruits are dry or fleshy schizocarps with two or four one-seeded nutlets or two, two-seeded parts or a drupe with one, two or four stones surrounded by the persistent calyx. Fruits are occasionally winged. Distribution: A nearly cosmopolitan family, they extend to the Arctic and Antarctic, but are absent from the driest and coldest regions. Phylogeny and evolution: On morphological grounds, Verbenaceae were always considered close to Lamiaceae, but molecular phylogenetic studies have shown that Verbenaceae form a clade with Lentibulariaceae and possibly Schlegeliaceae and Thomandersiaceae. Junellia coralloides, Royal Horticultural Society Alpine Show, London, UK [416]

Petraea volubilis, garden in Nairobi, Kenya [416]

LAMIALES

EUDICOTS

Rhaphithamnus spinosus, Royal Botanic Gardens, Kew, UK [416]

Stachytarpheta jamaicensis, Tapiraí, Brazil [416]

Verbena bonariensis, Royal Botanic Gardens, Kew, UK [416]

Verbenaceae appear to have originated in tropical South America, and fossil pollen grains are reported from Neogene deposits. Previously identified erroneously as sister to Bignoniaceae in some analyses, Petreaceae are now firmly placed as sister to the rest of Verbenaceae. Several genera formerly placed in Verbenaceae (e.g. Callicarpa, Clerodendrum, Congea and Vitex) have now moved to Lamiaceae, and Avicennia, also sometimes placed here, is now included in Acanthaceae. Some genera have been demonstrated not to be monophyletic, and reclassification of these will be necessary in the future.

used as a herb to flavour liquor; it has also been used to make a (sedative) tea and in the perfume industry as an effective lemonscent (commonly used in air-freshners and potpourri). Mexican oregano (Lippia graveolens) and Hispanic thyme (L. micromera) can be used as herbs to flavour food. Other Lippa species are locally used medicinally. The honeyherb (Phyla dulcis) has been used since historical times (it was known by the Aztecs) as a sweetener. The sweetness is caused by hernandulcin, a sesquiterpenoid compound. Common vervain (Verbena officinalis) has quinine-like properties and has been used against infections and fever in former times. Verbenalin, a compound found in Verbena, has sleep-promoting properties. The wood of Citharexylum (especially fiddlewood, bois fidèle, C. spinosum) is valued for carpentry. Several species are popular ornamental plants, especially golden dewdrop (Duranta erecta), garden verbena (Glandularia ×hybrida, often erroneously called ‘Verbena speciosa’), Spanish f lag (Lantana camara), trailing lantana (L. montevidensis), purple wreath (Petrea volubilis), lippia or frogfruit (Phyla nodiflora), red snakeweed (Stachytarpheta mutabilis), purpletop vervain (Verbena bonariensis) and slender vervain (V. rigida). Verbena bonariensis frequently escapes from gardens, and Lantana camara has become a problematic invasive, especially in tropical

areas and on islands. Berries of Lantana are not edible.

Genera and species: Verbenaceae include c. 32 genera and c. 1,000 species: Acantholippia (6), Aloysia (37), Baillonia (1), Bouchea (9), Casselia (11), Chascanum (27), Citharexylum (c. 70), Coelocarpum (7), Diostea (1), Dipyrena (1), Duranta (17), Glandularia (c. 95), Junellia (c. 40), Lampayo (3), Lantana (c. 150), Lippia (c. 200), Mulguraea (11), Nashia (7), Neosparton (4), Parodianthus (2), Petrea (11), Phyla (c. 11), Pitraea (1), Priva (c. 20), Recordia (2), Rehdera (3), Rhaphithamnus (2), Stachytarpheta (c. 100), Tamonea (7), Verbena (200–250), Xeroaloysia (1) and Xolocotzia (1). Uses: Lemon verbena (Aloysia triphylla, also sometimes sold as Lippia citriodora), is

Etymology: Verbena is a classical Latin word used for scented leaves of ceremonial plants. Sacred to the Romans, verbena was often presented on altars dedicated to Jupiter, but these offerings may have been any type of herb, not necessarily plants that we call verbena today. This association of divinity (Egyptians called it the ‘tears of Isis’) has been embedded in local culture, and verbena has thus been said to have staunched the wounds of Jesus Christ or to protect against the devil, witchcraft or even vampires.

418. LAMIACEAE Mint family

This family is composed of usually aromatic annual, biennial and perennial herbs, shrubs and sometimes trees, rarely vines. Young stems and branches are often quadrangular in cross-section. Leaves are usually opposite, Plants of the World

573

LAMIALES

Callicarpa bodinieri, private garden, Kingston upon Thames, Surrey, UK [418]

Pityrodia bartlingii, Mt Benia, Western Australia [418]

EUDICOTS

Hemiandra pungens, Mt Benia, Western Australia [418]

Vitex ovata, Singapore [418]

Clerodendron trichocarpa in fruit, Royal Botanic Gardens, Kew, UK [418]

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Prostanthera ovalifolia, Royal Botanic Gardens, Kew, UK [418]

Chloanthes parviflora, Australian National Botanic Garden, Canberra [418]

Tectona grandis, in fruit, Cameroon (CD) [418]

Dracocephalum argunense, Helsinki Botanical Garden, Finland [418]

Scutellaria orientalis, near Artvin, Turkey [418]

Salvia przewalskii, Yunnan, China [418]

Monarda didyma, De Border Nursery, Delden, the Netherlands [418]

LAMIALES

EUDICOTS

sometimes whorled, rarely alternate, mostly simple, sometimes pinnately or palmately compound or trifoliate, without stipules. Blades are serrate, crenate, lobed or entire, often leathery or bullate, sometimes inrolled and ericoid, with pinnate or palmate venation or one-veined. Glandular hairs with ethereal oils are often present, and some genera have extrafloral nectaries on the lower side of the lamina and petiole. Inflorescences are terminal or axillary panicles, racemes, spikes or heads often composed into thyrses, or the partial inflorescences in whorls, with bracts that are often large and sometimes petaloid (coloured). Flowers are bisexual and almost always zygomorphic and bilabiate. The (four or) five (to nine) sepals are often unequal in size, fused at the base and usually persistent. The (four or) five (to 16) petals are fused into a bilabiate corolla, usually with two upper and three lower petals or with five lower petals sometimes forming one lip. Usually four stamens are present, two longer (didynamous), sometimes with two, rarely five, six or up to 16 stamens, alternating with the petals and fused to the corolla tube. Anthers are basifixed or dorsifixed, often versatile and connivent, usually opening outwardly by lengthwise slits, rarely by apical pores. The connective is often prolonged, and in Salvia it is modified into a lever-like structure. Staminodia are

Phylogeny and evolution: Lamiaceae are the sister group to a clade uniting Orobanchaceae, Paulowniaceae, Phrymaceae and Mazaceae and are not closely related to Verbenaceae as previously assumed. However, several genera previously placed in Verbenaceae belong here, in particular Callicarpa, which was found as sister to the rest of the family. Therefore, the traditional subfamilial classification of Lamiaceae does not hold: Garrettia and relatives creating problems for recognising the separate subfamilies,

which below we have therefore subsumed into an enlarged Lamioideae (including Ajugoideae, Nepethoideae, Scutellarioideae, Lamioideae and the Garrettia group). This broadly defined Lamioideae are in turn sister to the Tectona clade (which should be elevated to a subfamily). These clades together are sister to Viticoideae (including Symphorematoideae) and Prostantheroideae (the former Chloanthaceae). With the inclusion of Viticoideae, Caryopteris, Clerodendrum and Callicarpa, it is clear that the traditional characters defining this family, the gynobasic style and schizocarpic fruit with four nut-like mericarps, have evolved several times (including in the distantly related Boraginaceae, Boraginales) and is not uniformly present within this circumscription of Lamiaceae. Generic delimitation is not clear in some cases and needs realignment, especially for genera such as Salvia (in which Mentha, Origanum, Rosmarinus and Thymus are embedded), Clerodendrum, Clinopodium, Leucas, Plectranthus, Stachys etc., all not monophyletic in the traditional sense. Trungboa poilanei was only collected once and originally described as Scrophulariaceae. This probably extinct species is placed here tentatively. Fossil leaves, flowers and fruits have been

Melittis melissophyllum, private garden, Hengelo, the Netherlands, [418]

Nepeta subsessilis, Ruissalo Botanical Garden, Turku, Finland [418]

Ocimum basilicum, private garden, Kingston upon Thames, Surrey, UK [418]

usually absent, and the nectar disk is entire or lobed and often developed on the lower side only. The superior ovary is composed of two (to five) carpels that each form a locule, in many species the locules parting in two upon maturation through ingrowth of the ovary wall (and then appearing tetraloculate). The ovary is sometimes stalked, and placentation is usually basal or axile, rarely apical. The style is single, usually bifid at the tip with a punctate stigma, usually gynobasic, in the middle of the four (false) locules or on the top of an unlobed ovary. Fruits are usually schizocarps with four single- (or two-)seeded nutlets (or small drupes in Prasium), sometimes a drupe (Viticoideae) or capsules. Distribution: Lamiaceae are cosmopolitan.

Plants of the World

575

LAMIALES

EUDICOTS

Origanum vulgare, Box Hill, UK [418]

Rosmarinus officinalis, private allotment, Kingston upon Thames, Surrey, UK [418]

Physostegia virginiana, Helsinki Botanical Garden, Finland [418]

Salvia pratensis, Helsinki Botanical Garden, Finland [418]

reported from the Eocene and later deposits in North America, Europe and India. Fossil pollen of Lamiaceae was found in Neogene layers. The age estimated for the first divergences in Lamiaceae is 38–49 million years ago. Genera and species: Lamiaceae include 241 genera with more than 6,800 species: ‘Callicarpoideae’ Callicarpa (c. 140); Prostantheroideae (14 genera) – Brachysola (2), Chloanthes (4), Cyanostegia (5), Dicrastylis (c. 25), Hemiandra (14), Hemigenia (c. 50), Hemiphora (1), Lachnostachys (6), Microcorys (c. 20), Newcastelia (9), Physopsis (5), Pityrodia (c. 40), Prostanthera (>100) 576

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Phlomis rotata, Yunnan, China [418]

Rostrinucula dependens, Royal Botanic Gardens, Kew, UK [418]

and Westringia (30); Viticoideae (7 genera) – Congea (7), Petitia (2), Pseudocarpidium (8), Sphenodesme (14), Symphorema (3), Teijsmanniodendron (23) and Vitex (c. 250); ‘Tectonoideae’ (7 genera) – Cornutia (12), Gmelina (c. 40), Paravitex (1), Premna (c. 50), Tectona (4), Tsoongia (1) and Viticipremna (5); Lamioideae (211 genera) – Acanthomintha (4), Acanthoprasium (2), Achyrospermum (c. 25), Acrotome (8), Acrymia (1, endangered), Aegiphila (c. 110), Aeollanthus (c. 40), Agastache (22), Ajuga (c. 45), Ajugoides (1), Alvesia (3), Amasonia (8), Amethystea (1), Anisochilus (c. 20), Anisomeles (3), Asterohyptis (4), Ballota (c. 30), Benguellia (1), Betonica (6), Blephilia (3), Brazoria (3),

Mentha spicata, Helsinki Botanical Garden, Finland [418]

Clerodendrum bungei, Royal Botanic Gardens, Kew, UK [418]

Bystropogon (7), Cantinoa (23), Capitanopsis (3), Caryopteris (7), Catoferia (4), Cedronella (1), Chaiturus (1), Chamaesphacos (1), Chelonopsis (16), Cleonia (1), Clerodendrum (c. 300), Clinopodium (c. 165), Colebrookea (1), Collinsonia (4), Colquhounia (c. 6), Comanthosphace (5), Condea (26), Conradina (6), Craniotome (1), Cuminia (2), Cunila (c. 15), Cyanocephalus (25), Cyclotrichium (6), Cymaria (3), Dauphinea (1), Dicerandra (9), Discretitheca (1), Dorystoechas (1), Dracocephalum (c. 70), Drepanocaryum (1), Elsholtzia (c. 40), Endostemon (19), Eplingiella (2), Eriope (>30), Eriophyton (1), Eriopidion (1), Eriothymus (1, extinct), Eurysolen (1), Faradaya (c. 5), Fuerstia (8), Galeopsis (10),

LAMIALES

EUDICOTS

Garrettia (1), Glechoma (6), Glechon (7), Glossocarya (c. 11), Gomphostemma (c. 35), Gontscharovia (1), Gymneia (7), Hanceola (8), Haplostachys (6), Haumaniastrum (c. 35), Hedeoma (c. 40), Hesperozygis (8), Hoehnea (4), Holmskioldia (1), Holocheila (1), Horminum (1), Hosea (1), Hymenocrater (c. 11), Hymenopyramis (7), Hypenia (c. 23), Hypogomphia (1), Hyptidendron (19), Hyptis (c. 145), Hyssopus (2), Isodon (c. 95), Isoleucas (2), Kalaharia (1), Karomia (9), Keiskea (c. 6), Killickia (4), Kurzamra (1), Lagochilus (c. 40), Lagopsis (4), Lallemantia (5), Lamium (c. 20), Lavandula (c. 40), Leocus (6), Leonotis (c. 40), Leonurus (c. 25), Lepechinia (c. 40), Leptohyptis (5), Leucas (c. 100), Leucosceptrum (1), Lophanthus (c. 20), Loxocalyx (3), Lycopus (14), Macbridea (2), Madlabium (1), Marmoritis (5), Marrubium (c. 40), Marsypianthes (6), Martianthus (4), Matsumurella (1), Medusantha (8), Meehania (6), Melissa (4), Melittis (1), Mentha (19), Meriandra (2), Mesosphaerum (c. 25), Metastachydium (1), Microtoena (c. 25), Minthostachys (c. 12), Moluccella (3), Monarda (16), Monardella (c. 20), Monochilus (2), Mosla (c. 22), Neoeplingia (1), Nepeta (>200), Obtegomeria (1), Ocimum (c. 65), Ombrocharis (1), Oncinocalyx (1), Oocephalus (14), Origanum (c. 35), Orthosiphon (c. 40), Otostegia (c. 10), Ovieda (1), Oxera (c. 20), Panzerina (6), Paralamium (1), Paraphlomis Caryopteris mongholica, Royal Botanic Gardens, Kew, UK [418]

(c. 20), Pentapleura (1), Perilla (1), Perillula (1), Peronema (1), Perovskia (7), Petraeovitex (8) Phlomidoschema (1), Phlomis (c. 75), Phlomoides (c. 175), Phyllostegia (c. 25), Physominthe (2), Physostegia (12), Piloblephis (1), Platostoma (c. 45), Plectranthus (c. 200), Pogogyne (7), Pogostemon (c. 85), Poliomintha (7), Prasium (1), Prunella (7), Pseudocaryopteris (3), Pseudomarrubium (1), Pycnanthemum (c. 20), Pycnostachys (37), Renschia (1), Rhabdocaulon (7), Rhaphiodon (1), Rhododon (2), Rosmarinus (3), Rostrinucula (2), Rotheca (c. 50), Roylea (1), Rubiteucris (2), Rydingia (4), Saccocalyx (1), Salvia (>900), Satureja (c. 40), Schizonepeta (3), Schnabelia (5), Scutellaria (c. 360), Sideritis (c. 140), Siphocranion (2), Spartothamnella (3), Stachydeoma (1), Stachyopsis (4), Stachys (c. 450), Stenogyne (c. 25), Suzukia (2), Synandra (1), Syncolostemon (c. 35), Tetraclea (2), Tetradenia (c. 17), Teucridium (1), Teucrium (c. 250), Thorncroftia (4), Thuspeinanta (2), Thymbra (4), Thymus (c. 220), Tinnea (19), Trichostema (c. 18), Tripora (1), Trungboa (1, possibly extinct), Volkameria (c. 30), Warnockia (1), Wenchengia (1, possibly extinct), Zataria (1), Zhumeria (1) and Ziziphora (c. 20). Uses: Lamiaceae are known for their various ethereal oils, and many species have fragrant Thymus pulegioides, Royal Botanic Gardens, Kew, UK [418]

leaves commonly used as kitchen herbs, in cooking, salads, drinks etc. The best known herbs are probably peppermint (Mentha ×piperita, a hybrid between M. aquatica and M. spicata), commonly used in drinks and its oil in cigarettes and confectionery (chewing gum, peppermint etc.); basil (Ocimum basilicum) used frequently in salads and as the basis for pesto; oregano (Origanum vulgare) frequently used in Italian cuisine, especially on pizza; rosemary (Rosmarinus officinalis) used in many meat dishes and baked goods and traded from southern to northern Europe since the 13th century; sage (Salvia officinalis), used in many dishes and cheese and to flavour drinks; summer savory (Satureja hortensis) traditionally used in broad beans (Vicia faba), but now a common ingredient of mixed herbs; and common thyme (Thymus vulgaris), used to flavour meat dishes with an extra benefit of working as a decongestant and relaxant. Less well known, but also delicious or frequently used herbs are: anise hyssop (Agastache foeniculum), basil thyme (Clinopodium acinos), yerba buena (C. douglasii), American dittany (Cunila origanoides), ground ivy or alehoof (Glechoma hederacea), hyssop (Hyssopus officinalis), lemon balm (Melissa officinalis), field mint (Mentha arvensis), watermint (M. aquatica), horse mint (M. longifolia), pennyroyal (M. pulegium), applemint (M. suaveolens), spearmint

Oxera neriifolia, New Caledonia [418]

Plants of the World

577

LAMIALES (M. spicata), orange mint (M. ×piperita ‘Citrata’), chocolate mint (M. ×piperita ‘Citrata Chocolate’), bergamot (Monarda didyma), hoary basil (Ocimum americanum), Greek basil (O. basilicum ‘Minimum’), Peruvian basil (O. campechianum), lemon basil (O. ×citriodorum), African basil (O. gratissimum), Thai basil (O. tenuiflorum), dittany (Origanum dictamnus), winter marjoram (O. heracleoticum), sweet marjoram (O. majorana), shiso (Perilla frutescens), Spanish sage (Plectranthus amboinicus), frosted mint (Poliomintha incana), shrubsage (Salvia fruticosa), winter savory (Satureja montana), mountain tea (Sideritis), lemon thyme (Thymus ×citriodorus), caraway thyme (T. herba-barona), broadleaved thyme (T. pulegioides), creeping thyme (T. serpyllum), tomillo salsero (T. zygis), kakoti (Ziziphora tenuior) etc. The scented oils produced by many species are popular as perfumes in the cosmetics and perfume industry. The best known and grown on a large scale in England and southern France are lavandin (Lavandula ×intermedia) and to a lesser extent, but with finer oil, true lavender (L. angustifolia). Lavender calms the body and was traditionally used in Roman baths, hence its name (from lavare, Latin, to bathe). Ninde (Aeollanthus myrianthus) and bergamot (Monarda) have seeds that yield an oil that is high in geraniol. Another important fragrant oil is patchouli, from Pogostemon cablin, mostly cultivated for this purpose in Sumatra. Chinese patchouli (Microtoena patchouli) yields an oil similar to patchouli. Rosemary oil is often used in perfumes such as ‘Eau de Cologne’. Clary (Salvia sclarea) yields an aromatic oil used in soaps and scents. Mint oil (Mentha) and tipo (Minthostachys mollis) yield mintoil with various applications in cosmetics and perfumes, as well as in teas and food flavourings. Serpolet oil is harvested from hoary thyme (Thymus polytrichus), which is used medicinally. White horehound (Marrubium vulgare) is a herb used for sore throats and in drinks, sweets and liqueurs. Apart from ethereal oils, some species are cultivated for their healthy cooking oils, especially chia (Salvia hispanica), which

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EUDICOTS

is high in alpha-linolenic acid and was a major crop of the Aztecs (Mexico). Other oil seeds are beni seed or black sesame (Hyptis spicigera), dragon’s head (Lallemantia iberica) and Pseudocaryopteris bicolor. Oil from yegoma (Perilla frutescens) is used in Japan to waterproof paper and umbrellas and in ink and paint. Seeds and leaves of Salvia apiana are also locally eaten. There are many medicinal plants in this family, too numerous to list here, but we highlight some. Fruits of chaste tree (Vitex agnus-castus) have been used as a pepper substitute, and were long believed to be an anaphrodisiac. For that reason, these trees were commonly planted in monasteries and used by monks to maintain celibacy, resulting in the other common name ‘monk’s pepper’. Medicinally, the fruit contains compounds like progesterone and are therefore sometimes recommended to treat premenstrual stress syndrome and cyclical breast pain. The functioning of the herb is not fully understood, but because it has been used since ancient times to treat sexual disorders, it is now being tested for gynaecological and prostate problems. Ajuga iva from the Mediterranean may have antimalarial properties, but this needs investigation. Gomphostemma javanicum and Hyptis suaveolens are said to have antitumour properties. Salvinorin A is a hallucinogen found in divine sage (Salvia divinorum), used in Mazatec rituals in Mexico, but it has now become a recreational drug in North America and Europe. Cat thyme (Teucrium marum), catnip (Nepeta cataria) and some other Nepeta species are irresistable to cats. Catnip can also have a mild hallucinogenic effect on humans. Hausa potato (Plectranthus esculentus and P. rotundifolius) has edible tubers. Pogostemon mutamba also has starchy tubers that can be eaten. Crosnes or Chinese artichoke (Stachys affinis) makes tubers that are edible when salted and boiled, whereas the tubers of betony (S. officinalis) are good against haemorrhoids. Fruits of Cornutia pyramidata are used to make a blue or red ink. The best-known timber species of this family is is teak (Tectona grandis),

commercially grown throughout the tropics for its valuable, light wood, which is used for boats, bridges, furniture, f looring, construction etc. Gmelina species also produce a teak-like timber but greyer in colour. Fiddlewood, ‘bois fidèle’, is harvested from Petitia and Vitex and makes good timber for poles. Headache tree (Premna serratifolia) has beautifully marked woods used in Indonesia for furniture. Many genera include popular garden ornamentals, often because of their long flowering period, attractiveness to bees, butterf lies and hummingbirds, and their often scented leaves. Genera that are frequently cultivated are Agastache (giant hyssop), Ajuga (bugle), Amethystea, Ballota (horehound), Blephilia (pagoda plant), Callicarpa (beautyberry), Caryopteris (bluebeard), Clerodendrum (glorybower), Colquhounia, Congea (orchid shower), Conradina (false rosemary), Dracocephalum (dragonhead), Elsholtzia, Glechoma (ground ivy), Holmskioldia (cup-and-saucer plant), Horminum (dragonmouth), Lamium (dead nettles), Lavandula (lavender), Leonotis (wild dagga), Leonurus (motherwort), Lepechinia, Meehania (Meehan’s mint), Melittis (bastard balm), Moluccella (bells of Ireland), Monarda (beebalm), Monardella (coyote mint), Nepeta (catmint), Origanum (oregano), Orthosiphon, Perilla, Perovskia (Russian sage), Phlomis (Jerusalem sage), Physostegia (obedient plant), Plectranthus (coleus), Premna, Prostanthera (mintbush), Prunella (selfheal), Pycnostachys (blueboys), Rotheca (Oxford and Cambridge bush), Salvia (ornamental sage), Scutellaria (skullcap), Sideritis (ironwort), Stachys (betony, lamb’s ears), Tetradenia, Teucrium (germander), Thymus (ornamental thyme), Trichostema (blue curls), Vitex (chaste tree) and many others. Species of Clerodendrum and Plectranthus are frequently grown as houseplants. Etymology: Lamium is the Latinised Greek of λαμία (lamia), a gaping mouth, in reference to the lipped flowers. Labiatae (derived from Latin; labium, lip) is an alternative name for this family, but it is not based on a genus name and is therefore better avoided.

LAMIALES

EUDICOTS

Lancea tibetica, Yunnan, China [419]

Mazus stachydifolius, Mt Huangshan, Anhui, China [419]

Mazus pumilus, Shanghai, China [419]

419. MAZACEAE

carpels forming a single locule. Placentation is usually more or less basal or axile. A single style terminates the ovary and has a bilobed or fan-shaped, sensitive stigma. Fruits are berry-like indehiscent capsules enclosed by the persistent and sometimes enlarged calyx.

Etymology: Mazus is the Latinised form of Greek μαστός (mastos or mazos), breasts, from the ridges on the lower lip of the flower.

Cupflower family

420. PHRYMACEAE Lopseed family

Distribution: These plants are found in Tibet, temperate East Asia, mountains in Malesia to New Guinea, southeastern Australia, Tasmania and New Zealand. Mazaceae are perennial and annual, rhizomatous herbs with stems square in cross-section. Leaves are usually opposite (rarely alternate and spiral), simple or lobed, without stipules. Leaf blades have serrate margins and pinnate venation. Inflorescences are terminal or axillary cymes, or flowers are borne singly in the leaf axils. Flowers are bisexual and zygomorphic. The five sepals are persistent and fused into a bell-shaped or bilabiate (three upper and two lower lobes) calyx. The (two to) four or five petals are fused into a tubular or bilabiate (with two upper and three lower lobes) corolla, sometimes with a spur. Nectaries are present at the base of the four (rarely two) stamens, of which two are longer (didynamous). Filaments are free from each other, alternate with the petals, and are fused to the floral tube. Anthers are dorsifixed and open by longitudinal slits. The superior ovary is composed of two fused

Phylogeny and evolution: Even though in earlier studies Mazaceae were found to be close to Mimulus and hence were placed in the recently expanded Phrymaceae, they are now known to be sister to a larger clade including Orobanchaceae, Paulowniaceae and Phrymaceae. Therefore this family is accepted here, although they are not morphologically divergent from many other families of Lamiales. Mazaceae differ chemically from Phrymaceae so there is character support for separation, although not immediately evident. Genera and species: Mazaceae include three genera and c. 23 species: Dodartia (1), Lancea (2) and Mazus (c. 20). Uses: Mazus is sometimes grown as a groundcover ornamental, and M. reptans is especially popular. Depgul (Lancea thibetica) is smoked as a narcotic in Kashmir.

These are annual and perennial rhizomatous herbs, rarely slightly woody at the base or becoming shrubs, mostly terrestrial, sometimes aquatic. Young stems are square in cross-section and bear opposite or alternate, simple leaves without stipules. Leaf blades have pinnate venation and serrate, sometimes lobed, margins. Inflorescences are terminal or axillary cymes, racemes or spikes, rarely in heads of flowers or flowers solitary in leaf axils. Flowers are usually zygomorphic, rarely almost actinomorphic. The (three to) five sepals are fused into a tubular or bilabiate, persistent calyx (with three upper and two lower lobes). The (two to) four or five petals are fused into a tubular or bilabiate corolla (with two upper and three lower lobes,

Plants of the World

579

LAMIALES

EUDICOTS

Mimulus douglasii has two upper lobes only), sometimes with a basal spur. Stamens are usually four, rarely two, usually two longer (didynamous). Filaments are fused to the corolla tube and alternate with the petal lobes. Anthers are dorsifixed and often confluent, and the thecae open by lengthwise slits. The superior ovary is composed of two carpels, with the lower carpel often reduced and sterile, usually unilocular and with axile placentation. A single simple style with a broadly bilobate (sometimes capitate) stigma tops the ovary. The stigma is sensitive in Mimulus. Fruits are usually loculicidal (rarely septicidal) capsules or a one-seeded nut, enclosed by a persistent calyx, with the calyx teeth often modified into prickles or hooks, rarely a berry. Distribution: Phrymaceae are found in North America, Mesoamerica, the Antilles, Erythranthe cardinalis, Royal Botanic Gardens, Kew, UK [420]

Diplacus pictus, Royal Horticultural Society Garden, Wisley, UK [420]

580

Christenhusz, Fay & Chase

western South America, the Galápagos and Juan Fernández Islands, southern and eastern Africa, Madagascar, temperate East Asia, tropical Asia, New Guinea, the Philippines, Australia and New Zealand, with few species in the humid tropics. Phylogeny and evolution: This unexpected grouping of genera is made up of Phryma, formerly placed in Verbenaceae, and Mimulus and their relatives, which used to be considered members of Scrophulariaceae s.l. Phryma may be embedded in Mimulus, resulting in the disintegration of the latter into ten genera, some of which were already in use. Mazaceae, which share many characters with this family, were previously included. This distinction appears to be supported by chemical differences: Mazus has iridoids whereas Mimulus does not. Phrymaceae are

Hemichaena fruticosa, Guatemala (MV) [420]

sister to the clade of Paulowniaceae plus Orobanchaceae. Genera and species: Phrymaceae include 13 genera and 187 species: Diplacus (46), Elacholoma (2), Erythranthe (111), Glossostigma (5), Hemichaena (5), Leucocarpus (1), Microcarpaea (1) Mimetanthe (1), Mimulus (7), Peplidium (4), Phryma (1), Thyridia (1) and Uvedalia (2). Uses: Some species of Leucocarpus and Mimulus are grown as ornamentals. As a result, Mimulus (especially M. guttatus and M. luteus) is widely naturalised outside its native range. Etymology: Phryma is a name of unknown origin, but may be a cognate with Greek φρύνος (phrynos), a toad. Phryma leptostachya, Saugatuck, Michigan, USA [420]

Peplidium aithocheilum, Western Australia [420]

LAMIALES

EUDICOTS

Paulownia kawakamii in fruit, Royal Botanic Gardens, Kew, UK [421]

Paulownia kawakamii, Royal Botanic Gardens, Kew, UK [421]

Paulownia tomentosa, Royal Botanic Gardens, Kew, UK [421]

421. PAULOWNIACEAE

Filaments are fused to the corolla tube, and anthers are basifixed, positioned head-to-head and opening by lengthwise slits. A disk with vascularised nectaries surrounds the ovary. The superior ovary is composed of two fused carpels, each forming a locule, topped by a single, simple, hollow style and a punctate stigma. Placentation is axile and protruding, with numerous ovules per carpel. Fruits are loculicidal (Paulownia) or loculicidalsepticidal (Wightia) capsules with a persistent, somewhat woody calyx tube and numerous flattened, winged seeds.

characters with Paulownia and is thus placed here tentatively. It shares the membranaceous seed wings typical for Paulownia (but these are also found in Brandisia, Orobanchaceae). The placement of this genus could further destabilise the family circumscription of Lamiales.

Empress-tree family

These are deciduous and evergreen trees and viny shrubs (Wightia). Leaves are petiolate, opposite (whorled in young shoots), simple, without stipules. Blades are entire or serrate and have pinnate venation, often large and basally cordate and often with with several terminal lobes. Extrafloral nectaries are often present on the leaf blades. Inflorescences are terminal thyrses (Paulownia) or axillary racemes (Wightia). Flowers are bisexual and zygomorphic. The five sepals are more or less basally fused, bell-shaped and covered by long hairs. The five petals are fused into a bilabiate corolla with nectar guides on the lip. Four stamens are present with two longer (didynamous); in terminal flowers, stamens are often five, alternating with the petal lobes.

Genera and species: The three genera of Paulowniaceae include eight species: Paulownia (6), Shiuyinghua silvestrii and Wightia speciosissima.

Distribution: This family occurs from temperate East Asia (Japan, Korea, China, Taiwan), extending into warm-temperate to tropical regions in the Himalayas, northeastern India, northern Burma, Southeast Asia and Malesia.

Uses: Paulownia produces reasonable timber and is sometimes cultivated for that purpose. Best known in horticulture is the empress tree (Paulownia tomentosa), which is commonly grown in gardens and city parks and which easily naturalises in North America and Europe.

Phylogeny and evolution: Fossils assigned to Paulownia have been described from the Miocene/Pliocene transition zone of Germany, but this identification may be problematic. Paulowniaceae are sister to Orobanchaceae. However, some studies have placed them as sister to Lamiaceae. The position of Wightia is still uncertain, but it shares

Etymology: Paulownia is named for Her Imperial Highness Anna Pavlovna of Russia (1795–1865), sponsor of the second expedition to Japan (1861) by Philipp von Siebold, who named this tree for her and introduced it into European horticulture. She was queen consort, wife of King William II of the Netherlands. Plants of the World

581

LAMIALES

EUDICOTS

Castilleja wrightii, Sea Ranch, California, USA [422]

Neobartsia laticrenata, Ecuador [422]

Broomrape family

These are mostly parasitic herbs, rarely woody herbs and shrubs, that are green or lack chlorophyll, and most turn black upon drying. They are rarely non-parasitic (Lindenbergia, Rehmannia and Triaenophora), but in the other species haustorial connections are made to the roots of their host plants. In photosynthetic species, plants only obtain Christenhusz, Fay & Chase

[422]

Euphrasia alpina, French Pyrenees [422]

422. OROBANCHACEAE

582

Triphysaria eriantha subsp. rosea, Sea Ranch, California, USA

water and nutrients from the host, whereas holoparasitic species also obtain organic compounds from the host phloem. Leaves are normally developed and green or scale-like and not green, opposite to alternate and then spirally arranged, without stipules. Blades are usually toothed to deeply lobed, often pinnatifid or bipinnatifid, sometimes entire, and venation is pinnate. Inflorescences are racemes or spikes, often with colourful bracts, each subtending a flower. The bisexual flowers are zygomorphic and often bilabiate, nearly actinomorphic in Buchnera. The two to five sepals are fused into a four- to five-lobed or toothed calyx. The usually five petals are fused into a salverform or campanulate, often galeate or bilabiate corolla. The four (or five) stamens are usually of two different lengths (didynamous), sometimes nearly equal, sometimes sterile (Lamourouxia). Anthers

Parentucellia trixago, Sicily, Italy [422]

open by lengthwise slits or pores (Euphrasia). The superior ovary is composed of two (rarely three) fused carpels topped with a single style and capitate or bilobed stigma. The fruit is a loculicidal to septicidal capsule. Distribution: Members of this family are found almost globally, apart from permanently frozen areas. Their major centres of diversity are in the Mediterranean, southern Africa, the Himalayas and western North America. Phylogeny and evolution: Traditionally, Orobanchaceae were a small family containing only parasitic genera without chlorophyll, including Cistanche, Conopholis, Epifagus, Hyobanche and Orobanche. Photosynthetic parasites of Lamiales were placed in Scrophulariaceae, but these were moved to Orobanchaceae after molecular

LAMIALES

EUDICOTS

results showed an exclusive relationship with them. The taxonomic problems of Orobanchaceae and Scrophulariaceae are only one aspect of the difficulties that family delimitation in the order Lamiales has presented to taxonomists. Changes in family circumscriptions following dismemberment of Scrophulariaceae have seen an increase in the size of Orobanchaceae with the inclusion of green parasitic genera, such as Castilleja, Euphrasia, Pedicularis and Rhinanthus. Rehmannia, Triaenophora and Lindenbergia are not parasitic but were found to be sister to the rest of the family. Brandisia is facultatively parasitic and may be sister to the core clade of photosynthetic Orobanchaceae. Brandisia has a seed coat with a reticulate pattern and membranous seed wings similar

Genera and species: Orobanchaceae include c. 104 genera and more than 1,960 species: Aeginetia (3), Agalinis (c. 40), Alectra (c. 40), Asepalum (1), Aureolaria (11), Bardotia (1), Bartsia (c. 13), Bartsiella

(1), Baumia (1), Bornmuellerantha (1), Boschniakia (1), Brachystigma (1), Brandisia (c. 13), Buchnera (c. 100), Bungea (2), Buttonia (3), Castilleja (>200), Centranthera (9), Chloropyron (4), Christisonia (17), Cistanche (16), Conopholis (3), Cordylanthus (18), Cyclocheilon (3), Cycnium (18), Cymbaria (4), Dasistoma (1), Dicranostegia (2), Epifagus (1), Eremitilla (1), Escobedia (c. 15), Esterhazya (5), Euphrasia (c. 170, plus many apomictic species), Gerardiina (2), Ghikaea (1), Gleadovia (6), Graderia (5), Harveya (c. 40), Hedbergia (1), Hiernia (1), Hyobanche (8), Kopsiopsis (2), Lamourouxia (28), Lathraea (7), Leptorhabdos (1), Leucosalpa (4), Lindenbergia (15), Macranthera (1), Macrosyringion (2), Magdalenaea (1), Mannagettaea (3), Melampyrum (35),

Melampyrum behariense, Romania [422]

Pedicularis densiflora, Sea Ranch, California, USA [422]

Orobanche caryophyllacea, Sicily, Italy [422]

to Paulownia (Paulowniaceae), which could be a reason for uniting the two families. Also Phrymaceae and Mazaceae could be included as they are closely related clades and share many morphological characters. Holoparasitism, which involves loss of chlorophyll, has occured several times in the green parasites in this clade. Monophyly of several genera has yet to be evaluated. Orobanche is sometimes divided into several genera. Castilleja is here treated in its broad sense.

Lamourouxia virgata, Napo, Ecuador (CD) [422]

Lathraea clandestina, Royal Botanic Gardens, Kew, UK [422]

Conopholis alpina var. mexicana, New Mexico, USA (DZ) [422]

Plants of the World

583

LAMIALES Melasma (35), Micrargeria (5), Micrargeriella (1), Monochasma (4), Necranthus (1), Neobartsia (47), Nesogenes (8), Nothobartsia (3), Nothochilus (1), Odontitella (1), Odontites (c. 30), Omphalotrix (1), Orobanche (c. 150), Orthocarpus (9), Paraharveya (1), Parasopubia (2), Parentucellia (2), Pedicularis (c. 550), Petitmenginia (2), Phacellanthus (1), Phelypaea (4), Phtheirospermum (7), Physocalyx (2), Platypholis (1), Pseudomelasma (1), Pseudosopubia (5), Pseudostriga (1), Pterygiella (4), Radamaea (5), Rehmannia (6), Rhamphicarpa (6), Rhaphispermum (1), Rhinanthus (c. 45), Rhynchocorys (6), Schwalbea (1), Seymeria (c. 25), Seymeriopsis (1), Sieversandreas (1), Silviella (2), Siphonostegia (4), Sopubia (41), Striga (c. 40), Tetraspidium (1), Thunbergianthus (2), Tomanthera (2), Tozzia (1), Triaenophora (3), Triphysaria (5), Vellosiella (3), Xylanche (1) and Xylocalyx (5).

584

EUDICOTS

Castilleja densiflora, Lathraea clandestina, Lindenbergia, Melampyrum nemorosum, Pedicularis and Rehmannia elata (Chinese foxglove). Several species are serious agricultural pests, especially Aeginetia indica (on sugarcane), Orobanche crenata (on beans), O. cumana (on sunflowers), O. ramosa (on hemp), Seymeria cassioides (on pines) and Striga (on C 4 cereals such as maize, millet, sorghum and sugarcane). Rhinanthus species are used in grassland restoration to reduce the competitiveness of dominant grasses.

Uses: Cistanche phelypaea is eaten like asparagus by the Tuareg in North Africa. Inflorescences of Orobanche and Pedicularis have been eaten by native Americans. Eyebright (Euphrasia officinalis) was once used to treat eye diseases. Some species are occasionally grown as ornamentals, although their parasitic relationships may cause problems for cultivating them without a host. Examples of plants cultivated in gardens include Agalinis purpurea, Aureolaria flava,

Parasitic sap-suckers: The vast majority of Orobanchaceae are root-parasites, making connections with the roots of host plants that are often specific to the species of parasite. When seeds germinate, the roots quickly find host-plant roots and make haustorial connections. Some species such as Orobanche arenaria have motile seeds that search for the host after germinating by spiralling into the soil, or seeds that germinate in response to host root exudates, remaining dormant in the soil for many years when no host is present. This can result in an agricultural field being rendered useless for certain crops due to the presence of a seed bank of Orobanchaceae in the soil. Partial parasitism is also common; here the parasite does have chlorophyll and can survive without a host. However, these so-called

Rehmannia elata, Royal Horticultural Society Garden, Wisley, UK [422]

Striga elegans, Trappes Valley, Eastern Cape, South Africa (CD) [422]

Christenhusz, Fay & Chase

‘hemiparasties’ will grow much better when a host connection is achieved. Parasitism has evolved just once in Orobanchaceae, but the total loss of photosynthetic ability has evolved at least five times within the family. It has been proposed that although the evolution of a haustorial connection with a host plant is a relatively rare event, the transition from a green, photosynthetic parasite to an achlorophyllous, obligate holoparasite may occur more readily through mutations leading to the loss of function of one or more genes involved in photosynthesis. Thus, genera that are radically different morphologically, such as holoparasitic, scaly Lathraea and green, leafy Rhinanthus are more closely related to each other than Lathraea is to other holoparasites such as Orobanche. Are these plants carnivores as well as parasites? Various authors, especially in the 19th century, have suggested that the glands and capitate hairs found in the cavities of the scale leaves of some Orobanchaceae, such as Lathraea, might be involved in the entrapment and digestion or absorption of small organisms. This has been discounted because these structures are involved instead in water excretion. Etymology: Orobanche is composed of the ancient Greek words ωρωvός (orobos), bitter vetch, and anche (άγχη), stress or anxiety. Lindenbergia grandiflora by M. Smith from Curtis’s Botanical Magazine, vol. 126: plate 7738 (1900) [422]

AQUIFOLIALES

EUDICOTS

AQUIFOLIALES Families 423 to 427 make up the order Aquifoliales, which can be recognised by their valvate free petals and one or two ovules per carpel. They have been estimated to have evolved c. 87–88 to 113 million years ago. Within this order, there are two main clades, one formed by Cardiopteridaceae and Stemonuraceae, most members of which were formerly placed in Icacinaceae. The other is composed of Aquifoliaceae, Helwingiaceae (ex-Cornaceae) and Phyllonomaceae (ex-Saxifragaceae). This is a good example of an order that includes members that were placed far apart based on an emphasis on selected morphological characters but found to be closely related on the basis of DNA studies. After more careful morphological study, similarities were found among these families. Epiphylly, where flowers emerge from the leaf blade has evolved three times independently in this order (Helwingia, Leptaulus and Phyllonoma), but in each case, the inflorescences are formed in different manners, so it is not parallel evolution in the strict sense.

Grisollea thomassetii, Seychelles [423]

Gomphandra quadrifida, Singapore (MN) [423]

423. STEMONURACEAE Buff-beech family

These are evergreen, unisexual shrubs and trees with entire, alternate, distichous leaves without stipules. Blades have entire margins and obscure pinnate venation. Inflorescences are axillary or cauliflorous, compound cymes with actinomorphic, unisexual flowers. The usually four or five (to seven) sepals are more or less basally fused, and the same number of petals are usually free (rarely slightly fused at

the base) and sometimes keeled on the outside and inflexed at the apex. Male flowers usually have five stamens alternating with the petals. Filaments are stout with club-shaped hairs and a well-developed connective, and anthers are basifixed, opening by lateral slits. Male flowers have a pistillode in the centre. Female flowers often have five staminodes (or absent), and the superior ovary is composed of two carpels of which only one is fertile, together forming a single locule. The ovary is topped by a broad sessile stigma. Fruits are drupes that are often lobed with a fleshy appendage on one side. Distribution: This is a nearly pantropical family found in northern South America, tropical West Africa, Madagascar, the Comores and Seychelles, southern India, Sri Lanka and tropical Southeast Asia and Malesia south to northern Australia.

Lasianthera africana, Monts de Cristal, Gabon (CD) [423]

Phylogeny and evolution: Previously the members of this family were included in Icacinaceae, but wood anatomy and molecular analysis confirmed their exclusion from that family and a relationship with Aquifoliales, in which they are probably sister to Cardiopteridaceae, another family for which most genera were formerly part of Icacinaceae. The family can be recognised by their distinctive cigar-shaped buds, a flattened drupe that often has a colourful appendage or lobe, club-shaped hairs on their filaments and distichous, entire leaves. Genera and species: Stemonuraceae include 12 genera and c. 103 species: Cantleya (1), Codiocarpus (2), Discophora (3), Gastrolepis (2), Gomphandra (65), Grisollea (3), Hartleya (1), Irvingbaileya (1), Lasianthera (2), Medusanthera (8), Stemonurus (c. 14) and Whitmorea (1). Plants of the World

585

AQUIFOLIALES

EUDICOTS

Uses: Some species make locally valuable timber. The wood of Cantleya corniculata is fragrant and sometimes used as a sandalwood replacement. Fruits of C. corniculata are edible. Etymology: Stemonurus is composed of the Greek words στήμονα (stemona), a stamen or thread, and ουρά (oura), a tail.

424. CARDIOPTERIDACEAE Churnwood family

axillary or rarely epiphyllous (Leptaulus) panicles or cymes. Flowers are bisexual and actinomorphic. The five sepals are fused, and the five petals are free or basally fused. The five stamens alternate with the petals and are sometimes fused to the petal tube. They have basifixed or dorsifixed anthers that open by lateral slits. The superior ovary is composed of three carpels, and there is a false locule in Citronella and Pseudobotrys. The ovary is topped by one or two slender styles with truncate or capitate stigmas. Fruits are drupes that are often lepidotehairy outside. Distribution: The family has a nearly pantropical distribution but is rare in Central America, patchy in South America and absent from the Indian subcontinent. They extend into subtropical Chile, Brazil and Australia.

These are bisexual, evergreen trees and shrubs, occasionally twining vines with milky sap, all parts covered in scale-like, stellate or simple hairs. Simple leaves are alternate, arranged in a plane (distichous) or spirals (Cardiopteris). Blades have entire or serrate margins with small symbiotic arthropod-housing structures (domatia) and pinnate venation. Inflorescences are terminal, Gonocaryum macrophyllum in fruit, Singapore (WA) [424]

586

Christenhusz, Fay & Chase

Phylogeny and evolution: This heterogenous group of genera has been found to be separate from other Icacinaceae in molecular studies, and characters shared by the genera are the slender, short styles and the more or less f lattened and sometimes ridged drupes. Cardiopteris is a vine that resembles Dioscorea (Dioscoreaceae) in general habit and leaf shape, but the flowers and fruits of Cardiopteris are five- not three-parted.

Citronella mucronata, Santiago Botanical Garden, Chile [424]

Genera and species: In their current circumscription, Cardiopteridaceae include five genera and 43 species: Cardiopteris (2), Citronella (22), Gonocaryum (11), Leptaulus (6) and Pseudobotrys (2). Uses: The leaves of Cardiopteris are locally eaten as a vegetable. Congonha (Citronella gongonha) is used to make a kind of ‘yerba maté’ (a tea, see Aquifoliaceae). Churnwood (Citronella moorei) from Australia produces valuable timber. Etymology: Cardiopteris is composed of the Greek words καρδιά (kardia), a heart, and πτέρης (pteris), a feather or fern.

425. PHYLLONOMACEAE Flowering-leaf family

These are glabrous trees and shrubs with simple, alternate, petiolate leaves and minute

Cardiopteris moluccana, Barron Gorge National Park, Queensland, Australia (CD) [424]

AQUIFOLIALES

EUDICOTS

Phyllonoma laticuspis, Chiapas, Mexico (CD) [425]

Phyllonoma laticuspis, Chiapas, Mexico (CD) [425]

Helwingia chinensis, Hortus Botanicus Leiden, the Netherlands [426]

Helwingia japonica, Royal Botanic Gardens, Kew, UK [426]

placentation, stipulate, simple leaves and similarity in seed structure. Their inflorescences have similarities to those of Helwingia (Helwingiaceae), and molecular studies indicate placement in this order close to Helwingiaceae and Aquifoliaceae.

Distribution: This is a Neotropical family found from Durango State in Mexico and Mesoamerica to the Andes of northwestern Bolivia.

shrubs, rarely small trees with simple, alternate, petiolate leaves. The stipules are entire or divided and soon deciduous. Blades have pinnate venation and glandular serrate or crenate margins. Inflorescences are sessile umbels, borne on the midvein of a leaf blade (epiphyllous), rarely on the petioles of leaves or on the upper part of young branches. Flowers are unisexual, green or purplish and actinomorphic. Sepals are entirely fused to form a toothed cup, but sometimes the flowers are interpreted as sepal-less. The three or four (or five) petals are free and spreading. The nectar disk is flat and fleshy. Male flowers are three to 20 per umbel, and each has three or four (or five) free stamens alternating with the petals. Anthers are dorsifixed and open by longitudinal slits. Female flowers are one to four per umbel. The inferior ovary is composed of three or four (rarely five) carpels, each forming a locule and topped with a single style and lobed stigma. Fruits are drupe-like berries crowned by a persistent calyx and style, with one to five ridged seeds.

Phylogeny and evolution: This family was formerly included in Saxifragaceae or Grossulariaceae based on their parietal

Distribution: The family is distributed in temperate East Asia, from the Himalayas to Japan and northern Vietnam.

stipules. Blades have pinnate venation, with up to ten veins on each side below the inflorescence insertion and entire to broadly serrate-dentate margins. Inflorescences are epiphyllous cymes, with a single central flower and several side branches, appearing as a short raceme borne on the upper surface of the leaf blade, often towards the leaf apex. Flowers are bisexual and actinomorphic. The five sepals are free and open, and the five petals are also free and spreading. Stamens alternate with the petals and are inserted in the same whorl atop a short hypanthium. Anthers are basifixed and open by lateral slits. A prominent nectar disk surrounds the style. The inferior ovary is composed of two carpels fused to form a single locule. The ovary is topped with a style that is fused to the nectar disk and has two recurved stigmas sessile on the disk. Fruits are three- to six-seeded berries.

Genera and species: The four species of this family are members of the single genus Phyllonoma. Etymology: Phyllonoma is composed of the Greek words φύλλων ( fyllon), leaf, and νομός (nomos), pasture or county, in reference to the epiphyllous inflorescences.

426. HELWINGIACEAE Flowering-rafts family

These are unisexual, evergreen and deciduous

Plants of the World

587

AQUIFOLIALES Phylogeny and evolution: Morphologically, this family is intermediate between Cornaceae and Araliaceae, but molecular studies place them as sister to Phyllonomaceae, which also have epiphyllous inflorescences, in which an axillary shoot has fused to the midvein of the leaf.

EUDICOTS

427. AQUIFOLIACEAE Holly family

Genera and species: The sole genus in Helwingiaceae is Helwingia with four species. Uses: Helwingia japonica and rarely other species are sometimes grown as curiosity shrubs in specialist collections, but they are popular in Japan and China and should perhaps be grown more widely. The young leaves are edible. Etymology: Helwingia is named in honour of Prussian botanist Georg Andreas Helwing (1668–1748), who was a specialist in Pulsatilla (Ranunculaceae) and introduced several plants into horticulture. Ilex cornuta in fruit, Royal Botanic Gardens, Kew , UK [427]

This is a family of evergreen and deciduous, unisexual trees and shrubs. Leaves are simple, usually petiolate, alternate, rarely opposite, with minute stipules. Leaf blades have entire, serrate or spinose margins and pinnate venation. Inf lorescences are axillary cymes or fascicles, but sometimes flowers are solitary. Flowers are unisexual by having aborted stamens or ovaries. The four to eight sepals are fused and persistent in fruit. Petals are as many as sepals and fused

Ilex aquifolia, Box Hill, UK [427]

Ilex paraguariensis, Matthaei Botanical Garden, Ann Arbor, Michigan, USA [427]

Christenhusz, Fay & Chase

Distribution: This is a pantropical family, most diverse in the American and Asian tropics, with only one species in Africa and several temperate representatives in eastern North America, Western Europe, Macaronesia, Anatolia and northern Iran, the Himalayas and Asia north to Sachalin.

Ilex integra, Royal Botanic Gardens, Kew, UK [427]

Ilex vomitoria in fruit, San Antonio Botanical Garden, Texas, USA [427]

Ilex anomala, Tahiti, French Polynesia [427]

588

up to half their length, forming a cup. Male flowers have four to eight stamens that are all similar, free from each other but fused to the petal tube and alternating with the lobes. Anthers are dorsifixed and open inwardly by lengthwise slits. A rudimentary ovary is present and has a short beak. Female flowers have sagittate or cordate staminodes that are fused with and alternate with the petals. The superior ovary is composed of four to eight (to ten) carpels, each forming a locule and topped by a nearly sessile, capitate, discoid or columnar stigma. Fruits are red, brown or black globose berries with a persistent style and several hard seeds.

ASTERALES

EUDICOTS

Phylogeny and evolution: Phylogenetic relationships in Aquifoliaceae are strongly correlated with geography, and the family was much more widely spread in the past, having for instance become extinct in New Zealand during the Tertiary. The South American species were found to be sister to the rest, and the species from Hawaii and New Caledonia were placed inside the American clade, indicating that long-distance dispersal must also happen in this family from time to time. The distinctive genus Nemopanthus was found to be deeply embedded in Ilex and is now a synonym. Previously, Phelline and Sphenostemon have been included in Aquifoliaceae, but Phelline is in its own family in Asterales, whereas Sphenostemon

Genera and species: Aquifoliaceae include a single genus Ilex with c. 400(–600) species. Extensive hybridisation in some areas makes species delimitation difficult.

northwestern South America and China, respectively. Ilex species supply hard, white, finegrained wood that is used for carpentry and furniture and inlay work. Several species, hydrids and cultivars of holly are grown as garden ornamentals, and female holly branches with berries are often used as decoration, especially for Christmas.

Uses: Yerba maté or Paraguay tea, popular in Latin America, is brewed from dried young leaves of Ilex paraguariensis. Dahoon (I. cassine) is also used for tea in North America. Feverbark (I. verticillata), guayusa (I. guayusa) and kudingcha (I. kaushue) are used as an alternative to tea in North America,

Etymology: Aquifolium is the Latin name for holly, derived from acus, needle, and folia, leaf, referring to the sharply pointed leaves of some species. It is a later synonym of Ilex, the Latin name for holm oak (Quercus ilex, Fagaceae), and is now applied to hollies.

is now in Paracryphiaceae (Paracryphiales). Aquifoliaceae were previously placed in a broad Celastrales. They are most closely related to Helwingiaceae.

ASTERALES Families 428 to 438 make up the order Asterales, with an estimated crown age of c. 90 million years. Campanulaceae and Rousseaceae are sister to the rest, in which a clear trend in inflorescence condensation can be observed toward the typical ‘flower-like’ composed heads found in Asteraceae. The order has been hypothesised to have originated in the Southern Hemisphere, and it has been postulated that diversification occurred in some lineages once they reached the Northern Hemisphere. Asterales include c. 13.6% of eudicot diversity.

428. ROUSSEACEAE Putaweta family

This is a family of evergreen shrubs and trees, sometimes climbing (Roussea), with simple unicellular hairs. They have simple, alternate or opposite (Roussea), petiolate leaves without stipules. Leaf blades have (glandular) serrate margins and pinnate venation. Inf lorescences are terminal or axillary panicles with minute bracts, or f lowers are solitary, often nodding. The bisexual or functionally female flowers are

actinomorphic. There are usually four or five (sometimes to seven) free or basally fused, often reflexed sepals and the same number of free or basally fused petals. The four or five stamens alternate with the petals and are free from each other and the perianth. Anthers are dorsifixed and open by lengthwise slits. A nectar disk between the stamens is often present. The superior or inferior ovary is composed of five (to seven) carpels, forming three to seven locules. It is topped by a single style with a capitate, globose or lobed stigma. Fruits are angular berries or loculicidal capsules with numerous seeds. Distribution: This is a Souther n Hemisphere family, with Abrophyllum and Cuttsia resticted to rainforests in eastern Australia, Carpodetus with a disjunct distribution in New Guinea and New Zealand, and Roussea endemic to Mauritius.

Phylogeny and evolution: Traditionally, Carpodetus and relatives (Carpodetaceae) were placed in a polymorphic Saxifragaceae (or Escalloniaceae). A relationship with Ericaceae, based on pollen morphology, had also been suggested. Molecular studies have shown that Carpodetus and relatives are sister to Roussea, which was previously of unknown affinity. They are sister to Campanulaceae, with which they share carpel characters. Genera and species: Rousseaceae include four genera and six species: Abrophyllum (2), Carpodetus (2), Cuttsia viburnea and Roussea simplex. Uses: The wood of putaweta (Carpodetus serratus) is strong and used for tool handles, but it is attractive to wood-boring insects, reducing its value. Abrophyllum ornans and Carpodetus serratus are sometimes grown as garden ornamentals. Plants of the World

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Roussea simplex pollinated by a blue-tailed day gecko (Phelsuma cepediana) on Mauritius (DN) [428]

Abrophyllum ornans, Mt Lewis, Queensland, Australia (CD) [428]

Carpodetus serratus in fruit, Leith Saddle near Dunedin, New Zealand (JC) [428]

An endangered plant in the land of the dodo: Roussea simplex is endemic to Mauritius and has no close relatives. Even though it used to be common on this distant Indian Ocean island, there are now only 85 plants left in the wild, and the species is considered Critically Endangered. For a long time, the life history of Roussea was an enigma. It has recently been discovered that its flowers are only pollinated successfully by a rare, endemic, blue-tailed day gecko (Phelsuma cepediana). This beautiful reptile climbs

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Carpodetus serratus, Royal Botanic Gardens, Kew, UK [428]

Carpodetus serratus, Botanical Reserve, Nelson, New Zealand (JC) [428]

the plant to feed on the abundant nectar, pollinating the f lowers in the process. However, the involvement of this gecko does not end there. The fruits have the shape of a teat of a baby’s bottle from which a pulp with numereous seeds is secreted and eaten by the blue-tailed gecko, which disperses the seeds in its droppings. The dramatic decline of this plant species has always been thought to be due to extensive deforestation and the introduction of pigs, monkeys and goats. A tiny invasive ant (Technomyrmex albipes),

native to the Indo-Pacific area, builds walls of clay that seal the flowers, creating a chamber in which it farms mealy bugs for the sugary sap they excrete. It is not the sap-sucking mealy bug that causes a problem, but the aggressive ants attack and chase off the geckos, preventing the plant from producing seeds. Attempts to cultivate Roussea have been unsuccessful until recently because the seeds need to be cleaned from the pulp and must pass through the gut of a gecko for germination. Embryos from an unripe fruit

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as there are locules. Fruits are capsules that open apically by lateral pores or loculicidally, sometimes a berry with numerous small seeds.

Campanulaceae include annual and perennial herbs, shrubs, vines and small trees (to 15 m tall), sometimes with swollen trunks (pachycaul), often with swollen underground roots. They are mostly terrestrial, sometimes aquatic or epiphytic. The sap is usually milky white, but sometimes differently coloured. Leaves are alternate, often with a basal rosette, rarely opposite or whorled, simple

or pinnately compound, usually petiolate (sometimes sessile), and without stipules. Blades are entire or toothed, sometimes variously dissected and sometimes sheathing at the base. Inflorescences are bracteate racemes, panicles, spikes, heads or umbels, or the f lowers are solitary. Flowers are actinomorphic or zygomorphic and usually bisexual. The five (sometimes three to ten) sepals are fused to the calyx forming a hypanthium atop the ovary, sometimes with reflexed appendages between the lobes. The usually five (sometimes four to ten) petals are fused to form a cup-shaped or bilabiate corolla, with the lobes fused or split to various levels, and sometimes with nectariferous spurs or sacs. Stamens are as many as petals, alternating with them, and fused to the base of the corolla tube, on the hypanthium rim or on the top of the ovary. Filaments are free or variably fused, and the basifixed (rarely dorsifixed) free or partially fused anthers open by longitudinal slits. Campanulaceae can be characterised by their special mechanism of secondary pollen presentation to pollinators, with pollen deposited on the style due to fusion of the introrse anthers around the often hairy style. The inferior (rarely superior or half-inferior) ovary is composed of two to five locules, forming one or many, occasionally up to ten, locules. The single style is hairy below the apex and topped by as many stigmas

Nemacladus glanduliferus var. orientalis, Las Vegas, Nevada, USA (SS) [429]

Campanula isophylla, Royal Botanic Gardens, Kew, UK [429]

Azorina vidalii, private garden, Kingtsonupon-Thames, Surrey, UK [429]

have now been isolated at the Royal Botanic Gardens, Kew, and plantlets are now growing in vitro in micropropagation. Some have now been released from their vials and grow well. This technique may help save the species from certain extinction, at least in cultivation. Etymology: Roussea commemorates Genevan philosopher Jean-Jacques Rousseau (1712–1778).

429. CAMPANULACEAE Bellflower family

Distribution: This is a nearly globally distributed family, but they are absent from many of the desert areas and permanently frozen parts of the world, although extending far into the Arctic. The greatest diversity is in warm temperate regions of the world, and there are some radiations on oceanic islands, especially at high elevation. Phylogeny and evolution: Few fossils of Campanulaceae are known. The most reliable are fossil seeds from the Miocene of Poland. It has been suggested that Lobelioideae originated in Africa and dispersed widely from there, with extensive diversification in South America, Africa and the Pacific. The Hawaiian radiation (c. 130 species) evolved from a single woody ancestor that arrived there c. 13 million years ago. Campanuloideae have diversified especially in western and Central Asia, with 100 species of Campanula in Turkey alone. There are complex diversification patterns that involved dispersal and isolation. Fleshy fruits have evolved independently several times in the family. The three monophyletic subfamilies used here are only tentative and need further study. Generic limits need attention, especially the clades

Plants of the World

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ASTERALES

Legousia speculum-veneris, Gran Meteora, Greece [429]

Canarina canariensis, private garden, Kingtson-upon-Thames, Surrey, UK [429]

Trachelium caeruleum, Sicily, Italy [429]

Jasione montana, Lake District, UK

including the paraphyletic Campanula and Lobelia, which will have to be expanded to establish monophyly. Genera and species: Campanulaceae include 84 genera and c. 2,300 species, in three subfamilies: Nemacladoideae – Nemacladus (13) and Pseudonemacladus (1); Campanuloideae – Adenophora (65), Asyneuma (33), Azorina (1), Berenice (1), Campanula (421), Canarina (3), Codonopsis (59), Craterocapsa (5), Cryptocodon (1), Cyananthus (23), Cyclocodon (2), Cylindrocarpa (1), Echinocodon (1), Edraianthus (13), Favratia (1), Feeria (1), Githopsis (4), Gunillaea (2), Hanabusaya (1), Heterochaenia (3), Heterocodon (1), Himalocodon (1), Homocodon (2), Jasione (16), Legousia (7), Merciera (6), Michauxia (7), Microcodon (4), Muehlbergella (1), Musschia (2), Namacodon (1), Nesocodon (1), Ostrowskia (1), Pankycodon (1), Peracarpa (1), Petromarula (1), Physoplexis (1), Phyteuma (22), Platycodon (1), Prismatocarpus (28), Pseudocodon (8), Rhigiophyllum 592

EUDICOTS

Christenhusz, Fay & Chase

[429]

Wahlenbergia capensis, Mt Benia, Western Australia [429]

Petromarula pinnata, Royal Horticultural Society Garden, Wisley, UK [429]

(1), Roella (20), Sergia (2), Siphocodon (2), Theilera (2), Trachelium (2), Treichelia (1), Triodanis (6), Wahlenbergia (260) and Zeugandra (2); Lobelioideae – Apetahia (4), Brighamia (2), Burmeistera (>100), Centropogon (213), Clermontia (22), Cyanea (78), Cyphia (64), Cyphocarpus (3), Dialypetalum (5), Diastatea (5), Dielsantha (1), Delissea (15), Downingia (13), Grammatotheca (1), Heterotoma (1), Hippobroma (1), Howellia (1), Isotoma (14), Legenere (1), Lobelia (>400), Lysipomia (30), Monopsis (15), Palmerella (1), Porterella (1), Ruthiella (4), Sclerotheca (6), Siphocampylus (>230), Solenopsis (6), Trematolobelia (4), Unigenes (1) and Wimmerella (10). Uses: Some species make f leshy edible fruits, such as Canarian bellflower (Canarina canariensis) that also makes an attractive flowering vine for shade in a winter-rain, frost-free climate and produces copious edible, faintly sweet, orange fruits with numerous tiny seeds. Most Campanula species have

edible leaves, and some also produce roots that can be eaten. One of the best vegetables is C. versicolor, which has leaves rich in vitamin C that are acceptable raw. Campanula latifolia and C. persicifolia also have leaves that are good in salads. Rampion (C. rapunculus) used to have a cultivar with a turnip-like root, but this clone is no longer extant, a lost vegetable. Adenophora latifolia and A. liliifolia produce particularly tasty sweet roots that can be eaten raw or cooked. Easily grown from seed in a warm, sunny position, they unfortunately resent root disturbance and cuttings may be difficult to establish. Codonopsis is usually grown as a curiosity vine, but it also has large swollen roots that can be eaten raw or cooked, with easily grown C. ussuriensis having a smell like that of fried food. Many Campanulaceae make wonderful, long-flowering garden plants, although some may be somewhat invasive. Many species have selected forms commonly used in horticulture, mainly the genera Campanula

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and Lobelia (with especially L. erinus cultivars being popular bedding plants), but numerous species of the genera Adenophora, Asyneuma, Brighamia, Campanula (including Symphyandra), Codonopsis, Edraianthus, Hippobroma, Jasione, Legousia, Lobelia (including Hypsela and Pratia), Michauxia, Physoplexis, Phyteuma, Plat ycodon, Trachelium and Wahlenbergia are popular in perennial borders, rock gardens and woodland gardens. Island radiation: Campanulaceae have some remarkable species that evolved on oceanic islands, often involving secondary woodiness and peculiar pollination syndromes. Rare Hawaiian olulu or alula (Brighamia insignis), also colloquially called ‘cabbage on a stick’ or ‘Hawaiian palm’, was nearly extinct in its native habitats on the islands of Kauai and Niihau, where in the 1990s, only c. 50 individuals remained. Its pollinator, an endemic hawkmoth, is now extinct, and thus Lobelia oligophylla, Royal Botanic Gardens, Kew, UK [429]

Lobelia dortmanna, Blea Tarn, Lake District, UK [429]

this species would have the same fate were it not for human intervention. It is easily handpollinated and grows quickly and easily from seed, so it is now rescued by horticulture from certain extinction. Although wild populations remain small and threatened, it is now widely cultivated as a novelty houseplant or tropical garden plant. Woody Cyanea are also endemic to the Hawaiian islands, where they have evolved into nearly 80 species of thorny bushes or trees with dimorphic leaves. This may have been an adaptation to prevent them being eaten by browsers (possibly birds) that used to occur on these islands before human settlement. Another example of island woodiness is Apetahia found across the Pacific (in the Marquesas, Raiatea and Tubuai). Predominantly herbaceous Wahlenbergia has an almost cosmopolitan distribution, but it diversified particularly in South Africa and the islands of the Pacific including New Zealand. Other islands in the Atlantic

also have small radiations, such as the four endemic species of Saint Helena (including the now extinct dwarf cabbage tree, Wahlenbergia roxburghii), which also evolved secondary woodiness. It seems that the closest relative of the St Helenan species of Wahlenbergia is W. berteroi from the Juan Fernández Islands in the eastern Pacific. The Macaronesian Islands also have several endemic genera, with Musschia endemic to Madeira and the Islas Desertas, Azorina vidalii only found in the Azores, and Canarina canariensis endemic to the laurel forests on the Canary Islands with its closest relatives in East Africa. Canarina canariensis has adapted to pollination by now extinct sunbirds, but pollination also appears to be effected by other pollinators as well. There is also a great diversity of Campanulaceae on the Mascarene Islands of the Indian Ocean, where on the island of Réunion, three species of Heterochaenia and Berenice arguta can be found. All are rare and locally threatened and could be

Centropogon grandidentatus, University of California Botanical Garden, Berkeley, USA [429]

Lobelia erinus, private garden, Kingston Lobelia giberroa, Taita Hills, upon Thames, Surrey, UK [429] Kenya [429]

Hippobroma longiflora, Singapore Botanical Garden [429]

Isotoma hypocrateriformis, Cervantes, Western Australia [429]

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Nesodocon mauritianus, National Botanic Gardens of Ireland, Glasnevin [429]

Pentaphragma ellipticum, Bukit Timah Nature Reserve, Singapore (MN) [430]

saved by bringing them into cultivation as was done for Brighamia. Closely related to Heterochaenia is the Mauritian Nesocodon mauritianus, which is only known from a single locality beside a waterfall. It was the first flower to be known to produce blood-red nectar, attracting the endemic red-whiskered bulbul (Pycnonotus jocosus). There are a few other species with red nectar, the others being Trochetia (Malvaceae, also from Mauritius) and a South American species of Jaltomata (Solanaceae). Woodiness is not restricted to islands. In East Africa, the genus Lobelia has radiated, and there are species at higher elevations there that grow into sizable trees.

asymmetrical with pinnate venation often with several main veins arising from the base, the margins serrate, rarely entire or sinuate. Inflorescences are bracteate axillary cymes or cincinni, usually scorpioid, or the flowers one to three together in the axils. Flowers are bisexual and actinomorphic except for the petal-like calyx, which is formed of five lobes fused into an bell- or tube-shaped structure, usually with two of the lobes wider than the other three. The five petals are fused into a corolla that is fused at the base to the calyx and has lobes that are partially fused or split nearly to the base. Alternate with the petals are five stamens fused to the corolla tube. Anthers are basifixed and open inwardly, but often appearing to open laterally due to the developing connective. The inferior ovary is composed of two or three carpels, each forming a locule and topped with a short thick style and a capitate or conical stigma. The ovary apex has nectar pockets between the septae. Fruits are indehiscent berries with numerous seeds and the remains of the perianth at the tip.

Etymology: Campanula is a Latin diminutive of campana, a bell, referring to the bellshaped flowers.

430.PENTAPHRAGMATACEAE Scorpion’s-tail family

Pentaphragma ellipticum, Singapore (KH) [430]

Distribution: This family is restricted to tropical Asia, from southern China and Thailand throughout Southeast Asia and Malesia to New Guinea.

This is a family of coarse, perennial, somewhat succulent herbs with clear sap and elongate, fat, more or less woody rhizomes. Leaves are simple, alternate and petiolate and lack stipules. Leaf blades are basally 594

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Phylogeny and evolution: Because of similarity of the fruit, the genus Pentaphragma has often been associated with Campanulaceae, but Pentaphragmataceae form an independent clade in Asterales, separate from Campanulaceae. Their flowers

are different, and they lack latex, so it is unsurprising that the two were not found to be closely related. Pentaphragmataceae are estimated to have evolved c. 67–84 million years ago. Genera and species: The sole genus included in this family is Pentaphragma with c. 30 species. Etymology: Pentaphragma is composed of the Greek words πέντε (pente), five, and φράγμα ( fragma), a barrier. The ovary is separated from the five-lobed hypanthium by nectar pits.

431. STYLIDIACEAE Triggerplant family

Stylidiaceae are annual and perennial herbs that are sometimes somewhat woody at the base or form cushion plants. Leaves are simple, alternate and spirally arranged, without stipules and often in basal rosettes or along the stem; stolons are often formed. Aerial roots (stilt-roots) are sometimes present, elevating the plant several millimetres above the substrate. Flowers are in axillary or terminal

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scapose racemes or cymes, with bracts, often in whorls. Flowers are zygomorphic or (nearly) actinomorphic with a tubular, often glandular-hairy calyx composed of five (rarely three to seven) fused sepals. The five (or ten) petals are free (Donatia) or fused into a tubular corolla with spreading lobes, but usually only four petal lobes are prominent, the uppermost often differentiated into a small lip. The two stamens are fused with the style to form an irritable column that snaps towards the petals when triggered by the pollinator. In Donatia, there are three free stamens. Anthers are dorsifixed and open by lengthwise slits. An inferior ovary is composed of two or three carpels, fused to each form a locule or a single locule by abortion or reduction of the septum. A simple irritable style is fused with Levenhookia pusilla, near Perth, Western Australia [431]

Stylidium rigidulum, Mt Benia, Western Australia [431]

the staminal tube and topped with two or three stigmas, or (in Donatia) there are two or three free styles. Fruits are many-seeded capsules enclosed by the persistent calyx. Distribution: This family is predominantly Australian, but a few species are found in the far south of South America, the Falkland Islands, Sri Lanka, Southeast Asia, Malesia and New Zealand. The centre of diversity is southwestern Western Australia, where they grow in open heathland on seasonally moist, sandy soils that are usually poor in nutrients. Phylogeny and evolution: Stylidiaceae have been estimated to have diverged c. 39 million years ago. Fossils are rare, but fossil pollen similar to pollen of Forstera has been found

in Australian Oligocene deposits. Placement of Stylidiaceae in Asterales is confirmed by molecular and morphological evidence. Donatia exhibits plesiomorphic characters, such as the free petals, stamens and styles, and is sister to the rest. Both major clades in the family are found in Australasia and South America, which indicates multiple long-distance dispersal events around the southern oceans. Flowers of Levenhookia and Stylidium have pollination mechanisms that are linked to specific pollinators. The upper petals are modified to form a labellum that is curved over the column, making the flower appear to be four-parted. The column moves rapidly when touched, depositing or collecting pollen (or both) on the abdomen of the visiting insect. The column returns to its original Stylidium crossocephalum, Mt Benia, Western Australia [431]

Levenhookia stipitata, near Perth, Western Australia [431]

Stylidium schoenoides, before and after the style was triggered, Albany, Western Australia [431]

Stylidium torticarpum, Mt Benia, Western Australia [431]

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position so it can strike again, the second time usually to collect rather than to deposit pollen so that self-pollination is excluded, although self-pollination might occur in some species. Stylidium schoenoides is called the “cow kick” because of the great force behind the released trigger.

Stylidiaceae are found. Like Drosera, the glands produce proteases and can thus digest the insects that have been caught. It has been shown that amino acids are absorbed by the surface of Stylidium, so at least some species of trigger plants appear to be fully carnivorous.

Genera and species: Stylidiaceae encompass four genera and c. 220 species: Donatia (2), Forstera (9), Levenhookia (8) and Stylidium (c. 200).

Etymology: Stylidium is deminutive Greek of στύλος (stylos), a pillar or style, in reference to the motile style of these species.

Uses: Species are occasionally cultivated as unusual garden ornamentals, especially in Australia. There is potential in horticulture for these beautiful and interesting plants. Carnivory: Several Stylidiaceae are known to trap small insects such as gnats and midges using mucilage-secreting glandular hairs on their inf lorescences and stems. Because these species are remarkable for their active pollination mechanism, attention has focused on that rather than on possible carnivory. However, Stylidium uses the same mechanism to trap their prey as Drosera or Byblis, both genera that occur in similar habitats and often in the same places where Alseuosmia macrophylla, Dunedin Botanical Garden, New Zealand (JC) [432]

432. ALSEUOSMIACEAE Toropapa family

These are terrestrial and epiphytic shrubs with alternate, nearly opposite or falsely whorled, simple leaves without stipules. Leaf blades have pinnate venation and entire or serrate margins. Inflorescences are alternate

Alseuosmia macrophylla in fruit, Dunedin Botanical Garden, New Zealand (JC) [432]

or terminal cymose fascicles, sometimes racemose or the flowers solitary, occasionally appearing from main axes (cauliflorous). Flowers are actinomorphic, bisexual or functionally unisexual (Periomphale) and have bracteate pedicels. The receptacle is fused to the ovary and terminated by four or five (to six) sepals fused at their base. The four or five (to six) petals are fused to form a urn-, funnelor bell-shaped corolla with winged lobes. Stamens are as many as and alternating with the corolla lobes, and filaments are fused to the base of the corolla tube or the anthers sessile (Platyspermation). Anthers are dorsifixed and open inwardly by longitudinal slits. A nectar disk is often present atop the ovary. The inferior (or half-inferior) ovary is composed of two or three fused carpels, each forming a locule. A single terminal style has a discoid, bi- or tri-lobed stigma. Fruits are berries, rarely capsules (Platyspermation) with one to several seeds. Distribution: This is an Australasian family (New Guinea, eastern Australia, New Caledonia and New Zealand). Phylogeny and evolution: Molecular evidence placed the genera of Alseuosmiaceae close to

Periomphale balansae, Ouaième, New Caledonia (JM) [432]

Periomphale balansae in fruit, Paéoua, New Caledonia (JM) [432]

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Alseuosmia quercifolia, New Zealand (JC) [432]

ASTERALES

EUDICOTS

Phelline comosa, New Caledonia [433]

Phelline comosa, New Caledonia [433]

Phelline macrophylla, New Caledonia [433]

Phellinaceae and Argophyllaceae in Asterales. The genera were always considered poorly fitting members of other families: Alseuosmia in Caprifoliaceae, Periomphale in Gesneriaceae and Wittsteinia in Ericaceae, for instance. Inclusion in a broad Escalloniaceae had also been proposed. Platyspermation, originally described in Myrtaceae, was poorly known and despite its morphological differences is also placed in this family, even though Platyspermatiaceae has been proposed for it.

Phellinaceae are evergreen, unisexual, understorey shrubs and trees. Their simple leaves are alternate, spirally arranged in false whorls at the branch tips, with petioles (sometimes short) but no stipules. Blades have pinnate venation with an intramarginal vein and an entire or crenate margin. Inflorescences are axillary or subterminal racemes or panicles. Flowers are unisexual and actinomorphic. The four to five (or six) sepals are fused at the base with the same number of free, valvate petals and stamens alternating with the petals. Anthers are dorsifixed and open by lengthwise slits. A pistillode is present in the middle of male flowers, whereas female flowers do not have staminodes. Carpels are as many as the sepals (usually four or five) and fused to form a multi-locular, superior ovary. Fruits are globose or lobed drupes containing up to five seeds.

Etymology: Phelline is derived from the Greek φελληνος ( fellinos), made of cork, referring to the corky fruits and seeds.

Genera and species: Alseuosmiaceae include five genera and ten species: Alseuosmia (5), Crispiloba (1), Periomphale (1), Platyspermation (1) and Wittsteinia (2). Uses: There are no economic uses of this family, but some species of Alseuosmia and Wittsteinia with wonderfully scented flowers would make attractive ornamental plants. Etymology: Alseuosmia is composed of the Greek words άλσος (alsos), a grove, and οσμή (osme), fragrance, referring to the strongly fragrant flowers.

433. PHELLINACEAE Corkfruit family

Distribution: The family is only found in New Caledonia. Phylogeny and evolution: The systematic position of Phelline was originally in Sapotaceae and later Rutaceae. It was then placed in Aquifoliaceae, from which it differed in numerous characters, varying from wood anatomy, and flower and inflorescence structure, to ovule position and leaf venation. However, it was maintained in Aquifoliaceae until molecular studies firmly placed it in Asterales, close to Alseuosmiaceae and Argophyllaceae, both Australasian families. Genera and species: The sole genus in this family is Phelline with 12 species.

434. ARGOPHYLLACEAE Silverleaf family

Argophyllaceae include shrubs and small trees with an indumentum of T-shaped hairs and sometimes zigzag branches. Leaves are simple, alternate or fascicled on short branches (brachyblasts) and lack stipules. Blades have pinnate venation and entire or serrate margins. Inflorescences are axillary or terminal panicles, racemes or cymose fascicles, or sometimes flowers are solitary. Flowers are actinomorphic and bisexual. The usually five (to eight) sepals are basally fused and persistent in fruit. The five (or six, rarely eight), yellow or white petals are free or fused basally and often have fringed appendages (corollar ligules) on the upper side. Stamens are as many as and alternating with the petals, the anthers dorsifixed and opening inwardly by lengthwise slits. The (half-) inferior ovary is composed of two to four carpels forming one to six locules. The apical style bears a capitate two- to five-lobed stigma. Fruits are loculicidal capsules or drupes. Plants of the World

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ASTERALES

Argophyllum vernicosum, New Caledonia (TW) [434]

EUDICOTS

Corokia cotoneaster, Royal Botanic Gardens, Kew, UK [434]

Distribution: This family is distributed in parts of Australasia, including eastern Australia, New Caledonia, New Zealand, Lord Howe Island, and Rapa (French Polynesia). Phylogeny and evolution: Even though the two genera share many characters, especially the T-shaped hairs and corollar ligules, they were previously placed in different families; Argophyllum in Saxifragaceae or Escalloniaceae and Corokia in Cornaceae. Morphological and molecular evidence placed these genera together as sister to Phellinaceae in Asterales. Genera and species: The two genera in Argophyllaceae include c. 21 species: Argophyllum (c. 15) and Corokia (6). Uses: New Zealand Corokia species and especially cultivars of the hybrid C. ×virgata are occasionally grown in gardens as ornamentals in New Zealand, Australia and the UK. Argophyllum nullumense is sometimes grown in Australia for its foliage. Many species are threatened, and species conservation through horticulture seems to be a suitable option. Etymology: Argophyllum is composed of the Greek words άργυρος (argyros), silver, and φύλλων ( fyllon), leaf. 598

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435. MENYANTHACEAE Bogbean family

This is a family of perennial and annual, usually aquatic herbs with emergent or floating leaves and creeping rhizomes or clustered rootstocks. The alternate (rarely opposite), petiolate leaves are simple or rarely trifoliate (Menyanthes), without stipules, and often forming a rosette. Blades are monomorphic or dimorphic, linear to cordate or reniform, with entire and often undulate, crenate or toothed margins, sometimes irregularly so. Inflorescences are simple or branched cymes, racemes, panicles, umbels, heads or clusters, sometimes the f lowers paired or solitary. Flowers are bisexual (rarely functionally unisexual) and actinomorphic. The usually five sepals are fused, at least at the base, and persistent in fruit. The (four or) five petals are fused, and the inner surfaces of the lobes often have a fimbriate, crested or fringed margin. The five

Corokia macrocarpa, Royal Botanic Gardens, Kew, UK [434]

(or four) stamens alternate with the petals, the filaments are inserted at the base of the tube. Anthers are dorsifixed and open by lengthwise slits. The superior or half-inferior ovary is composed of two carpels fused to form a single locule. Terminal styles are of various lengths, sometimes absent, and they are topped with a bilobed, papillate stigma. Fruits are dehiscent or indehiscent capsules, rarely a berry. Distribution: Menyanthaceae have an almost cosmopolitan distribution in wetland communities, but they are absent from the great deserts (Sahara, central Australia). Phylogeny and evolution: Traditionally placed close to Gentianaceae, Menyanthaceae have been shown to be anatomically, chemically, palynologically and molecularly close to Goodeniaceae in Asterales. Heterostyly in this clade is labile. Genera and species: The family includes six genera and c. 57 species: Liparophyllum (1), Menyanthes (1), Nephrophyllidium (1), Nymphoides (c. 40), Ornduffia (7) and Villarsia (c. 7). Uses: Species of Menyanthes, Nymphoides and Villarsia are occasionally cultivated as pond plants.

ASTERALES

EUDICOTS

Menyanthes trifoliata, Bix Bottom, UK [435]

Nymphoides beaglensis, Royal Botanic Gardens, Kew, UK [435]

Nymphoides cristata, private garden, Tustin, California, USA [435]

Nymphoides indica, Nouméa, New Caledonia [435]

Nymphoides peltata in fruit, Royal Botanic Gardens, Kew, UK [435]

Ornduffia parnassifolia, near Denmark, Western Australia [435]

Etymology: Menyanthes is derived from the Ancient Greek μενανθος (menanthos), the name for another aquatic plant (probably the water gentian, Nymphoides peltata). It is derived from μηνυος (menyos), small, and άνθος (anthos), flower.

but often with an axillary tuft of hairs. Leaf blades have entire to toothed margins and pinnate venation. Inflorescences are cymes, thyrses, racemes, spikes, false umbels or heads, the flowers rarely solitary. Flowers are bisexual and zygomorphic, rarely actinomorphic (Brunonia). The five, rarely three, sepals are free or fused or reduced to a little rim, sometimes appearing to be absent. The corolla is composed of five fused petals and is usually split on the upper side in one or two portions, becoming fan-like or bilabiate, sometimes pouched or spurred on one side. Petals are more or less winged, often with small lobes (auricles) around the stamens. Five stamens alternate with the petal lobes and are free or fused with the petal tube. Anthers are free or fused, basifixed and opening inwardly by lengthwise slits. The superior, half-inferior or inferior ovary is composed of four carpels fused to form two usually incomplete locules, rarely one or four. The style is entire with a hollow pollen cup (indusium) enclosing the more or less bifid tip, rarely two- or three-branched. Fruits are

drupes, nutlets or capsules, rarely a schizocarp separating transversely into single-seeded woody segments.

436. GOODENIACEAE Fanflower family

Herbs, shrubs and rarely scramblers make up this family. They are glabrous or variously hairy, the hairs sometimes glandular or viscid and becoming glossy with age. Leaves are simple and mostly alternate and spirally arranged, rarely opposite and without stipules,

Distribution: The greatest diversity in this family is centred in Australia and New Guinea where the bulk of the genera and species is found. Goodenia pilosa occurs throughout tropical Asia, extending north to Indochina, the Philippines and southern China, and three other species of Goodenia are found in Southeast Asia. Selliera radicans occurs in New Zealand and Chile. Some Scaevola species are an important component of the beach strand flora of islands and tropical coasts along the Pacific, Atlantic and Indian Oceans. Scaevola plumieri and S. taccada are particularly widespread. Phylogeny and evolution: Goodeniaceae have usually been associated with Campanulaceae because of their secondary presentation of pollen in a special structure on the style. On the basis of phytochemical and DNA results, a closer relationship to Calyceraceae Plants of the World

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ASTERALES

EUDICOTS

Dampiera teres, Mt Benia, Western Australia

Coopernookia georgei, Kings Park and Botanical Garden, Perth, Western Australia [436]

[436]

Goodenia amplexans, South Australia [436]

Velleia trinervis, Mt Benia, Western Australia [436]

Scaevola spinescens. Port Augusta, South Australia [436]

Lechenaultia formosa, Cocklebiddy, Western Australia [436]

Brunonia australis, Western Australia (KD) [436]

Scaevola phlebopetala, Mt Benia, Western Australia [436]

Scaevola plumieri, Galápagos Islands [436]

and Asteraceae has been proposed, and with Menyanthaceae as sister they form a morphologically well-defined clade. Crown group Goodeniaceae are estimated to have diversified c. 67 million years ago. There are two main clades, Lechenaultia and allies, and the other Scaevola, paraphyletic Goodenia and relatives. Within the second clade, Brunonia (formerly Brunoniaceae) is sister to the rest, and most of the numerous features by which it differs from other Goodeniaceae appear to be autapomorphies.

Lechenaultia (27), Pentaptilon (1), Scaevola (c. 130), Selliera (1), Velleia (21) and Verreauxia (3).

and collector with an interest in sedges and seaweeds.

Genera and species: Goodeniaceae include 12 genera, with c. 440 species: Anthotium (3), Brunonia (1), Coopernookia (6), Dampiera (c. 65), Diaspasis (1), Goodenia (c. 180), 600

Lechenaultia linarioides, Western Australia [436]

Christenhusz, Fay & Chase

Uses: Beach naupaka (Scaevola taccada) is sometimes used to prevent beach erosion. Several species from various genera are cultivated in Australia, but most commonly cultivated elsewhere is Scaevola aemula and cultivars, which has become a popular annual bedding and hanging-basket plant. Scaevola spinescens is being investigated for use as a cancer treatment. Etymology: Goodenia is named in honour of the Bishop of Carlisle Samuel Goodenough (1743–1827), who was an amateur botanist

437. CALYCERACEAE Balsamleaf family

This is a family of perennial, rarely annual, herbs that can be sometimes slightly woody,

ASTERALES

EUDICOTS

Boopis agglomerata, Chile (JH) [437]

often stemless with a leaf rosette and scapelike flowering shoots. Leaves are alternate, usually in a rosette and somewhat fleshy and sessile or narrowed basally to form a short petiole, without stipules. Blades are simple or (pinnately) dissected, with entire, wavy, toothed or serrate margins. Inflorescences are heads that are surrounded by triangular or lanceolate, more or less fused, involucral bracts that are arranged into scapose inflorescences. Flowers are actinomorphic and bisexual or rarely functionally male. The five (or four to six) sepals are fused to each other and to the top of the ovary, entire and unlobed or with teeth or short lobes. The four or five (rarely six) white or green petals are fused to form a cylindrical to funnel-shaped corolla with short lobes. Stamens are enclosed in the corolla tube, fused at various levels to the corolla and as many as and alternate with the petal lobes. Filaments are fused for almost their entire length into a tube, and anthers are also at least basally fused and open inwardly into the tube by lengthwise slits. Appendages are sometimes present atop the connective. Nectar glands are present at the base of the filament tube opposite the petal lobes. The inferior ovary is composed of two fused carpels forming a single locule. Fruits are achenes that are crowned with the calyx and sometimes somewhat woody and spiny. Distribution: Calyceraceae occur in southern South America, north to southern Peru,

Calycera crassifolia, Brazil (MT) [437]

Bolivia and Bahia state (Brazil) with a single species in the Falkland Islands. Most species are found in high-elevation grasslands and meadows, sometimes in coastal sand dunes and as agricultural weeds. A population of one species was found to have become naturalised in eastern North America, but it did not persist. Phylogeny and evolution: Originally Calyceraceae were placed in Dipsacales or Asteraceae, but with doubts expressed by the authors. Already in the 19th century, Calyceraceae were recognised and placed between Dipsacaceae and Asteraceae. Molecular studies have indicated a close relationship with Asteraceae, to which they are sister. Six genera have been widely accepted, but their circumscription is unsatisfactory. Acicarpha, Calycera and Gamocarpha are well defined, but the other genera (Acarpha, Boopis, Moschopsis and Nastanthus) are not and should be treated as a single genus, Boopis being the oldest name. Boopis in the broad sense diverged from the rest less than 50 million years ago. Genera and species: Calyceraceae include four genera and c. 60 species: Acicarpha (3), Boopis (c. 30), Calycera (c. 20) and Gamocarpha (7). Etymology: Calycera is composed of the Greek words καλυξ (kalyx), a calyx or chalice, and κέρας (keras), horn.

438. ASTERACEAE Daisy family

This large family includes annuals, biennials, perennials, subshrubs, shrubs, vines and trees, usually terrestrial, sometimes epiphytic or aquatic. Plants are sometimes gland-dotted and aromatic, and stems can be normally developed, herbaceous or woody, or variously swollen (pachycaul). Leaves are alternate or opposite, sometimes in basal rosettes, rarely in whorls, commonly petiolate and usually not stipulate, the leaf bases sometimes decurrent on the stems. Leaf blades are simple, rarely compound, but frequently the margins one- or two-times pinnatifid or palmatifid, sometimes lobed. Inflorescences are axillary or terminal indeterminate heads (capitula), with each head usually comprising surrounding whorls (or spirally arranged) of involucral bracts (phyllaries) on the inflorescence axis (receptacle) and (one to) five to >300 florets. Individual capitula can be sessile, or each is borne on a peduncle, the heads borne singly or in cyme-, corymb-, raceme- or spike-like Plants of the World

601

ASTERALES

EUDICOTS

ray flower

disc flower disc involucre phyllaries (bracts) peduncle

Capitulum, the Asteraceae inflorescence (Bellis perennis) [438]

compositions. Phyllaries are usually borne in one to five (or up to 15 or more) series. The receptacle is usually flat, cup-shaped, conical or columnar, often bearing scale-like bracts (palea) that individually subtend some or all of the florets. Flowers (florets) are bisexual, unisexual or sterile, the flowers of two types, actinomorphic disk flowers and zygomorphic ray flowers, with one or both kinds in a single head. Sepals are highly modifed, forming a pappus of bristles, awns or scales, sometimes different kinds found in a single flower. Petals are fused into a (three- to) five-lobed corolla, which is split on one side in ray flowers and fused into a tube in disk flowers. The (four or) five stamens alternate with the corolla lobes, and filaments are inserted on the corolla tube, the filaments usually distinct and the basifixed anthers usually fused to form a tube around the style (rarely filaments are connate and anthers free; e.g. in Ambrosiinae and Heliantheae), opening inwardly into the anther tube by lengthwise slits. Inferior ovaries are composed of two carpels fused to form a single locule with a single terminal style that is ringed by a nectary at the base. Through the anther tube, the style grows and pushes pollen out so that it is presented on the top of the style. The top of the style is branched in two with papillate stigmas that usually recurve. Fruits are single-seeded achenes (cypselae) that are usually dry and somtimes beaked or

602

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winged and often with a persistent pappus that helps with dispersal, rarely a fleshy drupe (Chrysanthemoides). Distribution: Asteraceae are a nearly worldwide family and especially rich in species and abundance in arid and semi-arid regions of subtropical and lower to middle temperate latitudes, but also in mesic montane and oceanic environments. There are marked radiations in the Americas, southern and eastern Africa, the Mediterranean, Central Asia and Australasia. Phylogeny and evolution: This family can be found in practically every environment. It is estimated to have separated from Calyceraceae c. 50 million years ago but initially would have been restricted to southern South America. A macrofossil (Raiguenrayun) from Patagonia, assignable to Mutisioideae or Asteroideae, is c. 47 million years old and together with fossil pollen evidence indicates that the early radiation in Asteraceae must have occurred in the Oligocene, when the pappus evolved, allowing seeds to drift on the winds and gain a wider distribution. From South America the first migrations must have occurred to southern Africa, and many of these Asteraceae that made the continental jump evolved features — shrubby or treelike habits etc. — common in island plants.

In Africa, many lineages arose from a second radiation and dispersal events, to Europe, Asia and Australasia and back into South America. Some lineages in Asteroideae have genera or generic pairs with wide continental disjunctions, which could have only been caused by long-distance dispersal events. Because of their easily recognisable composite inflorescence structure, Asteraceae (also called Compositae, ‘composites’ or ‘comps’) have long been recognised as a natural group, and circumscription of the group has never been controversial, although they have sometimes been divided into several families in earlier systems but then usually grouped in their own order. Molecular phylogenetic analyses changed this picture, and it has been shown that Calyceraceae are the closest relatives of Asteraceae; these two together with a couple of other smallish families form a clade with the campanulids currently circumscribed as Asterales. A Late Eocene origin has been hypothesised for Asteroideae in South America based on the early-branching South American Barnadesioideae. Fossil pollen data also point to an Eocene origin for Asteraceae in South America with migration to North America by the Oligocene, but other studies reported Mutisioideae pollen from the Palaeocene-Eocene of South Africa, suggesting an earlier, West Gondwana (southern Africa or Australia) origin for the family. This may indicate that Asteraceae arrived in North America via long-distance dispersal or island hopping before closure of the Panamanian Isthmus. Traditionally, a tribal classification within Asteraceae was adopted in the 19th and early 20th centuries. Molecular systematics established that a South Amercian clade is sister to the rest of Asteraceae, which was later elevated as subfamily Barnadesioideae. Further molecular and morphological phylogenetic analyses have deepened our knowledge of interrelationships of tribes in Asteraceae and led to a proposal of a phylogenetic classification for the family with 12 subfamilies (Bar nadesioideae, Mutisioideae, Stiftioideae, Wunderlichioideae, Gochnatioideae, Necastocleidoideae, Carduoideae, Pertyoideae, Gymnarrhenoideae,

ASTERALES

EUDICOTS

Cichorioideae, Corymbioideae and Asteroideae). However, because classification within Asteraceae has often been at only the tribal level and a dozen subfamilies seems unreasonable, we think it is sensible to tentatively accept Barnadesioideae and Mutisioideae at the subfamilial level because they are more morphologically and genetically distinct from the rest, and tentatively treat all others under Asteroideae. There are a large number of genera, many including one or only a few species, which is due to excessive splitting when a larger genus has been found to be paraphyletic to another smaller genus. This has led to the disintegration of several formerly wellestablished genera, such as Artemisia, Aster, ‘Cacalia’, Chrysanthemum, Centaurea, Eupatorium, Hieracium, Senecio, Vernonia etc., and to the expansion of others, such as Lactuca (now including Cephalorrhynchus, Cicerbita, Lactucella, Lagedium, Mulgedium, Mycelis and Scariola). Nevertheless, splitting cannot always be avoided, and thus we have accepted some genera here in spite of our opinion that there is a need for a general enlargement of generic concepts: e.g. Orchidaceae have a larger number of species but a much smaller number of genera than are recognised in Asteraceae. The literature on classification of Asteraceae is complex, and it is extremely difficult to come to a complete standard consensus generic classification simply because synantherologists (taxonomists of Asteraceae) do not always agree. However, Barnadesia arborea, Ecuador [438]

classification within families is not within the scope of this book, and we hope as a minimum to produce a good overview of the diversity of this family, which is indeed substantial. The list of genera and associated species numbers provided below is thus an approximation. Genera and species: Asteraceae are a large family, one of the two largest (the other Orchidaceae) with c. 1,627 genera and about 24,700 species in three subfamilies: Barnadesioideae (9 genera) – Arnaldoa (3), Barnadesia (18), Chuquiraga (23), Dasyphyllum (40), Doniophyton (2), Duseniella (1), Fulcaldea (1), Huarpea (1) and Schlechtendalia (1); Mutisioideae (c. 79 genera) – Achnopogon (2), Acourtia (c. 40), Adenocaulon (>5), Ainsliaea (50), Amblysperma (2), Ameghinoa (1), Aphylloclados (5), Berylsimpsonia (2), Brachyclados (3), Brachylaena (11), Burkartia (1), Calorezia (2), Catamixis (1), Cephalopappus (1), Chaetanthera (43), Chaptalia (56), Chimantaea (10), Chucoa (2), Cnicothamnus (), Criscia (1), Cyclolepis (1), Dicoma (57), Dolichlasium (1), Duidaea (4), Eriachaenium (1), Erythrocephalum (13), Gerbera (c. 35), Gladiopappus (1), Glossarion (2), Gochnatia (c. 60), Gongylolepis (15), Gypothamnium (1), Hecastocleis (1), Holocheilus (6), Hyalis (2), Hyaloseris (7), Ianthopappus (1), Jungia (26), Leibnitzia (4), Leucheria (46), Leunisia (1), Lophopappus (6), Lulia (1), Lycoseris (11), Macrachaenium (1), Macroclinidium (3), Marticorenia (1), Moscharia (2), Mutisia (59), Myripnois (1), Nassauvia (51), Neblinaea (1),

Onoseris onoseroides, Guatemala [438]

Nouelia (1), Oldenburgia (4), Onoseris (31), Oxyphyllum (1), Pachylaena (2), Panphalea (6), Pasaccardoa (4), Perdicium (2), Perezia (32), Pertya (15), Plazia (3), Pleiotaxis (26), Pleocarphus (1), Polyachyrus (7), Proustia (3), Quelchia (4), Richterago (15), Salcedoa (1), Stenopadus (14), Stifftia (6), Stomatochaeta (6), Tarchonanthus (2), Trichocline (22), Trixis (c. 65), Uechtritzia (3), Urmenetia (1) and Wunderlichia (5); Asteroideae (c. 1,535 genera) – Aaronsohnia (2), Abrotanella (18), Acamptopappus (2), Acanthocephalus (2), Acanthocladium (1), Acanthodesmos (1), Acantholepis (1), Acanthospermum (8), Acanthostyles (2), Achillea (130), Achnophora (1), Achyrachaena (1), Achyrocline (32), Achyropappus (1), Acilepidopsis (1), Acilepis (c. 36), Acmella (30), Acomis (4), Acrisione (2), Acritopappus (16), Actinobole (4), Acunniana (1), Adelostigma (2), Adenanthellum (1), Adenocritonia (1), Adenoglossa (1), Adenoon (1), Adenophyllum (10), Adenostemma (24), Adenostyles (4), Adenothamnus (1), Aedesia (2), Aequatorium (c. 20), Aetheorhiza (1), Afroaster (18), Ageratella (1), Ageratina (c. 245), Ageratinastrum (2), Ageratum (c. 40), Agoseris (11), Agrianthus (6), Ajania (34), Ajaniopsis (1), Akeassia (1), Alatoseta (1), Albertinia (1), Aldama (2), Alepidocline (4), Alfredia (5), Aliella (3), Allagopappus (2), Allardia (8), Allocephalus (1), Alloispermum (7), Allopterigeron (1), Almutaster (1), Alomia (5), Alomiella (2), Alvordia (4), Amauria (3), Amauriopsis (1), Ambassa (1), Amberboa (6), Amblyocarpum (1), Amblyolepis (1),

Perezia linearis, Royal Botanic Gardens, Kew, UK [438]

Plants of the World

603

ASTERALES

Mutisia oligodon, Royal Botanic Gardens, Kew, UK [438]

EUDICOTS

Mutisia subspinosa, Royal Botanic Gardens, Kew, UK [438]

Amblyopappus (1), Amboroa (2), Ambrosia (40), Amellus (12), Ammobium (3), Amolinia (1), Ampelaster (1), Amphiachyris (2), Amphiglossa (11), Amphipappus (1), Amphoricarpos (4), Anacyclus (12), Ananthura (1), Anaphalioides (7), Anaphalis (10), Anaxeton (9), Ancathia (1), Ancistrocarphus (2), Anderbergia (6), Andryala (c. 20), Anemocarpa (3), Angeldiazia (1), Angianthus (18), Anisocarpus (2), Anisochaeta (1), Anisocoma (1), Anisopappus (31), Anisothrix (2), Antennaria (45), Anteremanthus (1), Anthemis (c. 170), Antillanthus (17), Antillia (1), Antiphiona (2), Antithrixia (1), Anvillea (2), Apalochlamys (1), Aphanactis (8), Aphanostephus (4), Apopyros (2), Aposeris (1), Apostates (1), Apowollastonia (8), Arbelaezaster (1), Archanthemis (4), Archibaccharis (32), Archiserratula (1), Arctanthemum (1), Arctium (11), Arctogeron (1), Arctotheca (5), Arctotis (50), Argentipallium (6), Argyranthemum (24), Argyroglottis (1), Argyroxiphium (4+1 extinct), Arida (9), Aristeguietia (21), Arnica (29), Arnicastrum (2), Arnoglossum (8, the former Cacalia of America), Arnoseris (1), Arrhenechthites (6), Arrojadocharis (2), Arrowsmithia (1), Artemisia (c. 400), Artemisiella (1), Artemisiopsis (1), Asanthus (3), Ascidiogyne (2), Asplundianthus (17), Aster (c. 600), Asteridea (9), Asteriscus (9), Asteropsis (1), Asterothamnus (7), Astranthium (12), Athanasia (39), Athrixia (14), Athroisma (12), Atractylis (14), Atractylodes (11), Atrichantha (1), Atrichoseris (1), Austrobrickellia (3), Austrocritonia (4), Austroeupatorium (13), Austrosynotis (1), Axiniphyllum (5), Ayapana (14), Ayapanopsis 604

Christenhusz, Fay & Chase

Gerbera ×hybrida ‘Entourage’, Emsflower, Germany [438]

(15), Aynia (1), Aztecaster (2), Baccharis (c. 350), Baccharoides (24), Badilloa (10), Baeriopsis (1), Bafutia (1), Bahia (10), Bahianthus (1), Bahiopsis (12), Baileya (3), Bajacalia (3), Balduina (3), Baltimora (2), Barkleyanthus (1), Barrosoa (11), Bartlettia (1), Bartlettina (37), Basedowia (1), Batopilasia (1), Bebbia (2), Bechium (3), Bedfordia (3), Bejaranoa (2), Bellida (1), Bellis (8), Bellium (4), Belloa (9), Benitoa (1), Berardia (1), Berkheya (75), Berlandiera (5), Berroa (1), Bertilia (1), Bethencourtia (3), Bidens (c. 200), Bigelowia (2), Bishopalea (1), Bishopanthus (1), Bishopiella (1), Bishovia (2), Blainvillea (10), Blakeanthus (1), Blakiella (1), Blanchetia (1), Blennosperma (2), Blennospora (3), Blepharipappus (1), Blepharispermum (15), Blepharizonia (2), Blumea (c. 100), Blumeopsis (1), Boeberastrum (2), Boeberoides (1), Bolandia (2), Bolanosa (1), Boltonia (5), Bombycilaena (3), Borrichia (32), Bothriocline (c. 30), Brachanthemum (10), Brachionostylum (1), Brachyglottis (37), Brachyscome (c. 75), Brachythrix (5), Brenandendron (3), Brickellia (110), Brickelliastrum (2), Brocchia (1), Bryomorphe (1), Buphthalmum (2), Caatinganthus (2), Cabobanthus (2), Cabreriella (2), Cacaliopsis (1), Cacosmia (3), Cadiscus (1), Caesulia (1), Calanticaria (5), Calea (c. 110), Calendula (c. 12), Callicephalus (1), Callilepis (5), Callistephus (1), Calocephalus (11), Calomeria (1), Calostephane (6), Calotesta (1), Calotis (28), Calycadenia (10), Calycoseris (2), Calyptocarpus (2), Camchaya (5), Campova ssour ia (1), Camptacra (2), Campuloclinium (14), Canadanthus (1),

Cancrinia (4), Cancriniella (1), Capelio (3), Caputia (5), Cardopatium (2), Carduncellus (4), Carduus (91), Carlina (28), Carlquistia (1), Carminatia (3), Carpesium (25), Carphephorus (7), Carphochaete (6), Carramboa (5), Carthamus (51), Cassinia (c. 18), Castanedia (1), Castrilanthemum (1), Catananche (5), Catatia (2), Catolesia (1), Caucasalia (4), Cavalcantia (2), Cavea (1), Caxamarca (1), Celmisia (69), Centaurea (c. 736), Centaurodendron (3), Centauropsis (8), Centaurothamnus (1), Centipeda (5), Centrapalus (1), Centratherum (2), Centromadia (4), Cephalipterum (1), Cephalosorus (1), Ceratogyne (1), Ceruana (1), Chacoa (1), Chaenactis (18), Chaetadelpha (1), Chaetopappa (11), Chaetoseris (18), Chaetymenia (1), Chamaechaenactis (1), Chamaegeron (4), Chamaemelum (2), Chamaepus (1), Charadranaetes (1), Chardinia (1), Cheirolophus (23), Chersodoma (9), Chevreulia (5), Chiliadenus (9), Chiliocephalum (2), Chiliophyllum (2), Chiliotrichiopsis (4), Chiliotrichum (2), Chionolaena (6), Chionopappus (1), Chlamydophora (1), Chloracantha (1), Chondrilla (25), Chondropyxis (1), Chresta (11), Chromolaena (165), Chromolepis (1), Chronopappus (1), Chrysactinia (5), Chrysactinium (6), Chrysanthellum (13), Chrysanthemoides (2), Chrysanthemum (37), Chrysanthoglossum (2), Chrysocephalum (8), Chrysocoma (20), Chrysogonum (1), Chrysolaena (9), Chrysoma (1), Chrysophthalmum (3), Chrysopsis (11), Chrysothamnus (9), Chthonocephalus (6), Ciceronia (1), Cichorium (6), Cineraria (c. 16), Cirsium (c. 200), Cissampelopsis (10),

ASTERALES

EUDICOTS

Cladanthus (5), Cladochaeta (2), Clappia (1), Clibadium (29), Coleocoma (1), Coleostephus (3), Colobanthera (1), Cololobus (3), Columbiadoria (1), Comaclinium (1), Comborhiza (2), Commidendrum (4+1 extinct), Condylidium (2), Condylopodium (4), Conocliniopsis (1), Conoclinium (4), Constancea (1), Conyza (c. 30), Coreocarpus (9), Coreopsis (35), Corethamnium (1), Corethrogyne (1), Coronidium (9), Corymbium (9), Cosmos (26), Cota (40), Cotula (c. 50), Coulterella (1), Cousinia (c. 700), Cousiniopsis (1), Craspedia (23), Crassocephalum (24), Crassothonna (13), Cratystylis (3), Cremanthodium (70), Cremnothamnus (1), Crepidiastrum (7), Crepidifolium (4), Crepis (c. 200), Crinitaria (1), Criscianthus (1), Critonia (43), Critoniadelphus (2), Critoniella (6), Critoniopsis (28), Crocidium (1), Cronquistia (1), Cronquistianthus (29), Croptilon (3), Crossostephium (1), Crossothamnus (2), Crupina (3), Cuatrecasanthus (3), Cuatrecasasiella (2), Culcitium (6), Cullumia (15), Cuspidia (1), Curio (20), Cyanthillium (1), Cyathocline (3), Cyathomone (1), Cylindrocline (2), Cymbolaena (1), Cymbonotus (3), Cymbopappus (3), Cymophora (4), Cynara (8), Cyrtocymura (8), Dacryotrichia (1), Dahlia (35), Damnamenia (1), Damnxanthodium (1), Darwiniothamnus (3), Dasyandantha (1), Dasyanthina (2), Dasycondylus (8), Dauresia (1), Daveaua (1), Decachaeta (8), Decastylocarpus (1), Decazesia (1), Deinandra (21), Delairea (1), Delamerea (1), Delilia (2), Dendrocacalia (1), Dendrophorbium (c. 75), Dendrosenecio (4), Dendroseris (11), Denekia (1), Desmanthodium (7), Dewildemania (3), Diacranthera (2), Baccharis patagonica, Royal Horticultural Society Garden, Wisley, UK [438]

Dianthoseris (1), Diaphractanthus (1), Dicercoclados (1), Dichaetophora (1), Dichrocephala (3), Dichromochlamys (1), Dicomopsis (1), Dicoria (2), Dicranocarpus (1), Didelta (2), Dielitzia (1), Dieteria (3), Digitacalia (5), Dillandia (3), Dimeresia (1), Dimerostemma (12), Dimorphocoma (1), Dimorphotheca (9), Diodontium (1), Diplostephium (90), Dipterocome (1), Dipterocypsela (1), Disparago (9), Dissothrix (1), Distephanus (34), Disynaphia (16), Dithyrostegia (2), Dittrichia (2), Doellia (2), Doellingeria (8), Dolichoglottis (2), Dolichorrhiza (4), Dolichothrix (1), Dolomiaea (14), D orob a e a (3), D oronic u m (26 ), Dresslerothamnus (4), Dubautia (22), Dubyaea (10), Dugesia (1), Duhaldea (13), Dymondia (1), Dysaster (1), Dyscritothamnus (2), Dysodiopsis (1), Dyssodia (27), Eastwoodia (1), Eatonella (1), Echinacea (4), Echinocoryne (6), Echinops (c. 120), Eclipta (1), Edmondia (3), Egletes (6), Eirmocephala (3), Eitenia (2), Ekmania (1), Ekmaniopappus (2), Elachanthemum (2), Elachanthus (3), Elaphandra (13), Elekmania (9), Elephantopus (c. 15), Eleutheranthera (1), Ellenbergia (1), Elytropappus (10), Emilia (c. 90), Emiliella (5), Encelia (14), Enceliopsis (3), Endocellion (2), Endopappus (1), Engelmannia (1), Engleria (2), Enydra (10), Epaltes (14), Epilasia (3), Epitriche (1), Erato (5), Erechtites (c. 12), Eremanthus (18), Eremothamnus (1), Ericameria (36), Erigeron (c. 390), Eriocephalus (32), Eriochlamys (2), Eriophyllum (13), Eriotrix (2), Erlangea (6), Erodiophyllum (2), Erymophyllum (5), Eryngiophyllum (2), Erythradenia (1), Erythroseris (2), Espejoa (1), Espeletia Centaurea cyanus, Ruisalo Island, Turku, Finland [438]

(83), Ethulia (19), Eucephalus (10), Euchiton (17), Eumorphia (6), Eupatoriastrum (5), Eupatorina (1), Eupatoriopsis (1), Eupatorium (41), Euphrosyne (6), Eurybia (22), Euryops (97), Eutetras (2), Euthamia (5), Evacidium (1), Ewartia (4), Ewartiothamnus (1), Exomiocarpon (1), Exostigma (2), Facelis (3), Farfugium (3), Faujasia (4), Faujasiopsis (3), Faxonia (1), Feddea (1), Feldstonia (1), Felicia (85), Fenixia (1), Ferreyranthus (10), Ferreyrella (1), Filago (12), Filifolium (1), Fitchia (7), Fitzwillia (1), Flaveria (21), Fleischmannia (c. 80), Fleischmanniopsis (5), Florestina (8), Floscaldasia (2), Flosmutisia (1), Flourensia (18), Flyriella (4), Foveolina (5), Gaillardia (c. 16), Galactites (3), Galatella (c. 40), Galeana (3), Galeomma (2), Galinsoga (13), Gamochaeta (c. 50), Gamochaetopsis (2), Garberia (1), Garcibarrigoa (2), Gardnerina (1, possibly extinct), Garhadiolus (4), Garuleum (8), Gazania (16), Geigeria (20), Geissolepis (1), Geraea (2), Geropogon (1), Gibbaria (1), Gilberta (1), Gilruthia (1), Glebionis (2), Glossocardia (12), Glossopappus (1), Glyptopleura (2), Gnaphaliothamnus (10), Gnaphalium (c. 38), Gnephosis (8), Gnomophalium (1), Goldmanella (1), Gongrostylus (2), Goniocaulon (1), Gonospermum (5), Gonzalezia (3), Gorceixia (1), Gorteria (3), Goyazianthus (1), Grangea (10), Grangeopsis (1), Graphistylis (8), Gratwickia (1), Grauanthus (2), Grazielia (10), Greenmaniella (1), Grindelia (c. 30), Grisebachianthus (8), Grosvenoria (4), Guardiola (10), Guayania (6), Guevaria (4), Guizotia (6), Gundelia (2), Gundlachia (6), Gutenbergia (24), Gutierrezia (28), Guynesomia (1), Gymnanthemum (40),

Aster amellus, Helsinki Botanical Garden, Finland [438]

Plants of the World

605

ASTERALES Gymnarrhena (1), Gymnocondylus (1), Gymnocoronis (3), Gymnodiscus (2), Gymnolaena (3), Gymnopentzia (1), Gymnosperma (1), Gymnostephium (8), Gynoxys (c. 130), Gynura (c. 40), Gyptidium (2), Gyptis (7), Gyrodoma (1), Haastia (4), Haeckeria (3), Haegiela (1), Handelia (1), Haplocarpha (10), Haploësthes (3), Haplopappus (c. 70), Haptotrichion (2), Harleya (1), Harmonia (5), Harnackia (1), Hartwrightia (1), Hasteola (2), Hatschbachiella (2), Hazardia (13), Hebeclinium (20), Hedosyne (1), Hedypnois (3), Helenium (c. 32), Helianthella (9), Helianthus (51), Helichrysopsis (1), Helichrysum (c. 500), Heliocauta (1), Heliomeris (5), Heliopsis (18), Helminthotheca (4), Helogyne (8), Hemizonella (1), Hemizonia (1), Henricksonia (1), Heptanthus (7), Herderia (1), Herodotia (1), Herreranthus (1), Hertia (10), Hesperevax (3), Hesperomannia (4), Heteracia (2), Heteranthemis (1), Heterocoma (1), Heterocondylus (13), Heterocypsela (1), Heteroderis (1), Heterolepis (4), Heteromera (1), Heteromma (3), Heteroplexis (2), Heterorhachis (2), Heterosperma (7), Heterothalamus (8), Heterotheca (28), Hidalgoa (6), Hieracium (c. 90 + c. 1,000 microspecies), Hilliardia (1), Himalaiella (11), Hinterhubera (8), Hippia (5), Hippolytia (19), Hirpicium (13), Hispidella (1), Hoehneophytum (3), Hoffmannanthus (1), Hoffmanniella (1), Hofmeisteria (7), Holocarpha (4), Hololeion (3), Hololepis (2), Holoschkuhria (1), Holozonia (1), Homogyne (3), Hoplophyllum (2), Huberopappus (1), Hubertia (25), Hughesia (1), Hulsea (7), Hulteniella (1), Humbertacalia (8), Humeocline (1), Hyalochlamys (1), Hyalosperma (9), Hybridella (2), Hydroidea (1), Hydropectis (3), Hymenolepis (7), Hymenonema (2), Hymenopappus (11), Hymenostemma (1), Hymenostephium (c. 26), Hymenothrix (5), Hymenoxys (25), Hyoseris (5), Hypacanthium (3), Hypericophyllum (7), Hypochaeris (c. 60), Hysterionica (7), Hystrichophora (1, possibly extinct), Ichthyothere (18), Idiopappus (1), Idiothamnus (4), Ifloga (6), Iltisia (2), Imeria (2), Indocypraea (1), Inezia (2), Inula (c. 90), Inulanthera (10), Inuloides (1), Inulopsis (4), Io (1), Iocenes (1), Iodocephalus (2), Iodocephalopsis (2), Iogeton (1), Ionactis (5), Iostephane (4), Iotasperma (2), Iphiona (11), Iphionopsis

606

Christenhusz, Fay & Chase

EUDICOTS

(2), Iranecio (16), Irwinia (1), Ischnea (4), Ismelia (1), Isocarpha (5), Isocoma (16), Isoëtopsis (1), Isostigma (11), Iva (9), Ixeridium (13), Ixeris (20), Ixiochlamys (4), Ixiolaena (1), Ixodia (2), Jacmaia (5), Jaegeria (6), Jalcophila (3), Jaliscoa (3), Jamesianthus (1), Jaramilloa (2), Jasonia (1), Jaumea (2), Jefea (5), Jeffreya (2), Jeffreycia (5), Jensia (2), Jessea (4), Joseanthus (5), Jurinea (c. 200), Kalimeris (16), Karelinia (1), Karvandarina (1), Kaschgaria (2), Kaunia (14), Kemulariella (6), Keysseria (12), Kinghamia (5), Kingianthus (2), Kippistia (1), Klasea (c. 45), Kleinia (c. 40), Koanophyllon (c. 115), Koehneola (1), Koelpinia (5), Koyamasia (1), Krigia (7), Kyhosia (1), Kyrsteniopsis (4), Lachanodes (1), Lachnophyllum (2), Lachnorhiza (2), Lachnospermum (4), Lactuca (c. 130), Laennecia (18), Laestadia (6), Lagascea (8), Lagenocypsela (2), Lagenophora (14), Laggera (17), Lagophylla (4), Lagoseriopsis (1), Lamprocephalus (1), Lampropappus (3), Lamyropappus (1), Lamyropsis (8), Langebergia (1), Lantanopsis (3), Lapidia (1), Lapsana (1), Lapsanastrum (4), Lasianthaea (11), Lasiocephalus (10), Lasiolaena (6), Lasiopogon (8), Lasiospermum (4), Lasthenia (18), Launaea (54), Lawrencella (2), Layia (14), Lecocarpus (3), Lemooria (1), Leonis (1), Leontodon (c. 50), Leontopodium (59), Lepidaploa (116), Lepidesmia (1), Lepidolopha (9), Lepidolopsis (1), Lepidonia (7), Lepidophorum (1), Lepidophyllum (1), Lepidospartum (3), Lepidostephium (2), Leptinella (33), Leptocarpha (1), Leptoclinium (1), Leptorhynchos (10), Leptostelma (5), Leptotriche (12), Lescaillea (1), Lessingia (12), Lessingianthus (c. 110), Lettowia (1), Leucactinia (1), Leucanthemella (2), Leucanthemopsis (9), Leucanthemum (25), Leucoblepharis (1), Leucochrysum (5), Leucocyclus (2), Leucogenes (2), Leucophyta (1), Leucoptera (3), Leysera (4), Liabum (38), Liatris (37), Libinhania (13) Lidbeckia (3), Lifago (1), Ligularia (125), Ligulariopsis (1), Limbarda (1), Lindheimera (1), Linzia (10), Lipoblepharis (5), Lipochaeta (6), Litogyne (1), Litothamnus (2), Litrisa (1), Llerasia (14), Logfia (12), Lomatozona (4), Lonas (1), Lopholaena (19), Lordhowea (1), Lorentzianthus (1), Loricaria (20), Lourteigia (9), Lowryanthus (1), Loxothysanus (3), Lucilia (8), Luciliocline (5), Luina (2), Lundellianthus

(8), Lundinia (1), Lycapsus (1), Lychnophora (26), Lychnophoriopsis (4), Lygodesmia (5), Machaeranthera (30), Macowania (12), Macropodina (3), Macvaughiella (2), Madagaster (5), Madia (10), Mairia (3), Malacothrix (20), Malmeanthus (3), Malperia (1), Mantisalca (4), Manyonia (1), Marasmodes (4), Marshallia (9), Marshalljohnstonia (1), Matricaria (6), Mattfeldanthus (2), Mattfeldia (1), Matudina (1), Mauranthemum (4), Mausolea (1), Mecomischus (2), Melampodium (39), Melanodendron (1), Melanthera (35), Merrittia (1), Mesanthophora (1), Mesogramma (1), Metalasia (52), Mexerion (2), Mexianthus (1), Micractis (3), Microcephala (5), Microglossa (18), Microgyne (2), Microliabum (6), Micropsis (5), Micropus (4), Microseris (14), Microspermum (7), Mikania (c. 430), Mikaniopsis (c. 15), Milleria (2), Millotia (16), Minasia (6), Minuria (9), Miricacalia (1), Misbrookea (1), Miyamayomena (5), Mniodes (5), Monactis (9), Monarrhenus (2), Monoculus (2), Monogereion (1), Monolopia (5), Monoptilon (2), Montanoa (25), Monticalia (c. 70), Moonia (1), Moquinia (2), Moquiniastrum (24), Morithamnus (2), Msuata (1), Mtonia (1), Munnozia (43), Munzothamnus (1), Muschleria (1), Myanmaria (1), Myopordon (6), Myriactis (22), Myriocephalus (9), Myxopappus (2), Nabalus (15), Nananthea (1), Nannoglottis (8), Nanothamnus (1), Nardophyllum (5), Narvalina (4), Nelsonianthus (1), Nemosenecio (6), Neobrachyactis (3), Neocabreria (5), Neocuatrecasia (12), Neojeffreya (1), Neomirandea (27), Neonesomia (2), Neopallasia (3), Neotysonia (1), Nephrotheca (1), Nesampelos (3), Nesomia (1), Nestlera (1), Neurolaena (13), Neurolakis (1), Nicolasia (7), Nicolletia (3), Nidorella (15), Nikitinia (1), Nipponanthemum (1), Nivellea (1), Nolletia (10), Nordenstamia (20), Norlindhia (3), Nothobaccharis (1), Nothoschkuhria (1), Nothovernonia (2), Noticastrum (20), Notobasis (1), Notoseris (12), Novenia (1), Obliia (2), Ochrocephala (1), Oclemena (3), Odixia (2), Odontocline (6), Oedera (18), Oiospermum (1), Oldfeltia (1), Olearia (c. 180), Olgaea (16), Oligactis (12), Oliganthes (9), Oligocarpus (2), Oligochaeta (4), Oligothrix (1), Olivaea (2), Omphalopappus (1), Oncosiphon (8), Ondetia (1), Onopordum (c. 40), Oocephala (2), Oonopsis

ASTERALES

EUDICOTS

Actinobole uliginosum, near Oodnadatta, South Australia [438]

Ambrosia artemisiifolia, Belleville, Wisconsin, USA [438]

Polycalymma stuartii, South Australia [438]

Achillea millefolium, Hallinen, Turku, Finland [438]

Taraxacum officinale, Kingston upon Thames, UK [438]

Leontopodium alpinum, private garden, Kingston upon Thames, Surrey, UK [438]

Montanoa hexagona, San Francisco Botanical Garden, California, USA [438]

Zinnia acerosa, New Mexico, USA (DZ) [438]

Petasites fragrans, Lizard Peninsula, Cornwall, UK [438]

Plants of the World

607

ASTERALES

EUDICOTS

Taxonomic quarrels the circumscription of plant groups today, even when these result in a better overall understanding of plant evolution. It is, however, good that there are differences of opinion, which allow science to progress while being thoroughly scrutinised.

Sigesbeckia orientalis, Asteraceae, roadside weed on La Réunion

many ways. For example, Linnaeus named an insignificant, but troublesome, stinking weed Sigesbeckia, which is still known under that name today, whereas Siegesbeck used his powerful position in St Petersburg to block any Siberian and Russian plants and animals from reaching Linnaeus for his studies. Siegesbeck instead preserved these in his own collections; his herbarium is now kept in the Herzog August Library in Wolfenbüttel, Germany. When Linnaeus returned to Stockholm in 1748, he found that Siegesbeck’s mockery had some consequences. He had become the laughing stock of the Stockholm establishment and could not even find servants, let alone receive patients to be cured. Linnaeus did not partake in his defence, which was performed by Johann Browallius (1707– 1775), the later bishop of Turku, Finland, who defended the practicality of Linnaeus’ system, admitting that it is an artificial classification, bringing together distantly related plants in the same group. Disagreement in taxonomy is found in every age and any classifying discipline. We are bemused to see how strongly some plant scientists respond to necessary changes in

(4), Oparanthus (4), Ophryosporus (37), Opisthopappus (2), Orbivestus (10), Oreochrysum (1), Oreoleysera (1), Oreostemma (3), Oresbia (1), Oritrophium (21), Orochaenactis (1), Orthopappus (1), Osbertia (3), Osmadenia (1), Osmiopsis (1), Osmitopsis (9), Osteospermum (c. 65), Oteiza (3), Othonna (c. 130), Otopappus (14), Otospermum (1), Oxycarpha (1), Oxylobus (5), Oxypappus (2), Oyedaea (16), Ozothamnus (50), Pachystegia (3), Pachythamnus (1), Pacifigeron (1), Packera (c. 65), Pacourina (1), Palafoxia (12), Paleaepappus (1), Pallenis (3), Pappobolus (38), Papuacalia (17), Paquirea (1), Paracalia (2), Parafaujasia (2), Paragynoxys (15), Paranephelius (7), Parantennaria (1), Parapiqueria (1), Paraprenanthes (11), Parasenecio (>60, the former Cacalia of Asia), Parastrephia (3), Parasyncalathium (1), Parthenice (1), Parthenium (16), Pascalia (3), Paurolepis (1),

Pechuelloeschea (1), Pectis (c. 90), Pegolettia (9), Pelucha (1), Pentacalia (c. 200), Pentachaeta (7), Pentalepis (3), Pentanema (18), Pentatrichia (4), Pentzia (23), Pericallis (14), Pericome (2), Peripleura (9), Perityle (67), Perralderia (3), Perymenium (41), Petalacte (1), Petasites (19), Peteravenia (5), Petradoria (1), Petrobium (1), Peucephyllum (1), Phaenocoma (1), Phaeostigma (3), Phagnalon (43), Phalacraea (4), Phalacrocarpum (2), Phalacroseris (1), Phaneroglossa (1), Phanerostylis (5), Phania (2), Philactis (4), Philoglossa (5), Philyrophyllum (2), Phitosia (1), Phoebanthus (2), Phyllocephalum (3), Phymaspermum (18), Picnomon (1), Picris (c. 40), Picrosia (2), Picrothamnus (1), Pilbara (1), Pilosella (c. 20+60 microspecies), Pinaropappus (10), Pinillosia (1), Piora (1), Pippenalia (1), Piptocarpha (45), Piptocoma (18), Piptolepis (9), Piptothrix (5), Piqueria

(6), Piqueriella (1), Piqueriopsis (1), Pithecoseris (1), Pithocarpa (2), Pittocaulon (5), Pityopsis (7), Pladaroxylon (1), Plagiobasis (1), Plagiocheilus (7), Plagiolophus (1), Plagius (1), Planaltoa (2), Planea (1), Plateilema (1), Platycarpha (3), Platypodanthera (1), Platyschkuhria (1), Plecostachys (2), Plectocephalus (2), Pleiacanthus (1), Pleurocarpaea (2), Pleurocoronis (3), Pleuropappus (1), Pleurophyllum (3), Pluchea (c. 80), Podachaenium (2), Podanthus (2), Podocoma (8), Podolepis (20), Podotheca (6), Poecilolepis (2), Pogonolepis (2), Pojarkovia (1), Polyanthina (1), Polyarrhena (4), Polycalymma (1), Polychrysum (1), Polydora (8), Polymnia (3), Polytaxis (2), Porophyllum (c. 25), Porphyrostemma (4), Praxeliopsis (1), Praxelis (14), Prenanthes (26), Prestelia (1), Printzia (6), Prolobus (1), Prolongoa (1), Proteopsis (2), Psacaliopsis (5), Psacalium (42),

Johann Georg Siegesbeck (1686–1755) was a German physician and botanist who became professor of botany at the Russian Academy of Sciences in St Petersburg in 1735. He was an outspoken opponent of the Linnaean system of classification based on the reproductive organs of plants. He called it “such abominable lewdness in the Kingdom of Plants”, repugnant and immoral. Siegesbeck mocked Linnaeus’s idea that God would allow 20 men or even more (stamens) to have a single wife (pistil) in common, or that a wedded man would be allowed concubines (nearby female flowers). The animosity between Siegesbeck and Linnaeus was deep and expressed in

608

Christenhusz, Fay & Chase

Sigesbeckia in C. Linnaeus, Hortus Cliffortianus, ed. 1, tab XXIII. 1737. Illustrated by Jan Wandelaar

ASTERALES

EUDICOTS

Psathyrotes (3), Psathyrotopsis (3), Psednotrichia (3), Ps e p h e ll u s (9 0), Pseudelephantopus (2), Pseudobahia (3), Pseudoblepharispermum (2), Pseudobrickellia (2), Pseudoclappia (2), Pseudoconyza (1), Pseudognaphalium (84), Pseudogynoxys (12), Pseudohandelia (1), Pseudonoseris (3), Pseudopiptocarpha (2), Pseudostifftia (1), Pseudoyoungia (8), Psiadia (60), Psiadiella (1), Psilactis (6), Psilocarphus (5), Psilostrophe (7), Psychrogeton (20), Pterachaenia (1), Pterocaulon (17), Pterochaeta (1), Pterocypsela (11), Pteronia (70), Pterygopappus (1), Ptilostemon (14), Pulicaria (76), Pulicarioidea (1), Pycnosorus (6), Pyrrhopappus (5), Pyrrocoma (14), Pytinocarpa (2), Quadribractea (1), Quechualia (4), Quinetia (1), Quinqueremulus (1), Rachelia (1), Radlkoferotoma (3), Rafinesquia (2), Raillardella (3), Rainiera (1), Raoulia (26), Raouliopsis (2), Rastrophyllum (2), Ratibida (7), Raulinoreitzia (3), Rayjacksonia (3), Reichardia (8), Relhania (13), Remya (3), Rennera (4), Rensonia (1), Revealia (1), Rhagadiolus (2), Rhamphogyne (2), Rhanteriopsis (4), Rhanterium (3), Rhaponticoides (17), Rhaponticum (23), Rhetinocarpha (1), Rhetinolepis (1), Rhinactinidia (2), Rhodanthe (46), Rhodanthemum (12), Rhodogeron (1), Rhynchopsidium (2), Rhynchospermum (1), Rhysolepis (3), Richteria (3), Riencourtia (6), Rigiopappus (1), Robinsonecio (2), Roebuckiella (8), Rochonia (4), Rojasianthe (1), Rolandra (1), Roldana (c. 55), Roodebergia (1), Rosenia (4), Rothmaleria (1), Rudbeckia (23), Rugelia (1), Rumfordia (12), Russowia (1), Rutidosis (6), Sabazia (17), Sachsia (3), Salmea (10), Santolina (c. 10), Santosia (1), Sanvitalia (5), Sarcanthemum (1), Sartorina (1), Sartwellia (3), Saussurea (c. 400), Scalesia (15), Scherya (1), Schischkinia (1), Schistocarpha (10), Schistostephium (13), Schizogyne (2), Schizopsera (1), Schizotrichia (5), Schkuhria (2), Schmalhausenia (1), Schoenia (5), Sciadocephala (5), Sclerocarpus (8), Sclerolepis (1), Sclerorhachis (4), Scolymus (3), Scorzonera (c. 175), Scrobicaria (2), Selloa (3), Semiria (1), Senecio (c. 1,633), Sericocarpus (5), Serratula (c. 35), Shafera (1), Shangwua (3), Sheareria (2), Shinnersia (1), Shinnersoseris (1), Siapaea (1), Siebera (2), Sigesbeckia (12), Siloxerus (4), Silphium

(12), Silybum (2), Simsia (20), Sinacalia (4), Sinclairia (23), Sinosenecio (36), Sipolisia (1), Smallanthus (c. 20), Soaresia (1), Solanecio (16), Solenogyne (3), Solidago (116), Soliva (8), Sommerfeltia (1), Sonchella (2), Sonchus (62), Sondottia (2), Soroseris (7), Spaniopappus (5), Sparganophoros (1), Sphaeranthus (40), Sphaereupatorium (2), Sphaeromeria (9), Sphagneticola (4), Spilanthes (6), Spiracantha (1), Spiroseris (1), Squamopappus (1), Stachycephalum (2), Staehelina (5), Standleyanthus (1), Staurochlamys (1), Steiractinia (12), Steirodiscus (6), Stenachaenium (5), Stenocephalum (5), Stenocline (3), Stenophalium (4), Stenops (2), Stenoseris (5), Stenotus (4), Stephanodoria (1), Stephanomeria (15), Stevia (c. 240), Steviopsis (10), Steyermarkina (4), Stilpnogyne (1), Stilpnolepis (2), Stilpnopappus (24), Stizolophus (3), Stoebe (25), Stokesia (1), Stomatanthes (17), Stramentopappus (1), Streptoglossa (8), Strotheria (1), Stuartina (2), Stuckertiella (2), Stuessya (3), Stylocline (7), Stylotrichium (4), Symphyllocarpus (1), Symphyopappus (12), Symphyotrichum (98), Syncalathium (3), Syncarpha (c. 30), Syncephalum (5), Syncretocarpus (2), Synedrella (1), Synedrellopsis (1), Syneilesis (7), Synotis (50), Syntrichopappus (2), Synurus (1), Syreitschikovia (2), Tagetes (52), Talamancalia (4), Tamananthus (1), Tamaulipa (1), Tanacetopsis (21), Tanacetum (163), Taplinia (1), Taraxacum (c. 34 + c. 2,000 microspecies), Tarlmounia (1), Tehuana (1), Teixeiranthus (2), Telanthophora (14), Telekia (1), Telmatophila (1), Tenrhynea (1), Tephroseris (50), Tessaria (3), Tetrachyron (7), Tetradymia (10), Tetragonotheca (4), Tetramolopium (38), Tetraneuris (9), Tetranthus (4), Tetraperone (1), Thaminophyllum (3), Thamnoseris (1), Thelesperma (14), Thespidium (1), Thespis (3), Thevenotia (2), Thiseltonia (2), Thurovia (1), Thymophylla (13), Thymopsis (2), Tibetoseris (1), Tietkensia (1), Tilesia (3), Tithonia (11), Tolpis (13), Tomentaurum (1), Tonestus (4), Tourneuxia (1), Townsendia (27), Tracyina (1), Tragopogon (c. 110), Traversia (1), Trep a d oni a (2), Tr ic h a n th e mi s (9), Trichanthodium (4), Trichocoronis (3), Trichocoryne (1), Trichogonia (30), Trichogoniopsis (2), Trichogyne (9), Tricholepis (18),

Trichoptilium (1), Trichospira (1), Tridactylina (1), Tridax (22), Trigonopterum (1), Trigonospermum (5), Trilisa (1), Trioncinia (1), Tripleurospermum (38), Triplocephalum (1), Tripolium (2), Tripteris (20), Triptilodiscus (1), Troglophyton (6), Tuberculocarpus (1), Tuberostylis (2), Tugarinovia (1), Turaniphytum (2), Tussilago (1), Tuxtla (1), Tyrimnus (1), Ugamia (1), Uleophytum (1), Unxia (3), Urbananthus (2), Urbinella (1), Urolepis (1), Urospermum (2), Ursinia (39), Varilla (2), Varthemia (1), Vellereophyton (7), Venegasia (1), Verbesina (c. 300), Vernonanthura (65), Vernonia (c. 780), Vernoniastrum (12), Vernoniopsis (1), Vieraea (1), Viereckia (1), Vigethia (1), Viguiera (c. 150), Villanova (10), Villasenoria (1), Vinicia (1), Vittadinia (20), Vittetia (2), Vogtia (2), Volutaria (16), Waitzia (5), Wamalchitamia (5), Warionia (1), Wedelia (c. 100), Welwitschiella (1), Werneria (c. 25), Westoniella (6), Wilkesia (1), Willemetia (2), Wollastonia (1), Wyethia (8), Xanthisma (17), Xanthium (3), Xanthocephalum (6), Xenophyllum (21), Xeranthemum (6), Xerochrysum (5), Xerxes (1), Xiphochaeta (1), Xylanthemum (9), Xylorhiza (10), Yariguianthus (1), Yermo (1), Youngia (37), Zaluzania (10), Zemisia (1), Zexmenia (2), Zinnia (17), Zoegea (3), Zyrphelis (13), Zyzyura (1) and Zyzyxia (1). Uses: The great diversity and widespread occurrence of this family has resulted in many economic uses. Of course, we cannot list here the great variety of these, but we aim to at least mention the main species used as food, drink, condiments, oil, dye, timber, cosmetics, hallucinogens, medicines, cut flowers and ornamental plants. Salads and vegetables: let t uce (Lactuca sativa) was derived in the eastern Mediterranean from prickly lettuce (L. serriola), out of which the prickles and bitterness had been selected in ancient times. Lettuce was already a popular salad green in ancient Greece and Rome, where many selections were grown, and it is now the main salad green used worldwide, especially in temperate regions. It can also be cooked into soup. Golden samphire (Limbarda crithmoides) has succulent leaves that are sometimes eaten as a salad green. Milfoil

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609

ASTERALES (Achillea millefolium) and sneezewort (A. ptarmica) are effective insect repellents, and herbal treatments for arthritis but are sometimes also eaten as a salad plant. Young shoots and leaves of milk thistles, especially sowthistle (Sonchus oleraceus) are eaten locally as a vegetable. In mythology, Theseus ate sowthistle before he went into the labyrinth to kill the Minotaur. A peculiar salad green, recently becoming more popular is Brazilian cress or electric buttons (Acmella oleracea). This small herb has an ‘electric’ taste because it contains spilanthol, which gives a tingling sensation on the tongue. Leaves of dandelion (or pis-en-lit, Taraxacum officinale) are also sometimes eaten as salad greens, but this has to be done in moderation because they are mildly diuretic. Marigold flowers (Tagetes erecta and cultivars) make decorative and flavourful additions to salads as well. Their common name, African marigold, belies their origin in Central America. Leaves of corn marigold (Glebionis segetum) are sometimes added to salads in Greece. Sea aster (Tripolium pannonicum, formerly Aster tripolium) has young leaves that are picked from mudflats in Zeeland (the Netherlands), where they are eaten as an exclusive vegetable called lamsoren (lamb’s ears). In Greece, a variety of common brighteyes (Reichardia picroides), known there as galatsida, is eaten in salads, but more often boiled, steamed or cooked in olive oil. Several species are popular in Asian cuisines, especially Glebionis coronaria, which is usually known as chrysanthemum greens (shungiku, tong ho) and is common in Cantonese, Japanese and Taiwanese dishes. The leaf stalks of fuki (Petasites japonicus) are eaten as rhubarb or as a vegetable in Japanese cuisine; they have to be soaked in ash or baking soda first to remove their astringency, but fuki contains carcinogens and is better avoided. Okinawa spinach (Gynura bicolor) is stir-fried with sesame oil and ginger in China. Gallant soldiers (Galinsoga parviflora) is sometimes cooked as a vegetable in Asia, but it is originally from South America and has become a troublesome garden weed worldwide.

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The leaves of cardoon (Cynara cardunculus var. cardunculus) are blanched and eaten like chard. It is not as popular as the immature f lower heads of globe artichoke (C. cardunculus var. scolymus), which are cultivated on a large scale in the Mediterranean and California. The bases of the involucral bracts and receptacle of the flower-head are eaten, and the florets (called the ‘choke’) are discarded. Of a similar taste, the stems of Scotch thistle (Carduus nutans) have a thick pith that can be eaten when boiled, similar to the leaf bases and stems of swamp thistle (Cirsium palustre). The thistle is the emblem of Scotland, although this was most likely the spear thistle (C. vulgare) and not the exotic cotton thistle (Onopordum acanthium), which now usually receives this honour. Holy thistle (Silybum marianum) has edible leaves, stems and roots that can be boiled and buttered, and the inflorescences were eaten like globe artichokes. One of the bitter herbs of Passover is endive (Cichorium endivia), which, like lettuce, has been used as a vegetable since ancient times. It is slightly bitter and can be eaten fresh in salads or cooked in other dishes. There are three cultivar groups, friseé (Friseé Group), escarole (Scarole Group) and endive (Endivia Group). Witloof or true chicory (C. intybus) has also been cultivated for millennia and there are four main cultivar groups: Root Chicory, Pain de Sucre, Radicchio and Witloof, the last usually blanched. Chicory and endive are eaten fresh in salads or cooked or baked in various ways. The root chicories were roasted and used as a coffee substitute, especially during times of scarce coffee, such as during the Napoleonic wars. When coffee again became available, people often continued to add chicory root to coffee because they were used to the taste, and coffee remained expensive. These chicory supplements are still commercially available, but have lost popularity because coffee now comes in a great variety of flavours, and chicory is no longer needed. Roots of a number of other plants were also used to replace coffee (e.g. Scolymus hispanicus, Scorzonera hispanica, Taraxacum officinale). The seeds of Silybum marianum (kenguel

seeds) were sometimes roasted and used to replace coffee. Several Asteraceae are used to flavour other drinks. Wormwood (Artemisia absinthium) contains thujone, a compound with a similar effect on the body as cannabis. An alcoholic drink flavoured with fennel, anise and wormwood is called absinthe, which originated in Switzerland in the late 18th century and became popular in the late 19th and early 20th centuries. It was associated with the Bohemian lifestyle and was opposed by social conservatives. The psychoactive effects of absinthe have been exaggerated, thujone being present in the drink only in trace amounts, but the drink was villified and banned in 1915. A revival of absinthe in the 1990s has now resulted in it becoming a legal and fairly popular drink again, although it is not as popular as it was a century ago. Southernwood (A. abrotanum) has a fragrance that is more pleasant than that of absinthe, and it is sometimes used to make tea or as a flavouring for herbal bitters. Russian wormwood (A. pontica) is sometimes used to flavour vermouth, and alpine wormwood (A. glacialis) is found as a flavour in génepi liqueur. Burdock (Arctium lappa) has roots that are called gobō and eaten as a vegetable in Japan; it is reputed to be an aphrodisiac, and the young leaves are sometimes eaten in countries around the Arctic as a salad. Together with fermented dandelion (Taraxacum officinale) root, burdock has also been used since the Middle Ages in the UK to brew a drink. It originally was lightly alcoholic, but “dandelion and burdock” is now usually a type of soft drink. The unopened flower heads of dandelion can be pickled (like capers), and when opened they can be cooked into a syrup, good as cordial, in cakes or to make wine. Yarrow (Achillea erbarotta subsp. moschata) is used in liqueurs and other drinks and formerly was used in herbal medicine. The roots of pellitory (Anacyclus pyrethrum), medicinally called radix pyrethri, are used in mouthwashes and liqueurs and as a food flavouring, and alecost or camphor plant (Tanacetum balsamita) was formerly used to flavour ale.

ASTERALES

EUDICOTS

Native to Paraguay, caa-ehe (Stevia rebaudiana) has been used by the local people as a sweetener for centuries. It contains stevioside, which is 300 times sweeter than sucrose, and stevia is now much used as a natural sugarfree sweetener in soft drinks, chewing gum and confectionery, especially in Japan and the Nordic countries. It is usually propagated by cuttings because its seeds are difficult to germinate. Many Asteraceae make tasty root vegetables. Best known is perhaps topinambour or Jerusalem artichoke (Helianthus tuberosus), which has sweet tubers due to the high content of inulin. Inulin is a sweet compound that humans cannot digest, and in some people it can cause severe flatulence (hence the vulgar name ‘fartichokes’). These North American sunflower relatives were not introduced to England via Israel, as their name might seem to imply, but rather Terneuzen, the Netherlands, which was corrupted into Jerusalem, although the name is also said to have come from the Italian word for sunflower, girasole. Whatever the origin of the name, these roots were eaten by native Americans long before European contact and quickly adopted into the diet by early settlers. Helianthus tuberosus arrived in Europe in 1613 and naturalises easily from cultivation, especially along streams, where it sometimes forms large stands. Similar to Jerusalem artichoke, yacon are the edible tubers of Smallanthus sonchifolius, which also contain inulin. Eaten as a vegetable, mostly in Latin America, it is also used there to make alcohol. Dahlia roots have also been eaten in the Americas since ancient times, especially Dahlia coccinea, and species cultivars with large roots were selected for consumption. Commercial inulin (also called diabetic sugar) is harvested from Dahlia species. Horse-heal or elecampane (Inula helenium) has thick roots that have a camphorous floral scent and bitter taste. These plants, cultivated for their roots since ancient times in Europe, were used for food and medicine. It is also an alternative flavouring in absinthe. Scorzonera or black salsify (Scorzonera hispanica) roots are boiled, peeled and eaten as a vegetable. Roots of cardillo or Spanish oyster (Scolymus

hispanicus) and salsify or vegetable oyster (Tragopogon porrifolius) can be eaten in a similar way. The young flower shoots are sometimes cooked as chard. In Portugal a sweet is made from crystallised scorzonera roots. Yam daisy or murnong (Microseris lanceolata) has (after roasting) edible tubers, and these were once an important food source for Australian aboriginals. The plant suffered dramatically from the introduction of cattle, sheep and goats into Australia, resulting in food shortages for local people. The roots are said to taste sweet with a hint of coconut. Sunf lowers (Helianthus annuus) are commercially grown on a large scale for their seeds that yield a commonly used culinary oil. Sunflower oil is used in cooking, for deep frying, to make margarine and industrially for paint, lacquer, lubrication etc. The seeds are also roasted and salted to be eaten as a snack. Birds also love the seeds, and these are popular in bird feeders, but the seeds have allelopathic compounds that may make other plants wither so care must be taken. Another important oil seed is niger or ramtil (Guizotia abyssinica). Especially grown for oil used in local cuisine in India, it is elsewhere often given as a food to caged birds or on birdfood tables together with sunflower seeds, in Europe particularly to attract goldfinches (Carduelis carduelis). Oil from the seeds of lettuce species (Lactuca serriola, L. sativa) is also used in cooking. The seeds of goldfields (Lasthenia glabrata) can be ground and cooked into a porridge. Tarweed (Madia elegans) seeds were ground into flower, and it was a staple for native Americans. Madia sativa is grown for its seed oil in Chile, which is used as a substitute for olive oil. Because the leaves of many species are fragrant some can be used as herbs or spices. Tarragon (Artemisia dracunculus) and Russian tarragon (A. dracunculoides) are well-known kitchen herbs, eaten commonly with fish and chicken. Yomogi or Japanese mugwort (A. princeps) is added as a flavour to dumplings in Japan, and common mugwort (A. vulgaris) is used as a condiment. It is said to aid digestion and relieve depression. Many kitchen herbs also have medicinal properties, and best known for that is Roman chamomile (Chamaemelum nobile), which produces a

light blue oil used in liqueurs, shampoos, hair oil and other cosmetics. Its leaves and flower heads are used as a tea, but excessive use is associated with bladder cancer. Wild chamomile (Matricaria recutita) is also often used for this purpose, especially in colder climates. Steaming with chamomile may relieve colds and sinusitis, and chamomile oil can be used to treat psoriasis and boils. As limoncillo, the flowers of Pectis papposa are sold in the markets of Mexico to be used as a meat flavouring, and holy flax (Santolina rosmarinifolia) is used as a flavouring in the Algarve. Oil of lavender cotton (S. chamaecyparissus) is used in perfumery, and stems can be placed in the closet to scent clothes. There are numerous medicinal properties associated with Asteraceae, although many also contain toxic compounds, so care has to be taken when using pure herbs. We enumerate a few here that are perhaps best known. Artemisin is found in high concentrations in quinghao (Artemisia annua), a species from China that is used in traditional medicine to successfully treat malaria. It is now applied on a larger scale (usually via synthesised analogues that may not be as effective), but the available amount and application of artimisin remains limited. Arnica (Arnica montana) has a flavonoid compound (astragalin) that works as an antihistamine and is now promoted for jetlag. Scotch or pot marigold (Calendula officinalis) has long been used medicinally against warts etc. but is now mostly used to decorate salads, thicken soups, colour butter and in cosmetics. It is a cultigen of unknown origin, possibly derived from the hybridisation of two wild species. The rhizomes of purple cone flower plants (Echinacea purpurea and E. pallida var. angustifolia) have antiviral properties, and use stimulates the production of white blood cells. They are often used as temporary relief for cold symptoms, and as a food supplement they have now become an industry worth millions of dollars. Some studies demonstrate a lack of efficacy. Blessed thistle (Centaurea benedicta, formerly known as Cnicus benedictus) contains cnicin, a glucocide that used to be an important gout treatment. Its seeds are also a source of

Plants of the World

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ASTERALES

EUDICOTS

Artemisia annua, Chelsea Physic Garden, London, UK [438] Arctium lappa, Helsinki Botanical Garden, Finland [438]

oil. Oil from the curry plant (Helichrysum italicum) has antiviral properties. Sambong or ngai camphor (Blumea balsamifera) relieves symptoms of cold. ‘Radix helenae’ is the root of Inula helenium, used since ancient times in Europe to treat skin and chest diseases, now commonly used to treat asthma. Gumweed (Grindelia squarrosa) is used to cure dermatitis and burns. Dried heads of feverfew (Tanacetum parthenium) are effective in the treatment of migraine. Tansy (T. vulgare) deters insects, worms and flies and was used as a vermifuge. It also is an ingredient of drinsheen, an Irish blood-sausage. Petasine, a terpene found in butterbur (Petasites hybridus), is a strong anticonvulsive (better than papaverine), and that is the reason this plant is now present throughout Europe. Its leaves were also used to pack butter. Yellow fleabane (Dittrichia viscosa) was used for bone-setting and as an anti-inflammatory in Ancient Greece. Marigolds (Tagetes) were used medicinally by the Aztecs for tick removal. They have compounds that in modern medicine are effective in treating immune deficiency diseases. ‘Stinking roger’ (T. minuta) is also a good insecticide and can be used to protect crops against nematodes. Pyrethrine, an insecticide commonly used against plant pests, is harvested from Tanacetum cinerariifolium, commonly wild-collected or cultivated in Morocco and the uplands of East Africa for this compound. Thepelakano, also known as the leaves-of-god and dream herb (Calea ternifolia), contains caleicines

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and caleochromines that may induce sleep, and it is used in divination rituals in Mexico. Innocent-looking Mexican tarragon (Tagetes lucida) is reputed to be an hallucinogen, although leaves have a tarragon-like flavour with a hint of anise and can be used safely to flavour food. It was used by the Aztecs to flavour chocolate and ground with other herbs into a powder used to stupify victims of human sacrifice. It seems unlikely that this plant has much stupifying effect though, and its inclusion may have been symbolic. Apart from chamomile, echinacea and marigolds, several other species are used in cosmetics. Worth highlighting is North American mountain tail-leaf (Pericome caudata), which is used in cosmetic hairloss products. Pollen of golden leastdaisy (Pentachaeta aurea) has been used in make-up, and tincture of coltsfoot (Tussilago farfara) was an ingredient of some commercial hairsprays. Surprisingly edelweiss (Leontopodium nivale), national flower of Austria and Switzerland, is used in skin whitening creams, although this is unlikely to have any true effect on the skin. Many species yield good dyes, the best known are safflower (Carthamus tinctorius), which has flowers that were formerly used as a food and cosmetic dye. It is now sometimes used to adulterate saffron (Crocus sativus, Iridaceae), having a similar colour but not taste. As ‘kurdee’ it is known as an oil-seed crop, and this oil is frequently used in aromatherapy. Similarly, the stamens of cotton thistle (Onopordum acanthium) were sometimes used to adulterate

Carthamus tinctorius, Ruissalo Botanical Garden, Turku, Finland [438]

saffron, as were the flowers of spanish salsify (Scolymus hispanicus). An orange dye was formerly made from Coreopsis tinctoria, which also was used to make a type of hot drink. Golden chamomile (Cota tinctoria, formerly known as Anthemis tinctoria) yields a well-known yellow dye used for centuries to dye wool and cotton. Saw-wort (Serratula tinctoria) is mixed with alum to give a green or yellow dye used for wool. False daisy (Eclipta prostrata) produces a black dye used to colour hair or as a natural ink for tattoos. Several tree-like species produce good timber, such as Blepharispermum, Brachylaena, Erismanthus, Fulcaldea laurifolia, Montanoa quadrangularis, Olearia, Vernonia arborea and many others locally, often used for charcoal, fuel wood, paper making, hard wood for construction and termite-proof timber for flooring etc. Camphor wood (Tarchonanthus camphoratus) is valued to make musical instruments, but it is fragrant and also yields essential oils used as a scent by the Maasai in East Africa. In the 1920s leaf blight ruined the Brazilian rubber crop and as an alternative guayule (Parthenium fruticosum) was investigated to produce commercial rubber, which was again needed when Japan cut off the Malaysian rubber trade during World War II. The rubber of guayule is hypoallergenic and thus safer for rubber gloves than Hevea rubber. Synthetic rubber is now preferred for most uses. Asteraceae have long been popular cut flowers, most popular being chrysanthemums (Chrysanthemum ×morifolium, C. indicum,

ASTERALES

EUDICOTS

C. zawadskii and hybrids), which are sold in flower stands around the world, probably only surpassed in popularity by roses. Chrysanthemums were bred in various forms in ancient China and have remained popular ever since. Hybrid gerberas (Gerbera jamesonii × G. viridifolia) are also popular cut flowers and grown commercially for this purpose in greenhouses on a large scale. Solidaster (Solidago ×luteus, a spontaneous hybrid between S. canadensis and S. ptarmicoides) emerged in a nursery in Lyon and is now a popular cut flower grown especially for this purpose on a large scale in the Netherlands. Helianthus, Helenium, Liatris and many others are used as cut flowers on smaller scales. Paper daisies and everlastings are popular for dried flower arrangements because the colourful bracts keep their colour perfectly when dried. These are mostly species of Chrysocephalum, Coronidium, Helichrysum, Podolepis, Rhodanthe, Syn-

Chamaemelum nobilis, Royal Botanic Gardens, Kew, UK [438]

carpha, Xeranthemum and Xerochrysum. Of course, with this diversity of plants, hardiness and reliability of flowering, many Asteraceae have become popular garden ornamentals. As bedding plants the most popular are Ageratum houstonianum (bluemink), Arctotis (African daisy), Argyranthemum frutescens (bush marguerites), Bellis perennis (double forms; English daisy), Bidens aurea (bur-marigold), Brachyscome iberidifolia (Swan River daisy), Calendula officinalis (marigold), Callistephus chinensis (Chinese aster), Chrysanthemum ×morifolium (garden chrysanthemum or ‘mums’), Cosmos bipinnatus (garden cosmea), Dahlia ×pinnata (garden dahlia), Dimorphotheca sinuata (sun marigold), Erigeron (fleabane), Felicia amelloides (blue daisy) and F. bergiana (kingfisher daisy), Gerbera (gerbera), Gazania (treasure flower), Glebionis coronaria (crown daisy), Layia (tidy-tips), Osteospermum, Pericallis (florist’s cinerarias), Rudbeckia hirta (annual

Cichorium intybus, Ronda, Spain [438]

black-eyed Susan), Sanvitalia procumbens (Mexican creeping zinnia), Tagetes erecta (African and French marigolds), Tanacetum coccineum (garden pyrethrum), Zinnia elegans, Z. haageana and Z. violacea (zinnia). As perennial border and alpine plants, there are cultivars of Achillea (yarrow), Ajania pacifica (Pacific chrysanthemum), Amberboa moschata (sweet sultan), Amellus asteroides (African aster), Anacyclus (pellitory), Anaphalis (pearly everlasting), Antennaria (cat’s foot), Anthemis (dog-fennel), Arctanthemum arcticum (Arctic daisy), Aster (mountain asters), Baccharis (groundsel bush), Berlandiera lyrata (chocolate flower), Berkheya purpurea (purple berkheya), Boltonia asteroides (doll’s daisy), Buphthalmum (European ox-eye), Catananche caerulea (Cupid’s dart), Celmisia (New Zealand cotton daisy), Centaurea and Psephellus (bluebottle, knapweed, cornflower), Chrysogonum virginianum (goldenstar), Chrysopsis (golden aster), Cirsium

Helianthus annuus, Ruissalo Island, Turku, Finland [438]

Cynara cardunculus var. scolymus, Kensington Gardens, London, UK [438] Echinacea purpurea, Beth Chatto Gardens, UK [438]

Plants of the World

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ASTERALES

614

EUDICOTS

Scalesia affinis, Galápagos Islands [438]

Darwiniothamnus tenuifolius, Isabela, Galápagos Islands [438]

Commidendrum rugosum, Royal Botanic Gardens, Kew, UK [438]

(thistle), Coreopsis (tickseed), Cosmos atrosanguineus (chocolate cosmea), Cota (dyer’s chamomile), Cotula (golden buttons), Craspedia uniflora (billy buttons), Dahlia excelsa (tree-dahlia), Doronicum (leopard’s bane), Echinacea (cone flower), Echinops (globe thistles), Eupatorium (boneset, hemp-agrimony, thoroughwort), Gaillardia (blanket flowers), Helenium (sneezeweed), Helianthella (small sunflower), Helianthus (sunflowers), Helichrysum italicum (curry plant), H. petiolare (liquorice plant), Heliopsis (American oxe-eye), Inula (spikenard), Isocoma (goldenbush), Kalimeris (Korean aster), Leontopodium nivale subsp. alpinum (edelweiss), Leptinella squalida (brass buttons), Leucanthemella serotina (autumn oxeeye), Leucanthemum hybrids (shasta daisy), L. vulgare (ox-eye daisy or marguerite), Liatris spicata (gayfeather), Ligularia (golden ray, leopard plant), Lindheimera texana (Texas star), Marshallia (Barbara’s buttons), Nipponanthemum nipponicum (Nippon daisy), Olearia (daisy bush), Othonna capensis (Cape ragwort), Ozothamnus (alpine everlasting, kerosine bush), Petasites fragrans (winter heliotrope), Pilosella aurantiaca (fox and cubs), Piqueria trinervia (tabardillo, winter stevia), Raoulia australis (New Zealand scab plant), Ratibida columnifera (yellow cone flower), Rudbeckia (black-eyed susan), Verbesina (crown-beard), Vernonia (iron weed), Wyethia (mule’s ears), Senecio and Jacobaea (ragwort), Sinacalia tangutica (goldstaff), Solidago (goldenrod), Stokesia laevis (Stoke’s aster), Symphyotrichum (autumn asters), especially S.

ericoides (white aster), S. novae-angliae (New England aster), S. novibelgii (New York aster) and S. oblongifolium (aromatic aster) and their cultvars, Syneilesis palmata (Japanese umbrella plant), Telekia speciosa (heart-leaves oxe-eye), Tithonia diversifolia (tree marigold) and many others. As houseplants or outside in tropical countries one may also find Bartlettina sordida (blue mist flower), Cassinia, Curio rowleyanus (string-of-pearls), Delairea odorata (Cape ivy), Farfugium japonicum (leopard plant), Gynura procumbens ‘Aurantiaca’ (purple velvet plant), Kleinia (succulent senecios), Mikania (hempvine), Pseudogynoxys chenopodioides (formerly Senecio confusus, Mexican flame vine) etc. Their popularity as ornamental plants and the ease of escape thanks to wind-dispersed seeds have resulted in the escape of many species into the wild with severe consequences in some cases. This is especially problematic when the plants cause bad allergic reactions in humans due to their pollen (particularly hogbrake or ragweed, Ambrosia artemisiifolia and mugworts, Artemisia) or dermatitis upon contact, such as Mexican devil (Ageratina adenophora), Singapore daisy (Sphagneticola trilobata) and Parthenium hysterophorus. In addition the pollen of P. hysterophorus can make other flowers sterile when it accidentally gets transferred to their stigma, which happens frequently because insects are rarely completely selective. This may cause native species to have a poor seed set and allow an invasive to dominate the vegetation.

The ‘Darwin’s finches of botany’: Because woodiness has often been seen as a primitive character in Asteraceae and woodiness of predominantly herbaceous lineages is common on islands, it has been assumed that shrubby island species of Asteraceae are relics of a ‘primitive’ vegetation that has otherwise become extinct on the mainland. The Canary Islands have often served as a good example with their woody Sonchus species. Other examples are the eight shrubby Senecio species from the Juan Fernández Islands (formerly placed in the genus Robinsonia, named for Robinson Crusoe), which have their closest relatives among Senecio species on mainland South America. Also on the Californian Channel Islands one can find woody Asteraceae, Munzothamnus blairii and Coreopsis gigantea, with their closest relatives being herbaceous members on the American continent. It seems more likely that they evolved woodiness secondarily on islands rather than this being a primitive state. Woody Asteraceae have radiated in many places, including the Atlantic island of St Helena, where gumwoods (Commidendrum) radiated into five species (one extinct); there are other endemic woody species on St Helena, such as she cabbage (Lachanodes arborea), black cabbage (Melanodendron integrifolium) and he cabbage (Pladaroxylon leucadendron). The last has its closest relatives in South America and Australasia, whereas another woody species, whitewood (Petrobium arboreum), is most closely

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related to species from the Pacific. Such long-distance dispersal events are relatively common in Asteraceae. All woody Asteraceae have in the past been used for construction on St Helena, and most are now severely threatened or extinct. Other widely disjunct taxa are found on Lord Howe Island (between Australia and New Zealand), where Lordhowea insularis has its closest relative in South Africa (Phaneroglossa bolusii). Moonia heterophylla from Sri Lanka is most closely related to Dahlia, a Neotropical genus, and Pacific genera Oparanthus and Fitchia are related to certain Caribbean genera. These disjunctions are probably due to the

long-distance dispersals made possible by the evolution of pappus (and thus wind dispersal), although similar disjunctions are known in species with seeds decidedly poorly adapted for dispersal. Once they reached the island or archipelago, radiation is likely as there are many unfilled ecological niches. Good examples of such adaptive radiations are the Hawaiian silverswords (Argyroxiphium; related to North American Madia) and of course tree-like Scalesia (related to herbaceous Helianthus) that radiated on the Galápagos Islands in a manner similar to the adaptive radiation that made Darwin’s finches famous.

Etymology: Aster is Latin for star, referring to the flower heads. Compositae (and its derivative ‘composite’ in English) is an alternative name, based on the flower heads that are composed of many flowers, but this is not based on a genus name and is thus better avoided. Invention: Burdock fruits have stiff hooks, with which they cling to fur of animals and so disperse themselves. A fruiting head of Arctium lappa got stuck in the fur of a dog owned by a Swiss engeneer, George de Mestral, which inspired him to develop velcro in 1945.

ESCALLONIALES This heterogeneous order has an estimated age of c. 110 million years, possibly older. It now consists of the single family Escalloniaceae.

Anopterus glandulosus, Singapore (WA) [439]

Eremosyne pectinata, Western Australia (DM) [439]

439. ESCALLONIACEAE Currybush family

Escallonia alpina, Royal Botanic Gardens, Kew, UK [439]

Escallonia laevis, National Botanic Gardens of Ireland, Glasnevin [439]

These evergreen shrubs and trees, sometimes pachycaul (Anopterus), rarely annual herbs (Eremosyne), have simple, alternate leaves (rarely opposite in Polyosma), occasionally with prickles along the stem. Leaves are often leathery with a spicy scent (Escallonia). Blades are usually pinnately veined (rarely palmate), and margins are toothed or lobed and often glandular. Inflorescences are bracteate terminal racemes or cymes or the f lowers axillary or terminal and solitary (Tribeles). Flowers are bisexual and actinomorphic. The usually five sepals are Plants of the World

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ESCALLONIALES fused (rarely free), and the same number of petals are usually free (rarely fused), often forming a cup- or urn-shaped perianth. Stamens are usually five and free from each other and the petals, but sometimes an additional whorl of staminodes is present. Anthers are basifixed and open inwardly by lengthwise slits. There usually is a nectar disk inside the stamen whorl. The inferior ovary (or nearly superior in Anopterus) is composed of usually two to four fused carpels topped by a single style and stigma. The fruit is a capsule, berry or drupe. Escallonia plants often have a smell reminiscent of curry, but in some species this is only detectable in dried material (e.g. herbarium specimens). Distr ibut ion: Escalloniaceae occu r in t ropical and temperate Cent ral and South America, on Réunion, in temperate and tropical East Asia from the eastern Himalayas and southern

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China to northeastern Australia and New Zealand. Phylogeny and evolution: Escalloniaceae are the sole family in Escalloniales, and they form a heterogeneous group of plants that have racemose inflorescences and free petals and stamens in common. The family was previously included in Saxifragaceae or Grossulariaceae based on similarities in floral structure, but molecular studies placed them in their own family and order distant from Saxifragales, and although the order is not placed with certainty, it appears to be close to Asterales and Aquifoliales. The family evolved over 110 million years ago, and therefore some of its distribution may be explained by continental drift and extinction in Antarctica, but the occurrence on the volcanic oceanic island of La Réunion indicates that long-distance dispersal is also not uncommon. Tribeles, Polyosma and

Eremosyne were previously placed in their own families, but they are strongly supported in molecular studies, with Escallonia. Forgesia and Valdivia are sometimes included in Escallonia. Genera and species: The family has seven genera and 103 species: Anopterus (2), Eremosyne (1), Escallonia (37), Forgesia (1), Polyosma (60), Tribeles (1) and Valdivia (1). Uses: Escallonia rubra and various cultivars and hybrids are grown as ornamentals, frequently as a hedge plant in the UK and Ireland. Anopterus macleayanus, a pachycaul with long leaves, is sometimes grown in curiosity collections. Etymology: Escallonia is named for Spanish explorer and plant hunter Antonio Escallón (born 1738), who discovered E. myrtilloides in 1773 in Colombia.

BRUNIALES Families 440 and 441 form the order Bruniales, a clade that is thought to be c. 110 million years old and composed of two families, one restricted to South Africa and the other South America, this disjunction most likely the result of long-distance dispersal.

440. COLUMELLIACEAE Andean-holly family

Shrubs and trees with evergreen, simple, opposite leaves without stipules make up this family. Leaf blades have pinnate venation, and margins are prickly toothed (Desfontainia) or entire and sometimes glandular, then with the blade asymmetrical (Columellia). 616

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Inf lorescences are few-f lowered axillary or terminal cymes or the flowers solitary. Flowers are bisexual and actinomorphic or slightly zygomorphic. A hypanthium is present to which the usually five (rarely four to eight) sepals are fused. The same number of petals are fused into a five- (rarely fourto eight-)lobed tube that is entirely yellow or the tube is red and lobes yellow. The two (Columellia) or five (Desfontainia) stamens alternate with the petal lobes. Anthers are fused to a broad connective, the large pollen sacs with undulate margins, folded and twisted (in Columellia), the filaments fused to the corolla throat and anthers basifixed and opening by lateral slits in Desfontainia. The ovary is inferior and composed of two carpels forming a single locule in Columellia

or the ovary is superior and composed of three to seven (usually five) fused carpels in Desfontainia. The ovary has a terminal short thick style and a two- or four-lobed broad stigma. Fruits are dry dehiscent (septicidal or valvate) capsules (Columellia) or fleshy, yellowish-white berries (Desfontainia). Distribution: The family is found in the Andes of South America from Colombia to southern Chile and in the paramó of Costa Rica (Central America). Phylogeny and evolution: The two genera differ in many characters and have been placed in their own families for that reason in the past. Desfontainia was formerly placed in Loganiaceae, but wood anatomy, molecular

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Desfontainia spinosa in fruit, private garden, Kingston upon Thames, Surrey, UK [440]

Desfontainia spinosa, private garden Kingston upon Thames, Surrey, UK [440]

Brunia nodiflora, South Africa (Wikipedia Commons) [441]

Desfontainia steyermarkii, Ecuador [440]

Berzelia abrotanoides, South Africa (JA) [441]

phylogenetics and palynology have placed this genus close to Columellia. Together they are sister to Bruniaceae.

Etymology: Columellia honours Lucius Junius Moderatus Columella (c. 4–70 AD), a Roman agricultural writer.

Genera and species: This family consists of two genera and three to seven species: Columellia (2–4) and Desfontainia (1–3). Some authors reduce the four or five species of Columellia to two and others accept three species in Desfontainia. A molecular and morphological study of this group would be appropriate.

441. BRUNIACEAE

Uses: Chilean holly (Desfontainia spinosa) is sometimes cultivated as an ornamental in suitably mild climates. Tea from its leaves is allegedly hallucinogenic, but this has not been confirmed.

Buttonbush family

These ericoid shrubs (rarely trees) have closely set, spirally arranged (usually in

Columellia oblonga, Ecuador (MW) [440]

Berzelia lanuginosa, Royal Botanic Gardens, Kew, UK [441]

five ranks), simple, needle-like or scale-like leaves with entire margins and often a black leaf tip. Leaves often have minute stipules, long unicellular hairs and short or absent petioles, and venation is parallel, usually with three, but sometimes with five or up to 20 parallel veins. Inflorescences are usually bracteate heads, sometimes spikes or the flowers solitary, rarely in racemes. Flowers are bisexual, actinomorphic and usually subtended by two bracteoles. The (four or) five sepals are free or fused, persistent in fruit or reduced. The (four or) five petals are usually free (rarely fused in some Brunia), often basally clawed. Stamens are as many as and alternate with the petals, usually free or the filaments occasionally fused to the petal claws forming a tube. Anthers are Plants of the World

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BRUNIALES dorsifixed and open inwardly by lengthwise slits, sometimes with a noticeably sterile tip (connective). The ovary is inferior or halfinferior (rarely superior) and composed of one or two (or three in Audouinia) carpels, each forming a locule. Each carpel has a terminal style that is free or only basally fused, each terminated by one, two or three stigmas. Fruits are achenes (if there is a single carpel) or a capsule, schizocarp or nutlet. Distribution: This family is restricted to the Republic of South Africa, with the greatest diversity in the Cape Province and only one

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species (Brunia trigyna) found along the coast of KwaZulu-Natal. Phylogeny and evolution: The family is estimated to have emerged c. 59–99 million years ago, but much of the diversification occurred during the last 3–18 million years. There are some strikingly similar fossils from the Upper Cretaceous in Sweden. Genera and species: The family has six genera and c. 81 species: Audouinia (5), Berzelia (15), Brunia (c. 40), Linconia (3), Staavia (11) and Thamnea (7).

Uses: Species of Berzelia and Brunia are cultivated as ornamentals and cut flowers often called ‘Cape greens’. They make good everlastings in dried arrangements. Etymology: Brunia is probably named for Dutch traveller and artist Cornelis de Bruijn (1652–1727), who made several expeditions to the Middle East, Russia and Java (via the Cape). His illustrations of the Great Pyramid and Jerusalem were the first images of these places known in Western Europe.

PARACRYPHIALES This order evolved c. 100 million years ago and had a much wider distribution in the past, the current diversity being relictual. Formerly three families were accepted, which are now merged in Paracryphiaceae. The genera are closely related but morphologically divergent.

442. PARACRYPHIACEAE Possumwood family

This is a family of shrubs and trees with simple, leathery, alternate leaves, often in false whorls along the stem. They lack stipules and have blades with pinnate venation and entire to finely serrate margins. Inf lorescences are axillary or terminal racemes or spikes, sometimes in panicles. The bisexual, actinomorphic flowers are sessile or shortly stalked, in Paracryphia male flowers are found in the lower part of the inflorescence. The usually four to five (to 12) sepals are free, the petals absent in Paracryphia and some Sphenostemon with four or five free petals. The five (Quintinia) or eight (rarely to 11) stamens have free filaments and basifixed anthers that open by 618

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lengthwise slits. The superior (Paracryphia) or inferior (Quintinia, Sphenostemon) ovary is composed of two (Sphenostemon), three to five (Quintinia) or eight to 15 (Paracryphia) fused carpels topped with as many sessile stigmas as there are carpels. Fruits are capsules in Quintinia and Paracryphia, a drupe or berry in Sphenostemon. In Paracryphia the capsule opens from the base, and the valves are attached to a central column at the tip.

correct, the current Australasian distribution is relictual. The three genera have often each been placed in their own families that were, previous to molecular results, not usually associated with each other: Paracryphiaceae in Theales, Quintiniaceae in Saxifragales and Sphenostemonaceae in Celastrales or Sphenostemon included in Aquifoliaceae, Icacinaceae or Trimeniaceae. Sphenostemon is particularly poorly known.

Distribution: This family is found along the south eastern fringe of the Pacific in the Philippines, Sulawesi, Seram, New Guinea, New Caledonia, eastern Australia and New Zealand.

Genera and species: Paracryphiaceae consist of three genera with 36 species: Paracryphia alticola, Quintinia (25) and Sphenostemon (10). The greatest diversity occurs in New Caledonia where all three genera and 13 species can be found.

Phylogeny and evolution: Apart from tetramerous flowers and septicidal capsules, this family is polymorphic and placed in their own order among the asterids. They are estimated to have evolved c. 100 million years ago. A number of fossils are known from the Northern Hemisphere, most notably Silvianthemum suecicum and Bertilanthus scanicus in Sweden from the Late Cretaceous (83.5 million years ago), which resemble Quintinia to some extent. If this relationship is

Uses: Possumwood or opossum wood is the hard, pinkish, close-grained wood of Quintinia sieberi, which is occasionally harvested for timber in Australia. Etymology: Paracryphia is composed of the Greek words παρά (para), merely, and κρυφίως (kryfios), secret, referring to its umbrella-like fruits that resembles those of Medusagyne (Ochnaceae).

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Quintinia parviflora, New Caledonia (JM) [442]

Paracryphia alticola, Panié, New Caledonia (JM) [442]

Sphenostemon pachycladus, New Caledonia (TW) [442] Paracryphia alticola, Panié, New Caledonia (JM) [442] Quintinia sieberi, showing leaf domatia, Beaumont, New South Wales, Australia [442]

DIPSACALES Two closely related families, 443 and 444, compose the order Dipsacales. Formerly this clade was more highly divided, with Dipsacaceae, Morinaceae and Valerianaceae usually maintained as separate. In order to prevent the recognition of several small families, an expanded Caprifoliaceae was favoured in APG III (2009), but the mostly shrubby genera Sambucus and Viburnum were moved to the formerly exclusively herbaceous Adoxaceae, blurring the lines between these families somewhat. However, Adoxaceae can be recognised by having actinomorphic flowers and three to five (nearly) sessile stigmas, whereas Caprifoliaceae have usually zygomorphic flowers and one to three stigmas on a single elongate style. The split between these families may have occurred during the Mid Cretaceous (102–111 million years ago). The order probably originated in the Northern Hemisphere, but their diversification is relatively recent (perhaps just 10 million years ago).

443. ADOXACEAE Elder family

Adoxaceae are shrubs, sometimes tree-like, and rhizomatous perennial herbs. Their

leaves are opposite (rarely whorled), simple or pinnately or twice-ternately compound, usually long-petiolate and with or without stipules. Blades or pinnae have pinnate or palmate venation and entire or toothed to lobed margins. Inflorescences are terminal panicles, true or false umbels, spikes (Tetradoxa) or head-like cymes (appearing like a cube with four on each side and one on top in Adoxa). Flowers are usually bisexual and actinomorphic, and sometimes have different numbers of parts in the same inflorescence; in some species with (pseudo-) umbellate inflorescences the marginal flowers

are zygomorphic and enlarged (some species of Viburnum). The (three to) five sepals are fused and so are the four or five petals that often have a nectary at their base. The four or five stamens are adnate to and alternate with the petals. An inner whorl of four or five staminodes is sometimes present. Anthers are basifixed and open inwardly by lengthwise slits. The inferior or half-inferior ovary is composed of two to five fused carpels that form one or several locules. Each carpel has a free terminal style, or the style is absent and three to five stigmas are sessile. Fruits are fleshy indehiscent capsules or berries. Plants of the World

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Adoxa moschatellina, private garden, Kingston upon Thames, Surrey, UK [443]

Sambucus ebulus, Helsinki Botanical Garden, Finland [443]

Sambucus nigra, Sandbjerg, Denmark [443]

Sambucus nigra, Wavrons, France [443]

Viburnum davidii, Royal Botanic Gardens, Kew, UK [443]

Viburnum plicatum, Ruissalo Botanical Garden, Turku, Finland [443]

Distribution: This family has a mostly Northern Hemisphere distribution. It is found throughout North America, Central America and Andean and Atlantic southeastern South America. They occur in the Macaronesian Islands and throughout Europe, northwestern Africa, western Asia, Russia, Siberia and throughout temperate and subtropical Asia and tropical Southeast Asia. They have outliers in the mountains of East Africa (rare), southeastern Australia and Tasmania. Phylogeny and evolution: The placement of herbaceous Adoxa, Sinadoxa and Tetradoxa has been problematic in the past, although a relationship with woody Sambucus and Viburnum in Caprifoliaceae had been suggested early on, based on morphological characters. This was confirmed by molecular studies, not at all close to Cornales, 620

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with which they were often previously associated (Viburnum has occasionally been misidentified as Cornaceae). Adoxaceae are estimated to have diverged some 70–85 million years ago, and they diversified some 30–60 million years ago. Some deep splits are known between tropical and temperate clades, with Malesian Viburnum clemensiae sister to the rest of the mostly temperate genus. There has been a suggestion that this family should be renamed Viburnaceae, which may become its name in the future, again unnecessarily destabilising the nomenclature of this family. Genera and species: This is a family of five genera and c. 225 species: Adoxa (1), Sambucus (10), Sinadoxa (1), Tetradoxa (1) and Viburnum (c. 210).

Uses: Elder (Sambucus nigra) has deliciously scented flowers that are made into a cordial and other drinks or fermented into a fizzy drink, occasionally eaten in pancakes or other baked goods and confectionery. Elderflower water was previously used as a cosmetic skin and eye cleanser. Berries can also be eaten but are inferior to the American elder (S. canadensis), which is grown for its edible fruit that is delicious in jams, jellies, sauces, juice, drinks and wine. Danewort (S. ebulus) was previously used medicinally against dropsy and as a blue dye for leather. Several Viburnum species also have edible berries, e.g. wild raisin (V. cassinoides), squashberry (V. edule), nannyberry (V. lentago), Guelder rose (V. opulus), blackhaw (V. prunifolium) and American cranberry bush (V. trilobum), but some species have poisonous fruit (e.g. V. darwinii, V. tinus). The young leaves of V.

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setigerum are used as a tea substitute locally in China. The wood of Sambucus nigra is used to make the mouth piece of a ‘midwinterhoorn’, a traditional musical instrument from Twente (eastern Netherlands), where it is still played during yuletide. The young wood of Sambucus and Viburnum is soft and pliable, whereas the old wood is tough and hard, giving it numerous local applications. Many species of Viburnum and some Sambucus are grown as garden ornamentals for their decorative flowers, fruits, leaf shapes and autumn colour. Adoxa moschatellina (moschatel) is occasionally planted in woodland gardens as a curiosity. Its alternative name, ‘townhall clock’, refers to the four fiveparted flowers, facing north, south, east and west, and the terminal four-parted flower facing upwards. Elder magic: Gather some pulled teeth, some hair and fresh fingernails and bury these at sunrise between the roots of an elder bush so that your home and body will be protected from the influences of the devil. This is only one of many stories related to the beneficial magical powers of an elder bush. One of the best known is probably the story of Frau Hulda, one of the Grimm fairytales. In the Grimm version, she lived in a well, but being

Lonicera caprifolium, Romania [444]

the queen of elves and ruling over life and death, her house was originally in an elder bush; these were often planted near a well because the scent of the leaves repels flies (and the devil), which is useful if you want the water in your well to remain pure. That Hulda lived there with her cats, shaking pillows to make it snow on earth, and deciding about life and death, is something that one has to take for granted. However, always pick some flowers of elder at midsummer and keep these in case a devil wanders by. That you can make a fabulous drink from the flowers is an extra bonus! Why do the fruits of Sambucus taste bitter? Before Judas hanged himself on an elder tree, the fruits tasted delicious and sweet, but after his act they turned bitter. Judas also is said to have cut off his ear, and this is why elder bushes sometimes have a jelly fungus growing from their wood that closely resembles a human ear. It is called Auricularia auricula-judae and is edible, even though in Europe it is only used in folk medicine, whereas in China it is popular as an ingredient in many recipes. Etymology: Adoxa is composed of the Greek words α (a), without, and δόξα (doxa), glory, probably in reference to the diminutive stature of this plant. Lonicera caerulea var. edulis, Ruissalo Botanical Garden, Turku, Finland [444]

444. CAPRIFOLIACEAE Honeysuckle family

These are shrubs, trees, woody vines and perennial, annual and biennial herbs. Leaves are simple, pinnatifid or pinnately compound and opposite or in whorls along the stem, sometimes the bases fused to each other across the stem (perfoliate), sometimes petiolate or sheathing, stipules usually absent (but present in Leycesteria). Leaf blades are entire, toothed or lobed, and they usually have pinnate venation, sometimes with several main veins emerging from the base or the venation palmate (Morina); leaves on vegetative stems are often different in shape from those of flowering stems (and these may be interpreted as bracts, the distinction between bracts and leaves not always being clear). Inflorescences are bracteate simple to compound cymes and heads, the flowers paired or solitary. Involucral bracts are

Lonicera pileata in fruit, Royal Botanic Gardens, Kew, UK [444]

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DIPSACALES common. Flowers are usually zygomorphic, rarely actinomorphic, bisexual or functionally unisexual. The (two to) four or five sepals are usually fused and persistent. The (three to) five petals are fused into a lobed corolla that is sometimes spurred. The one to five stamens are fused to the corolla tube (two staminodes are sometimes present in Morina), sometimes two of the anthers longer (didynamous stamens). Anthers are dorsifixed and open by lengthwise slits. The usually inferior (sometimes part-inferior) ovary is composed of five (or two to eight) fused carpels with a single terminal style and a one- to fivelobed capitate or truncate, wet stigma. Fruits are achenes, cypselas, capsules, berries or drupes. Distribution: Caprifoliaceae are widespread and found on most continents (except Australia, New Zealand and Antarctica), but they are most diverse in the temperate zones of the Northern Hemisphere. Phylogeny and evolution: Dipsacaceae and Valerianaceae, two well-known and wellestablished families, were found in molecular studies to be embedded in a paraphyletic Caprifoliaceae, which were then split to produce several small families, such as Diervillaceae, Morinaceae and Linnaeaceae, in order to maintain Dipsacaceae and Valerianaceae. These families share many characters, and the position of Heptacodium, which may well be of hybrid orgin between Caprifolioideae and Linnaeoideae, causes issues in accepting smaller families in this clade. A larger family concept was therefore accepted, excluding Sambucus and Viburnum, which were placed in Adoxaceae, and Silvianthus, which is placed in Carlemanniaceae (Lamiales). Linnaea has been expanded to include all Linnaeoideae (the former genera Abelia, Diabelia, Dipelta, Kolkwitzia and Vesalea), except Zabelia, which falls in an unresolved position close to Morina, and Heptacodium is tentatively placed in Caprifolioideae. An age estimate for Caprifoliaceae has varied between 35–86 million years. Genera and species: Caprifoliaceae include 28 genera and over 825 species in five subfamilies: Diervilloideae – Diervilla (3) and Weigela 622

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Twin teapots — Carolus Linnaeus (1707–1778) Famed for his epic works describing all plants and animals of the world and simplifying nomenclature by allowing only two words to name a species, Carolus Linnaeus or Carl von Linné is probably one of the best-known biologists of all time. He was controversial in his lifetime, establishing a classification of plants based on their sexual parts. He explained his ’sexual system’ with the numbers of ’men’ and ’women’ that shared a (flower-) bed and arranged them on this basis. These were matters that were frowned upon during the time of the enlightenment, but as a consequence his lectures were popular with students! He described all of God’s creations (plants in 1753, animals in 1758, dates considered as the start of modern nomenclature) and organised them in an artificial way based on their reproductive organs; useful for identification purposes, but it did place species together that were otherwise obviously unrelated. This is why new systems were created fairly soon afterwards, aiming to be ʻmore natural’, although the Linnaean system prevailed for many decades. The twin f lower, Linnaea borealis, was named by Linnaeus’ friend Gronovius because Linnaeus Painting of Carolus Linnaeus in the traditional liked this plant so much. Linnaeus dress of the Saami people of Lapland, holding the twin flower, Linnaea borealis, by Hendrik himself had already named it RudHollander (1853) after an original by Martin beckia, after his professor, but he Hoffman (1737) at the Hartekamp adopted the name Linnaea for it in his Species Plantarum (using Rudbeckia for a genus of Asteraceae), and it soon became his emblem. In the house of Linnaeus in Uppsala, now a museum, there are two china tea sets decorated with Linnaea borealis. Linnaeus had these commissioned in Canton in the 1760s. The first survived the journey from China to Sweden, but some pieces broke on the trip from Gothenburg to Uppsala. It was also ‘killed’ because the flowers were too red and not accurately drawn. Therefore Linnaeus ordered a second set (costing a small fortune), that arrived intact, and in which the flowers are more accurate and pink instead of red. Photo of the tea service with twin flowers (wikimedia, open source)

DIPSACALES

EUDICOTS

Weigela middendorffiana, Helsinki Botanical Garden, Finland [444]

Diervilla rivularis, Ylioppilaskylä, Turku, Finland [444]

Heptacodium miconioides, Royal Botanic Gardens, Kew, UK [444]

Linnaea spathulata, Royal Botanic Gardens, Kew, UK [444]

Linnaea borealis, Hallinen, Turku, Finland [444]

Morina nepalensis, Yunnan, China [444]

(10); Caprifolioideae – Heptacodium (1), Leycesteria (6), Lonicera (c. 180), Symphoricarpos (17) and Triosteum (6); Linnaeoideae – Linnaea (>23), Morina (13) and Zabelia (4); Dipsacoideae – Bassecoia (3), Cephalaria (c. 65), Dipsacus (15), Knautia (c. 60), Pterocephalus (30), Scabiosa (c. 80), Succisa (1), Succisella (4) and Triplostegia (2); Valerianoideae – Aligera (15), Centranthus (9), Fedia (3), Nardostachys (1), Patrinia (17), Plectritis (4), Pseudobetckea (1), Valeriana (c. 200) and Valerianella (c. 50). Uses: Lamb’s lettuce or mache (Valerianella locusta) is often grown as a pot herb and for salads. Similarly horn-of-plenty (Fedia cornucopiae) is eaten as a salad plant. Honeyberry (Lonicera coerulea var. edulis)

is cultivated for its berries that taste somewhat like blueberries. In the Balkans, the starchy seeds of Morina persica are eaten like rice. The dried rhizome of valerian (Valeriana officinalis) has been used since ancient times as a nerve relaxant and sedative. It has a rancid smell attractive to mammals such as cats and rats (it contains actinidine, see Actinidiaceae) and may have been the scent used by the Pied Piper of Hamelin, a Mediaeval story popularised by the brothers Grimm, in which the Piper lures rats (and later children, when the good citizens do not pay him) away from the city. Ethereal oil from the roots of Indian nard or spikenard (Nardostachys jatamansi) was highly priced in Roman society and is now sometimes used in the perfume industry

Triosteum himalaicum, Yunnan, China [444]

Dipsacus fullonum, Turku, Finland [444]

or medicinally (a relaxant, like valerian). Teasel heads (dried inf lorescences of Dipsacus fullonum or its cultivated cousin D. sativus) were formerly used to raise the nap of wool fabrics, now largely replaced by metal cards (a cognate with Carduus, a thistle; similarly teasel is a cognate with thistle). The heads are sometimes used in dried flower arrangements, and the UK National Trust places them on historical chairs to prevent visitors from sitting on them. The scent of honeysuckle (Lonicera caprifolium), woodbine (L. periclymenum) and Japanese honeysuckle (L. japonica) are sometimes used in perfumes and shampoos. They are also ornamentals popular for their highly scented flowers. A number of other species are also common garden ornamentals, Plants of the World

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Succisella petteri, private garden, Kingston upon Thames, Surrey, UK [444]

Scabiosa prolifera in fruit, Qana, Lebanon [444]

Fedia cornucopiae, Sicily, Italy [444]

Patrinia gibbosa, Royal Botanic Gardens, Kew, UK

Centranthus ruber, Aberystwyth, Wales, UK

Valerianella locusta, private allotment, Kingston upon Thames, Surrey, UK [444]

Valeriana officinalis, Mackinac Island, Michigan, USA [444]

Valeriana rigida, Ecuador [444]

Knautia dispacifolia, Helsinki Botanical Garden, Finland [444]

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[444]

[444]

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especially red valerian (Centranthus ruber), giant scabious (Cephalaria gigantea), bush honeysuckle (Diervilla lonicera), seven sons plant (Heptacodium miconioides), widow flower (Knautia, especially K. macedonica), Himalayan honeysuckle (Leycesteria formosana), twin flower (Linnaea species), especially garden abelia (L. ×grandiflora), rosy dipelta (L. dipelta), beauty bush (L. amabilis), Mexican abelia (L. floribunda), honeysuckles (Lonicera species), morina (Morina longifolia), Patrinia triloba, scabious (Scabiosa, especially S. atropurpurea), blue

buttons or Devil’s bit (Succisa pratensis), snowberry (Symphoricarpos albus), tinker’s weed (Triosteum species), Valeriana and Weigela species (especially W. florida). Carnivory: The lower leaves of the common teasel (Dipsacus fullonum) are united around the stem, forming cup-shaped structures that fill with water after rain. All sorts of debris and insects get trapped in these, but no evidence of digestive enzymes or foliar nutrient absorption has been found. It was shown that fluid in the leaf sheaths has a

lower surface tension, which could be an adaptation to kill prey, but the benefit has not been shown. Other animals come and visit the ‘pitchers’ to drink and eat trapped prey, so they can form micro-environments in the somewhat dryish environments where teasel sometimes grows. Etymology: Caprifolium is composed of the Latin capra, a goat, and folium, a leaf. It is an old name for Lonicera, in turn named after Adam Lonitzer (1528–1586), a German botanist and physician.

APIALES Families 445 to 451 make up the order Apiales, which accounts for c. 2.4% of eudicot diversity. The umbels, Apiaceae (or the alternate name Umbelliferae), were the subject of the first botanical monograph (Morison, 1672) and are among the families with the longest period of recognition. Their characteristic inflorescence structure makes many members of this order easy to recognise, although not all families have this type of inflorescence. Even though this is the last order in the linear sequence of the APG classification, it does not mean it is the most highly evolved. The clades of Apiales and Dipsacales are of equal size and could easily be swapped, the position here as last is merely convention.

445. PENNANTIACEAE Kaikomako family

These are unisexual or bisexual shrubs and trees with buttressed trunks, rarely vines. They have alternate, petiolate leaves without stipules, placed spirally along zigzagging branchlets. Leaf blades have pinnate venation and entire or toothed margins. Inflorescences are terminal or caulif lorous panicles or cymes, or the flowers are solitary. Flowers are unisexual and actinomorphic. The five sepals are fused into a tube (appearing absent in

Pennantia cunninghamii), and the five petals are free or fused and white. Male flowers have five stamens that alternate with the petals and are free or fused to them. Anthers are dorsifixed and open by lengthwise slits. A pistillode (sterile ovary) is sometimes present. Female flowers sometimes have five staminodes surrounding the superior ovary, which is composed of two or three carpels fused to form a single locule (rarely two). One or two styles are sometimes present and topped by a trilobed or disk-like stigma (but the stigma sometimes sessile). Fruits are fleshy or dryish, black drupes with a single trigonous seed. Distribution: This small family is found in tropical and subtropical rainforests of eastern Australia, Norfolk Island, New Zealand, each area with a single species, and a possible fourth on Three Kings Island, north of New Zealand.

Phylogeny and evolution: Pennantia was previously included in Icacinaceae, but molecular evidence placed it as sister to all other Apiales. Genera and species: The sole genus in this family is Pennantia with three, possibly four, species. Uses: Pennantia corymbosa (kaikomako) and P. cunninghamii (brown beech) have hard timber used for cabinetry and tool handles. The other two species are of conservation concern. Etymology: Pennantia is named for Thomas Pennant (1726–1798), a Welsh zoologist, naturalist, writer, traveller and antiquarian. He is most famous for his A tour in Scotland and voyage to the Hebrides (1774–1776), Synopsis of quadrupeds (1771) and Account of London (1790), promoting early tourism.

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APIALES

Pennantia baylisiana in fruit, Wellington Botanical Garden, New Zealand (JC) [445]

Aralidium pinnatifidum, Singapore Botanical Garden [446]

EUDICOTS

Pennantia corymbosa, Dunedin Botanical Garden, New Zealand (JC) [445]

Aralidium pinnatifidum, Singapore Botanical Garden [446]

Aralidium pinnatifidum, flowers, Singapore Botanical Gardens [446]

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Pennantia cunninghamii, Melbourne Botanical Garden, Victoria, Australia (JC) [445]

Aralidium pinnatifidum, Singapore Botanical Garden [446]

Torricellia angulata var. intermedia, Yunnan, Torricellia angulata var, intermedia, China (BY) [446] Kunming Botanical Garden, China [446]

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446. TORRICELLIACEAE Ivy-palm family

Trees and shrubs with simple, alternate, petiolate leaves that sheath at the base and lack stipules make up this family. Blades have pinnate (Aralidium, Melanophylla) or palmate (Torricellia) venation, and the margins are entire, serrate, toothed, lobed or dissected. Inflorescences are terminal panicles or racemes or pendulous thyrses. Flowers are functionally unisexual and actinomorphic. The (three to) five sepals are basally fused, and the three to five petals are fused (absent in female flowers of Torricellia). The five stamens alternate with the petals and are free from them, with basifixed, rarely dorsifixed, anthers that open by lateral slits. The inferior ovary is composed of two to four fused carpels with three, sessile, persistent terminal stigmas (Torricellia) or with two to four free styles each with a stigma (Aralidium, Melanophylla). Fruits are drupes with a single seed and sometimes with two empty chambers (Melanophylla).

Griselinia jodinifolia, National Botanic Gardens of Ireland, Glasnevin [447]

Distribution: Torricellia is found in warm temperate parts of the eastern Himalayas and western China (Yunnan), Aralidium occurs in western Malesia (southern Thailand, Peninsular Malaysia, Singapore, Sumatra, Anambas, Borneo) and Melanophylla is endemic to northeastern Madagascar. Phylogeny and evolution: The family is estimated to be at least 56 million years old, and Torricellia is known to have been widespread in the Northern Hemisphere, being known from Eocene deposits in Europe and northwestern North America. The members of this family were difficult to place because they share characters of both Araliaceae and Cornaceae, and many authors placed them in their own families. Molecular and morphological studies have shown that the three are closely related, and they are now treated in a single family. Aralidium is sister to the rest. Genera and species: This family consists of three genera and 11 species: Aralidium pinnatifidum, Melanophylla (7) and Torricellia (3). Uses: The leaves of Aralidium pinnatifidum repel insects, and the wood is sometimes used for flooring. Etymology: The name Torricellia honours Italian physicist and mathematician Evangelista Torricelli (1608–1647), who is best known for inventing the barometer.

Griselinia littoralis, Lizard Peninsula, Cornwall, UK [447]

447. GRISELINIACEAE Kapuka family

These are unisexual, ter restrial and epiphytic shrubs, vines and trees. Leaves are alternate and simple, petiolate and without stipules, but the base of the petiole is slightly sheathing, leaving prominent leaf scars on the stems. Blades are leathery and have three prominent veins emerging from the base and entire, dentate or spinose margins, the tip sometimes minutely trifid. Inflorescences are axillary racemes or panicles, and flowers are unisexual and actinomorphic. Male flowers have a minute calyx with five teeth and five free petals. Stamens are five, alternating with the petals and surrounding a fleshy disk. Anthers are dorsifixed and open by lateral slits. Female flowers have five short, fused sepals and lack petals and stamens. The inferior ovary is composed of three carpels fused to form one or two locules, only one fertile. The ovary has three free terminal styles, each bearing a capitate stigma. The fruits are blackish drupes with a single seed.

Griselinia racemosa, Royal Botanic Gardens, Kew, UK [447]

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APIALES Distribution: A disjunct family found in southern South America (Chile, Argentina, southeastern Brazil, Uruguay) and New Zealand. Phylogeny and evolution: In the past Griselinia was often placed in Cornaceae or Garryaceae, but it differs from those families in many characters. Molecular studies place it in Apiales. An age of 33.9 million years for this family has been suggested, excluding the possibility of it being a member of the Antarctic flora. It has more likely to have achieved its current distribution by long-distance dispersal. Fossil evidence is wanting.

EUDICOTS

in New Zealand. Griselinia littoralis is the species most frequently cultivated, and it makes evergreen salt-tolerant hedges, good for protecting other plants against salt spray in the UK, New Zealand and British Columbia. Etymology: Griselinia is named for Italian historian, naturalist and traveller Francesco Griselini (1717–1783).

448. PITTOSPORACEAE Cheesewood family

Genera and species: The only genus in this family, Griselinia, has six species, four in South America and two in New Zealand. Uses: Timber of tree-like Griselinia species has been used for railway sleepers

Billardiera cymosa, near Adelaide, South Australia [448]

Pittosporum tobira, Helsinki City Winter Garden, Finland [448]

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This is a family of trees, shrubs and vines with coloured or clear sap and often containing essential oils that make the plants fragrant when crushed. Plants are self-supporting or climbing with stems that vine in an anticlockwise direction. Stems are sometimes spiny. Leaves are simple, evergreen, alternate or in whorls and often aromatic, lacking stipules, but usually with petioles that clasp the stems somewhat. Leaf blades have entire, undulate or serrate margins and pinnate venation. Inf lorescences are terminal or axillary corymbs or thyrses, or the flowers are solitary. Flowers are bisexual (rarely unisexual) and actinomorphic (weakly zygomorphic in Cheiranthera). The five sepals are free or sometimes basally fused and deciduous, and the five petals are free or fused into a short tube, sometimes slightly clawed. The five stamens are free and alternate with the petals. Anthers are dorsifixed or nearly basifixed and open by pores or short slits. The superior ovary is composed of two (rarely

Bursaria spinosa, Australian Arid Lands Botanic Garden, Port Augusta, South Australia [448]

Pittosporum senacia in fruit, Réunion [448]

Hymenosporum flavum in fruit, Australian National Botanical Garden, Canberra [448]

Hymenosporum flavum, planted in Perth, Western Australia [448]

APIALES

EUDICOTS

three to five) fused carpels that usually form a single locule (rarely up to five). A single style with a wet stigma tops the ovary. Fruits are dry, loculicidal capsules or fleshy berries. Distribution: The family is distributed across the Old World tropics extending into the Pacific to Hawaii and French Polynesia and into temperate East Asia (China, Korea, southern Japan), Tasmania and New Zealand. There is an isolated population on the Canary Islands (Pittosporum coriaceum). Pittosporaceae have their greatest diversity in Australia. Phylogeny and evolution: The family was formerly placed in Rosales due to its misleading f loral characters. Their true affinities lie in Apiales, where they are sister to the Araliaceae-Apiaceae clade, from which they diverged c. 41–48 million years ago. Generic delimitation has been problematic. Genera and species: The seven genera of Pittosporaceae include c. 245 species: Auranticarpa (6), Bentleya (2), Billardiera (24), Bursaria (7), Cheiranthera (5), Hymenosporum (1) and Pittosporum (c. 200). Uses: Seeds of petroleum nut (Pittosporum resiniferum) are pressed for oil that burns brightly and is used for lighting locally in the Philippines and Malaysia. Wollum wollum wood (Hymenosporum flavum) is sometimes used for cabinetry. Cheesewood (Pittosporum undulatum) is commonly used to make golf clubs, but the species has become an aggressive invader in many parts of the world. A few species are used as garden ornamentals due to their scented flowers or attractive foliage. Most frequently cultivated are Australian bluebell or sollya (Billardiera heterophylla), karo (Pittosporum crassifolium), tarata (P. eugenioides), kohuhu (P. tenuifolium), Japan-ese cheesewood (P. tobira) and Australian cheesewood (P. undulatum). In Australia, purple appleberry (Billardiera longifolia), Christmas bush (Bursaria spinosa) and Australian frangipani (Hymenosporum flavum) also can be found in gardens and as street plantings. Etymology: Pittosporum is composed of the

Greek words πίσσα (pissa or pitta), pitch or tar, and σπορο (sporo), seed, referring to its sticky seeds.

449. ARALIACEAE Ivy family

These are shrubs, trees, vines (Hedera) and perennial herbs (some Aralia, Panax, Hydrocotyle, Neosciadium and Trachymene) with short stems that usually branch sparsely and bear prominent leaf scars. Leaves are usually alternate and spirally arranged, rarely opposite, and pinnately, bipinnately or palmately compound or simple and then often palmately lobed, sometimes peltate (Hydrocotyle), occasionally aromatic and gland-dotted. Blades have entire, lobed, toothed or serrate margins, and venation is pinnate or palmate. Petioles are usually long and well-developed, and stipules are absent or minute. Inflorescences are usually simple or compound umbels or heads, rarely spikes or racemes, but the umbels are often arranged in racemose or spicate inflorescences, rarely epiphyllous. The bisexual (occasionally unisexual) flowers are actinomorphic. The usually five (rarely three to 12, or absent) sepals are free or fused into a small untoothed rim. The five or ten (rarely three to 12) petals are free or partially fused. Stamens are as many as the petals and alternate with them or numerous (in Plerandra) and surrounding a nectar disk. Anthers are dorsifixed and open by lengthwise slits. The inferior, rarely half inferior or superior (Tetraplasandra, now within Polyscias) ovary is composed of two to ten (or up to 15 in Plerandra) carpels fused to each form a locule, topped with as many styles as carpels that are free or partially fused. Fruits are drupes or berries (Aralioideae) or schizocarps (Hydrocotyloideae).

Distribution: Araliaceae are widespread and most diverse in the tropics. They extend north into Arctic North America and Antarctic South America, but in Europe and Asia they do not extend as far north, remaining in the temperate zones, apart from the Russian Far East, where they can be found in the Amur region and Sakhalin. There are radiations in the mountains in tropical Africa, tropical America, New Zealand and the Pacific Islands. Phylogeny and evolution: The traditional distinction between woody Araliaceae and herbaceous Apiaceae no longer holds, and this character was always strange considering that there are several herbaceous Aralia and Panax species that were always placed in Araliaceae despite their herbaceous habit. Hydrocotyle and Trachymene (including Uldinia), previously in Apiaceae but now in Araliaceae, were found to be most closely related to Neosciadium, a genus unusual in Araliaceae for having dry fruits rather than berries and thus a character which matches Hydrocotyloideae. Other genera formerly associated with Hydrocotyloideae (Azorella, Centella and Xanthosia) remain in Apiaceae. Formerly placed in this family were Diplopanax (now in Nyssaceae, Cornales), Homalosciadium (part of Platysace in Apiaceae) and Delarbrea and Myodocarpus (now Myodocarpaceae). Genera of Araliaceae are often poorly circumscribed and need much taxonomic and nomenclatural attention (the list below is tentative). At least it is known that Gastonia is embedded in Polyscias, and the species names have thus been transferred. Also Schefflera is widely polyphyletic, with five clades found throughout the Araliaceae tree corresponding to geographical regions, and these need to be recognised at generic levels. Only the Malesian clade has been revised so far and is now called Plerandra, which includes the common houseplant false aralia (P. elegantissima). This will change the circumscription of Schefflera, but this is yet to be undertaken. It has been estimated that Araliaceae diversified c. 80 million years ago, although ages half this old have also been suggested. Herbaceous and bicarpellate Hydrocotyloideae

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APIALES

Aralia continentalis, Helsinki Botanical Garden, Finland [449]

EUDICOTS

Hedera helix, Twickel Estate, Delden, the Netherlands [449]

Schefflera actinophylla, planted in Guayaquil, Ecuador [449] Trachymene glaucifolia, William’s Creek, South Australia [449]

are sister to the rest of the family, and these may well have been the plesiomorphic states of the family, woodiness having probably evolved secondarily. Genera and species: Araliaceae include 40 (+3) genera and c. 1,650 species in two subfamilies: Hydrocotyloideae – Hydrocotyle (c. 130), Neosciadium (1) and Trachymene (c. 50); Aralioideae – Anakasia (1), Apiopetalum (2), Aralia (74), Astrotricha (20), Brassaiopsis (47), Cephalaralia (1), Cheirodendron (6), Cheniopanax (2), Cussonia (20), Dendropanax (98), Eleutherococcus (37), Fatsia (3), Gamblea (4), Harmsiopanax (3), Hedera (17), Heteropanax (10), Kalopanax (1), Macropanax (17), Merrilliopanax (4), 630

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Meryta (29), Metapanax (2), Motherwellia (1), Neopanax (5), Oplopanax (3), Oreopanax (146), Osmoxylon (60), Panax (13), Plerandra (>37), Polyscias (175), Pseudopanax (7), Raukaua (6), Schefflera (c. 600, polyphyletic), Seemannaralia (1), Sinopanax (1), Tetrapanax (1), Trevesia (11) and Woodburnia (1). Uses: The most famous plant of this family is ginseng, species of the the genus Panax, mainly Asian ginseng (P. ginseng), Japanese ginseng (P. japonica), North American ginseng (P. quinquefolius) and Siberian ginseng (Eleutherococcus senticosus). Folk medicine attributes ginseng as a cure for all sorts of ailments (Panax means ‘all-heal’ in Greek), but little scientific proof for its benefit

Hydrocotyle bonariensis naturalised on a beach in New South Wales, Australia [449]

Pseudopanax laetus, Trebah Garden, Cornwall [449]

exists. The plants contain ginsenosides, unique to the genus Panax, which have a complex pathway in the human body, but are believed to protect cells against stress. There is some proof that they work in viral outbreaks and in stimulating white blood cell production, but insomnia can be a common side-effect of ginseng use. The name is derived from Chinese renshen, meaning person-root, referring to the forked shape of ginseng root. The pith of the rice paper plant, Tetrapanax papyrifer, is used to make a fine-quality paper in Asia, especially in 19th century Guangdong, China. The paper was commonly used for gouache paintings traded with westerners visiting at the time, and it was mistakenly believed that the paper was

APIALES

EUDICOTS

made from rice, hence the common name ‘rice paper’. The texture of the paper makes it unsuitable for writing, but it is excellent for water colours or crafting paper flowers, for which it is still often used. The wood of Araliaceae is often weak, but timber and firewood can be harvested from some species of Eleutherococcus, Meryta and Schefflera. Many species are grown as ornamental plants, especially Plerandra, Polyscias, Schefflera, Hedera and Fatsia, which are often found grown as potplants or as garden plants in suitable climates. A hybrid between the last two, ×Fatshedera lizei, has been developed and is also sometimes grown as a house or garden plant. Hedera helix (ivy) can be an aggressive invasive outside (and within) its native range. Other popular garden plants belong to the genera Aralia, Dendropanax, Eleutherococcus, Kalopanax, Meryta, Oplopanax, Oreopanax, Osmoxylon, Polyscias, Scheff lera, Trachymene and Tetrapanax. Hydrocotyle (pennywort) is often grown as an aquarium or pond plant and escapes easily, some species clogging waterways. Etymology: Aralia is the Latinised form of aralie, an old French Canadian name for the plant, probably derived from a native American language.

Myodocarpus fraxinifolius, New Caledonia [450]

450. MYODOCARPACEAE Mousefruit family

Myodocarpaceae are a family of trees and shrubs with alternate, spirally arranged, pinnately compound (rarely simple) leaves without stipules. Blades are entire, rarely toothed with pinnate venation. Inflorescences are compound umbels arranged in panicles or racemes. Flowers are unisexual or bisexual and actinomorphic. The five sepals and petals are each free. The five stamens are free and alternate with the petals, the dorsifixed anthers opening by lengthwise slits. The inferior or partially inferior ovary is composed of two fused carpels that each form a locule topped with a free style. Fruits are drupes (Delarbrea) or schizocarps that fall apart to form laterally flattened samaras (Myodocarpus), resembling mouse-ears. Distribution: This family is nearly restricted

to New Caledonia, with a few Delarbrea species found on other Melanesian islands and in eastern Indonesia (Timor, Moluku), New Guinea and extending to northeastern Australia. Phylogeny and evolution: Myodocarpus is similar in wood anatomy to Cornaceae but in other aspects resembles Apiaceae more than Araliaceae, where it was previously placed. It has always been a morphologically intermediate taxon and is now found to be sister to Apiaceae, with which it shares chemistry and anthers that are inflexed in bud. It is estimated that Myodocarpaceae diverged from Apiaceae c. 29–32 million years ago and that the crown group is c. 25.4 million years old. Their current distribution is undoubtedly the result of island hopping. Genera and species: This family includes only two genera with 15 species in total: Delarbrea (7) and Myodocarpus (8). Uses: Pachycaul species of Delarbrea are sometimes cultivated as curiosities. Etymology: Myodocarpus is composed of the Greek words μυώδες (myodes), mouselike, and καρπός (karpos), fruit, referring to the ‘eared’ samarras.

Myodocarpus fraxinifolius, New Caledonia [450]

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APIALES

451. APIACEAE Carrot family

Apiaceae are a family of bisexual and (occasionally) unisexual, annual, biennial and perennial herbs and shrubs, rarely trees (Heteromorpha, Steganotaenia), with pithfilled furrowed stems and often with oil canals. Alternate leaves are usually pinnately or palmately compound, sometimes simple. Petioles are usually sheathing at the base, clasping the stem. Leaf blades have pinnate, parallel or palmate venation and entire, toothed, serrate, crenate or prickly margins; they are often fragrant due to the presence of ethereal oils. Inflorescences are bracteate simple or compound umbels, sometimes contracted into heads with whorls of bracts that are green or sometimes colourful (an involucre). Flowers are actinomorphic or Anthriscus sylvestris, sheathing leaf bases, Richmond, Surrey, UK [451]

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weakly zygomorphic (sometimes one petal enlarged on the outside of the umbel, creating a ‘false flower’ (pseudanthium). The five sepals are fused to the ovary and free or fused to each other, often toothed or weakly lobed, sometimes entire. The five petals are free, often clawed, sometimes absent, sometimes unequal in shape and size. Stamens alternate with the petals, placed on a conspicuous epigynous nectar disk. Anthers are dorsifixed and open by lengthwise slits. An inferior ovary is composed of two fused carpels, each forming a single locule and topped with a persistent style often swollen at the base (stylopodium). Fruits are flat to round, distinctly ribbed, often stalked schizocarps, parting into two nutlets. Oil-tubes (vittae) are often present between or below the ribs of the fruits, occasionally hooked or spined, rarely fleshy and drupe-like.

Apiaceae have been variously circumscribed in the past and often included Araliaceae and Myodocarpaceae. Especially now that several genera of Apiaceae have been found to belong to Araliaceae and vice versa, the distinction between the families is blurred and poorly ref lected in morphological characters. The families are here maintained because of convention. Four subfamilies (Apioideae, Azorelloideae, Mackinlayoideae, Saniculoideae) are currently accepted, but these are not supported in all current phylogenetic studies, with Platysace (including Homalosciadium) and Hermas forming separate clades apart from the established subfamilies. In addition, some genera are in need of recircumscription.

Phylogeny and evolution: Apiaceae diversified c. 54–87 million years ago. Australian, South American and African origins of the family have all been suggested.

Genera and species: Apiaceae include c. 443 genera and c. 3,575 species: Aciphylla (39), Acronema (25), Actinolema (2), Actinotus (c. 20), Adenosciadium (1), Aegokeras (1), Aegopodium (7), Aethusa (1), Afrocarum (1), Afroligusticum (1), Afrosciadium (18), Afrosison (3), Agasyllis (1), Ainsworthia (2), Alepidea (40), Aletes (c. 15), Alococarpum (1), Ammi (4), Ammodaucus (1), Ammoides (2), Ammoselinum (3), Anethum (1), Angelica (c. 110), Anginon (12),

Angelica archangelica subsp. archangelica, Helsinki Botanical Garden, Finland [451]

Azorella trifurcata, Royal Botanic Gardens, Kew, UK [451]

Distribution: This family has a nearly worldwide distribution but is most diverse in the Mediterranean, Central Asia, the Himalayas, southern Africa and the Andes.

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Coriandrum sativum, private allotment, Kingston upon Thames, Surrey, UK [451]

Cryptotaenia canadensis, Bergius Botanical Garden, Stockholm, Sweden [451]

Angoseseli (1), Anisotome (15), Annesorhiza (12), Anthriscus (9), Aphanopleura (6), Apiastrum (2), Apiopetalum (2), Apium (25), Apodicarpum (1), Arafoe (1), Arctopus (3), Arcuatopterus (8), Arpitium (2), Arracacia (55), Artedia (1), Asciadium (1), Astomaea (2), Astrantia (8), Astrodaucus (2), Astydamia (1), Athamantha (6), Aulacospermum (14), Austropeucedanum (1), Autumnalia (2), Azilia (1), Azorella (26), Berula (1), Bifora (3), Bilacunaria (4), Billburttia (2), Bolax (5), Bonannia (1), Bowlesia (15), Brachyscias (1), Bunium (48), Bupleurum (c. 185), Cachrys (4), Calyptrosciadium (1), Canaria (1), Capnophyllum (5), Carlesia (1), Caropsis (1), Carum (c. 30), Caucalis (1), Cenolophium (1), Centella (c. 50), Cephalopodum (3), Cervaria (4), Chabrea (1), Chaerophyllopsis (1), Chaerophyllum (33), Chamaele (1), Chamaesciadium (1), Chamaesium (3), Chamarea (5), Changium (2), Chesneya (30), Chlaenosciadium (1), Choritaenia (1), Chuanminshen (1), Chymsydia (2), Cicuta (8), Cnidiocarpa (2), Coaxana (2), Conioselinum (18), Conium (6), Conopodium (6), Coriandrum (3), Coristospermum (3), Cortia (8), Cortiella (3), Cotopaxia (2), Coulterophytum (5), Crithmum (1), Cryptotaenia (5), Cuminum (4), Cyathoselinum (1), Cyclorhiza (2), Cyclospermum (3), Cymopterus (32), Cynapium (1), Cynorhiza (4), Cynosciadium (2), Dahliaphyllum (1), Dasispermum (1), Daucus (35), Demavendia (1), Dethawia (1), Deverra (10),

Dichosciadium (1), Dickinsia (1), Dicyclophora (1), Didiscus (1), Dimorphosciadium (1), Diplaspis (3), Diplolophium (c. 6), Diplotaenia (2), Diposis (3), Distichoselinum (1), Dolpojestella (2), Domeykoa (4), Donnellsmithia (c. 17), Dorema (12), Dracosciadium (2), Dregea (c. 12), Drusa (1), Ducrosia (5), Dystaenia (2), Echinophora (9), Ekima (1), Elaeoselinum (4), Elaeosticta (24), Eleutherospermum (1), Enantiophylla (1), Eremocharis (9), Eremodaucus (1), Ergocarpon (1), Erigenia (1), Eriosynaphe (1), Eryngium (c. 250), Erythroselinum (1), Eurytaenia (2), Exoacantha (1), Ezosciadium (1), Falcaria (5), Fergania (1), Ferula (130), Ferulago (43), Ferulopsis (2), Foeniculum (5), Frommia (1), Froriepia (2), Fuernrohria (1), Geocaryum (c. 13+2 extinct), Gingidia (13), Glaucosciadium (1), Glehnia (2), Glia (1), Glochidotheca (1), Gongylosciadium (1), Gongylotaxis (1), Grammosciadium (7), Guillonea (1), Gymnophyton (6), Hacquetia (1), Halosciastrum (1), Haloselinum (1), Hansenia (3), Haplosciadium (1), Haplosphaera (2), Harbouria (1), Harrysmithia (2), Haussknechtia (1, possibly extinct), Hellenocarum (3), Heptaptera (6), Heracleum (65), Hermas (8), Heteromorpha (7), Hladnikia (1), Hohenackeria (2), Holandrea (4), Homalocarpus (6), Homalosciadium (1), Horstrissea (1), Huanaca (4), Hyalolaena (10), Hymenidium (35), Hymenolaena (3), Imperatoria (3), Indoschultzia (2), Itasina (1), Johrenia (15), Johreniopsis (4), Kadenia (1),

Daucus carota subsp. carota, Sicily, Italy [451]

Kafirnigania (1), Kailashia (2), Kalakia (1), Kamelinia (2), Kandaharia (1), Karatavia (1), Karnataka (1), Kedarnatha (5), Kelussia (1), Keraymonia (4), Kitagawia (5), Klotzschia (3), Komarovia (1), Korshinskia (5), Kosopoljanskia (3), Kozlovia (4), Krubera (1), Kundmannia (2), Kuramosciadium (1), Ladyginia (3), Lagoecia (1), Lalldhwojia (4), Laretia (2), Laser (8), Laserpitium (6), Lecokia (1), Ledebouriella (2), Lefebvrea (6), Leiotulus (9), Lereschia (1), Leutea (6), Levisticum (1), Lichtensteinia (7), Ligusticopsis (14), Ligusticum (c. 50), Lilaeopsis (14), Limnosciadium (2), Lipskya (1), Lisaea (3), Lomatium (74), Lomatocarpa (4), Mackinlaya (5), Macroselinum (1), Magadania (2), Magydaris (2), Mandenovia (1), Margotia (1), Marlothiella (1), Mastigosciadium (1), Mathiasella (1), Mediasia (1), Meeboldia (5), Melanosciadium (1), Meum (3), Micropleura (2), Microsciadium (1), Modesciadium (1), Mogoltavia (2), Molopospermum (1), Mozaffariania (1), Mulinum (20), Musineon (3), Mutellina (3), Myrrhidendron (5), Myrrhis (1), Nanobubon (2), Naufraga (1), Neogoezia (5), Neomuretia (2), Neonelsonia (2), Neoparrya (2), Neosciadium (1), Niphogeton (18), Nirarathamnos (1), Normantha (1), Nothosmyrnium (2), Notiosciadium (1), Notobubon (12), Notopterygium (6), Oedibasis (4), Oenanthe (40), Oligocladus (2), Oliveria (1), Opoideia (1), Opopanax (3), Opsicarpium (1), Oreocome (6), Oreocomopsis (3), Oreomyrrhis (23), Oreonana (3), Plants of the World

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Orlaya grandiflora, Bergius Botanical Garden, Stockholm, Sweden [451]

Oreoschimperella (3), Oreoxis (4), Orlaya (3), Ormopterum (2), Ormosciadium (1), Ormosolenia (1), Orogenia (2), Oschatzia (2), Osmorhiza (10), Ottoa (1), Oxypolis (7), Pachypleurum (1), Palimbia (3), Pancicia (1), Paraselinum (1), Parasilaus (2), Pastinaca (14), Pastinacopsis (1), Paulita (3), Pedinopetalum (1), Perideridia (13), Perissocoeleum (4), Petagnaea (1), Petroedmondia (1), Petroselinum (2), Peucedanum (c. 110), Phellolophium (1), Phlojodicarpus (4), Phlyctidocarpa (1), Physospermopsis (15), Physospermum (2), Physotrichia (11), Pilopleura (2), Pimpinella (c. 200), Pinacantha (1), Pinda (1), Platysace (26), Pleurospermum (2), Podistera (4), Polemannia (3), Polemann-iopsis (1), Polytaenia (2), Polyzygus (1), Portenschlagiella (1), Postiella (1), Pozoa (2), Prangos (38), Prionosciadium (8), Psammogeton (7), Pseudocarum (2), Pseudocymopterus (7), Pseudopimpinella (1), Pseudoridolfia (1), Pseudoselinum (1), Pseudotrachydium (5), Pternopetalum (32), Pterygopleurum (1), Pteryxia (5), Ptilimnium (5), Ptychotis (1), Pycnocycla (12), Pyramidoptera (1), Registaniella (1), Rhabdosciadium (5), Rhizomatophora (1), Rhysopterus (3), Ridolfia (1), Rivasmartinezia (1), Rumia (1), Rutheopsis (1), Sajanella (1), Sanicula (39), Saposhnikovia (1), Scaligeria (3), Scandix 634

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Eryngium giganteum, Ruissalo Botanical Garden, Turku, Finland [451]

(7), Scaraboides (1), Schizeilema (1), Schoenolaena (2), Schoenoselinum (1), Schrenkia (12), Schtschurowskia (2), Schulzia (4), Sciothamnus (4), Sclerochorton (1), Sclerotiaria (1), Scrithacola (1), Selinopsis (2), Selinum (2), Semenovia (18), Seseli (c. 110), Seselopsis (2), Shoshonea (1), Siculosciadium (1), Siebera (2), Silaum (1), Siler (1), Silphiodaucus (1), Sinocarum (c. 24), Sinolimprichtia (1), Sison (2), Sium (14), Smyrniopsis (1), Smyrnium (7), Sonderina (4), Spananthe (1), Spermolepis (5), Sphaenolobium (3), Sphallerocarpus (1), Spuriodaucus (3), Stefanoffia (2), Steganotaenia (3), Stenocoelium (3), Stenosemis (2), Stenotaenia (6), Stewartiella (2), Stilbocarpa (3), Stoibrax (3), Symphyoloma (1), Synclinostyles (2), Szovitsia (1), Taenidia (2), Tamamschjanella (2), Tamamschjania (1), Tauschia (31), Tetrataenium (8), Thamnosciadium (1), Thapsia (8), Thaspium (3), Thecocarpus (2), Thyselinum (1), Todaroa (1), Tommasinia (1), Tongoloa (15), Tordyliopsis (1), Tordylium (18), Torilis (15), Trachydium (1), Trachyspermum (15), Trepocarpus (1), Tricholaser (2), Trigonosciadium (4), Trinia (c. 10), Trochiscanthes (1), Turgenia (1), Vanasushava (1), Vesper (6), Vicatia (3), Villarrealia (1), Xanthosia (25), Xatardia (1), Xyloselinum (2), Yabea (1), Zeravschania (6), Zizia (4) and Zosima (4).

Uses: Apiaceae include well-known root and leaf vegetables, herbs, spices, medicines and ornamental plants. One of the best known root crops is carrot (Daucus carota), which is not palatable in its wild state, but cultivated forms (subsp. sativus) are commercially grown, sweeter and less fibrous. Carrots can be white, yellow or purple, but are most commonly orange, derived from 16th century Dutch stock. Purple ones were first domesticated in Afghanistan, from where they spread across Asia, whereas orange carrots have been more popular in cultivation in the west. They have been crossed with D. capillifolius to breed resistance against carrot fly. Similar to carrots, parsnip (Pastinaca sativa) is a common root vegetable, especially in the USA and UK, although it is slowly gaining popularity again in other countries as a ’forgotten vegetable’. Crummock or skirret (Sium sisarum) has a parsnip-like root that tastes like sweet potato. It has been cultivated in the eastern Mediterranean since ancient times, where it was probably introduced from China. It has lost popularity but is promising as a new commercial vegetable, although the core has to be removed before cooking, which some people may find cumbersome. Turnip-rooted chervil (Chaerophyllum bulbosum) has an edible, carrot-like tuber

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and is in Europe locally cultivated for this on a small scale. The roots increase in sweetness after storage for a few months. Great pignut (Bunium bulbocastanum) has edible tubers, leaves and seeds and was formerly much cultivated, but it too has lost popularity. The tubers taste of sweet chestnut, the leaves are like parsley and the seeds, which are called ”black cumin”, have a smoky, earthy taste and are used as a culinary spice in the Indian subcontinent. Similarly small pignut or earthnut (Conopodium majus) has tubers that taste like chestnuts after roasting. Biscuitroots or cousroots (Lomatium ambiguum, L. cous, L. geyeri, L. macrocarpum) have roots that can be eaten raw or pounded into cakes or biscuits, a traditional native American food. Flour from the tubers of yampah (Perideridia gairdneri) was formerly also much eaten by native Americans and early settlers. Conioselinum pacificum (wild carrot or hemlock-parsley) was eaten in western North America. An important commercial root crop in the Andes, Peruvian parsnip or arracacha (Arracacia xanthorrhiza) is eaten like parsnip. It locally replaces potatoes in importance and is used in much the same way, boiled, as dumplings, in flour, purées, soups etc. It has a small starch grain that is easily digestible, and this crop deserves more attention as an alternative food source.

The swollen fleshy stems of several species are also popular foods, especially the petioles of celery (Apium graveolens var. dulce) that are commercially grown and commonly available in supermarkets. They are eaten raw or cooked and are important in French, Creole and Cajun cuisine. Celeriac (Apium graveolens var. rapaceum) is a variety of the same species that has a swollen root, which can be eaten like a parsnip, cooked or baked, or sliced and mixed with mayonnaise to make celeriac salad. It also is a main ingredient of traditional splitpea soup. Leaves of celery and celeriac can be used as a herb; the seeds also yield a volatile oil that is used in perfumery and industry. Seeds of Apium graveolens (celery) are used as a spice, usually to produce celery salt. It is also used to relieve arthritis and rheumatism. Jellico (Sium bracteatum) is an endemic of St Helena that has hollow stems similar to celery and was harvested from the wild and eaten locally. It has become increasingly rare there due to habitat fragmentation and invasive mammals, but it has potential as a crop. Another fragrant herb of which the leaves can be blanched like celery and eaten in a similar way is lovage (Levisticum officinale), which has been cultivated in the Eastern Mediterranean since ancient times. Its leaves are also used as a flavouring for soups, and its fruits and roots are used to flavour liqueurs

Astrantia major, private garden, Kingston upon Thames, Surrey, UK [451]

etc. Scots lovage (Ligusticum scoticum) is a potherb similar in use to celery and lovage, and alexanders (Smyrnium olusatrum) also have edible roots and stems that were consumed like celery. Alexanders were a common vegetable during Roman times in the UK and elsewhere in Europe, but it is no longer consumed and now naturalised in several areas, especially southern Britain, Ireland, the Netherlands and Bermuda. Another well-known stemvegetable is fennel (Foeniculum vulgare) that has varieties that make swollen stems and leaf bases. Other varieties are selected for the seeds and yield an anise-like essential oil commonly used in alcoholic drinks such as anisette, ouzo, pastis, rakı etc. Angelica (Angelica archangelica and to a lesser extent A. atropurpurea) has leaves that are eaten as a vegetable, especially in Siberia and Lapland. The hollow leaf and inflorescence stalks can be candied, especially in France, which is why it is called French rhubarb. The candied stems are used in cake decoration, often together with candied cherries and citrus rind. Angelica is also used to flavour alcoholic drinks such as Benedictine, Chartreuse and other liqueurs, gins, wines and tonics. The hollow stems of angelica are said to have been the inspiration for the grooved Doric columns in Greek classical architecture. Eryngo root, harvested from field eryngo

Sanicula arctopoides, Sea Ranch, California, USA [451]

Xanthosia rotundifolia, Albany, Western Australia [451]

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APIALES (Eryngium campestre) and sea-holly (E. maritimum), was formerly candied in the UK and marketed as ’kissing comfits’. It was said to have aphrodisiac properties. Sea samphire (Crithmum maritimum) leaves were formerly pickled, having a particular spicy and salty taste, and it is a shame it is no longer popular. Many species of Apiaceae have leaves that can be cooked like spinach and are sometimes important local vegetables. Best known of these is perhaps mitsuba or Japanese parsley (Cryptotaenia canadensis), which is cultivated on a large scale in China and Japan as a salad green. Its roots are sometimes used in stirfries. Minari or Chinese celery (Oenanthe javanica) is an important vegetable in East Asia, especially Taiwan. Asian pennywort or gotu kola (Centella asiatica) is a pantropical weed with edible leaves, especially used in Sinhalese and other Asian cuisines, often in salads or steamed dishes. Once planted it can be a troublesome weed. Recently, Centella has been applied as a hair lotion, together with green coffee bean extract and an antioxidant (often rosemary, Rosmarinus officinalis), to reverse hair loss. Apart from celery, fennel and lovage, there are many other species with fragrant leaves that make excellent herbs. Parsley (Petroselinum crispum), often presented as a food garnish, is one of the richest sources of vitamin C. It is also popular in salads and meat dishes and sausages. A cultivar P. crispum ’Tuberosum’ (Hamburg parsley) has an edible tuber. Dill (Anethum graveolens) originated in India but has been cultivated in Europe since 400 BC. It is a popular herb accompanying fish, especially smoked salmon. Chervil (Anthriscus cerefolium) is a delicate herb with a slight anise flavour used in salads, soups, omelettes and other dishes, particularly in Belgium. Like many other potherbs from Europe, it has been cultivated since ancient times. Similar to chervil is sweet cicely (Myrrhis odorata), a potherb with anise-flavoured leaves that is delicious in salads. It was formerly used to polish oak panelling and to deter insects from closets. An unlikely herb is goutweed or ground elder (Aegopodium podagraria). Best known as a troublesome garden pest, it was originally

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introduced to many parts of Europe by the Romans as a potherb. Young leaves have a good flavour and can be boiled like spinach or made into an excellent variant on pesto. It was originally used medicinally to treat gout, hence the colloquial name bishop weed, in an allusion to the opulent lifestyle that bishops were believed to have. Mediterranean hartwort (Tordylium apulum) has edible leaves that are used as a condiment or salad green in Italy and Greece. Probably one of the best-known herbs in this family is coriander (Coriandrum sativum), which has been cultivated since ancient times for its seeds. Together with celery seeds, they were preserved in Tutankhamun’s tomb (1325 BC), and coriander is one of the bitter herbs of the Jewish Passover feast. Ground seeds of coriander, known as dhani, are a main ingredient of Asian curries, and fresh leaves, often called cilantro, are popular as a garnish or flavouring in many dishes in Latin American, Mediterranean and several Asian cuisines. Not everyone appreciates this herb, including the Ancient Greeks, who named it in reference to a stink bug (‘korís’) because of the foetid smell of the leaves and unripe fruits. There are reports that some people are genetically predisposed to dislike cilantro, but little is still known about cilantrophobe genetics. In fact, many of the aldehydes that produce the scent and taste of cilantro are also found in some types of soap, and this may be why some people associate cilantro with the taste of soap. However, the seeds are a popular spice, and it is grown commercially on a large scale for use in gin, candies, breads, perfumes and indeed soap. In some tropical countries, cilantro is often replaced by false cilantro (Eryngium foetidum), which has a similar, somewhat stronger scent, but the flavour remains when the leaves are dried. A spice that has also been used for 4,000 years to flavour our foods and drinks is anise (Pimpinella anisum), the seeds of which are used to flavour milk or alcoholic drinks (anisette, absinthe, anis, aquavit, ouzo, pastis, rakı etc.). It is also used in confectionery, in which seeds are coated in a sugary substance, a popular treat in the Netherlands (called little mice “muisjes”) when a child is born. Aniseed oil is popular in aromatherapy and

also used medicinally. Caraway seed (Carum carvi) is another common spice used to flavour bread, cheese, sauerkraut and Kümmel liqueur. Cumin (Cuminum cyminum) has been cultivated for its fruits since the 13th century BC in Crete. It is still used occasionally to flavour some Dutch cheeses, cakes and liqueurs and in curry powder but has largely been replaced by caraway. An important spice in India is asafoetida (Ferula assa-foetida, F. foetida and F. narthex) of which the seeds and gum resin are used as a spice in Hare Krishna cuisine to replace onions. It is also an important ingredient of Worcestershire Sauce. Another common spice in India is ajwain (Trachyspermum ammi) of which the fruits, called carom seeds, are roasted and commonly eaten with beans and other legumes, reducing their gas-producing effect on the body. It is also the major spice of Bombay mix flavoured nuts. Ajmud or radhuni (Trachyspermum roxburghianum) fruits have a smell similar to parsley or celery and can be used in pickles or spice mixtures and curries, but it is strong and easily overpowers other spices. In North Africa Ammodaucus leucotrichus is sometimes used as a condiment. Candy carrot (Athamanta cretensis) is a condiment and used to flavour a liqueur. Because of the numerous ethereal oils found in this family, many gum exudates are used in the perfume industry. Gum ammoniacum is mainly harvested from Dorema ammoniacum and used as a scent for incense and in medicine. It is collected in Iran and India from stems that have been damaged by insects. Gum opopanax is harvested from Opopanax chironium and also used in scent making, just like gum galbanum, a biblical resin harvested from Ferula gummosa and F. rubricaulis. There are many other medicinal properties that cannot be enumerated here. As often with plants used in medicine, poisons are common too. Cowbane (Cicuta virosa) is probably being the most violently poisonous plant in the North Temperate zone, but hemlock water-dropwort (Oenanthe crocata) is almost as deadly. Both have been used in the past, either on purpose or accidentally, with unfortunate results. Hemlock (Conium maculatum) is also a poisonous plant and is

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said to have poisoned Socrates. Hemlock has spread dramatically along motorway verges in the southern UK in recent decades. All three have been used as rodent poison in the past. Also poisonous, Aethusa cynapium is called fool’s parsley as it has been mistaken for true parsley or celery with disastrous results. Llareta (Azorella compacta) was an important fuel source in the Andes because it burns without much smoke, but the species is now endangered due to overharvesting. Many plants are used as garden ornamentals for annual or perennial bedding or as speciality plants in alpine collections. The best known are Aciphylla, Actinotus helianthi (flannel flower), Ammi majus (bullwort), Angelica gigas, Astrantia (masterwort),

Scandix pecten-veneris, Ronda, Spain [451]

Azorella trifurcata, Bupleurum (hare’s ears, thorow-wax), Chaerophyllum hirsutum (hairy chervil), Eryngium (eryngo, rattlesnake master), Ferula communis (giant fennel), Foeniculum (ornamental fennel), Hacquetia epipactis, Heracleum (hogweed), Meum athamanthicum (baldmoney), Mathiasella bupleuroides, Molopospermum peloponnesiacum (striped hemlock), Orlaya, Osmorhiza (sweetroot), Sanicula (sanicle) and Zizia aurea (golden alexanders). Giant hogweeds (Heracleum mantegazzianum, H. speciosum, H. persicum, H. pubescens) were introduced to western Europe as garden ornamentals in the 19th century and soon naturalised. They are large plants, crowding and shading out native vegetation, and they cause severe skin

Myrrhis odorata, private garden, Kingston upon Thames, Surrey, UK [451]

inflammations when touched and the skin then exposed to sunlight. Eradication of these exotic species has been costly and to date has not been successful. In Norway, giant hogweed has been referred to as Tromsø palm, due to it having first naturalised around that city. Etymology: Apium is the classical Latin name for celery and parsnip, derived from apis, a bee. Umbelliferae (which means ‘umbel carriers’) is an alternative name for this family, but it is not based on the genus Umbellifera, but rather on Apium as well. In our opinion, it is better to have a family name that reflects the genus on which it is based, especially since this is not the only family with umbellate inflorescences.

Heracleum lanatum, Sea Ranch, California, USA [451]

Hacquetia epipactis, Royal Botanic Gardens, Kew, UK [451]

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GLOSSARY The following glossary includes many common terms used in botany and botanical descriptions. These terms are not all used in this book because we have attempted to use as few terms as possible to make this work widely accessible. However the glossary below provides a good start for getting access to botanical terminology and understanding some of the references cited in this book. Additional botanical terms are explained and illustrated in the excellent work by Beentje H. 2010. The Kew plant glossary, an illustrated dictionary of plant terms. Kew Publishing, Richmond. From this work we extracted many of the definitions given below. Please consult this work for more in-depth definitions of botanical terminology or when a term is absent from our list below. A acaulescent = without a stem acerose = needle-shaped (Fig. 21a) achene = a small nut, a nutlet (Fig. 35i) acroscopic = facing towards the tip of the leaf actinomorphic = radially symmetrical (Fig. 31a) acuminate = tapering to a long tip (Fig. 25k) acute = sharply pointed or tapering to a short tip (Figs. 23c, 25h) adnate = fused with a surface of another type (as opposed to connate) adventitious = roots or buds growing from places other than normal aerial roots = roots emerging from the plant above ground aerophore = a projection or swelling along the petiole or rachis usually for gas exchange (especially in Thelypteris) aestivation = the way sepals and petals are folded in bud agglomerated = densely crowded but not fused alate = winged alkaloid = organic basic nitrogenous compound alpine = growing in high mountainous areas above the tree line alternate = inserted at different levels along the axis (Fig. 20a) amber = fossil plant resin amorphous = without regular form amphibious = adapted to live on land and in water amplexicaul = embracing the stem (Fig. 20f) anastomosing = with veins merging where they come into contact forming a network (Fig. 27e) anatomy = internal structure androgynophore = a stalk carrying both stamens and carpels above the petal insertion, as in Passiflora angiosperm = flowering plant, the ovules being enclosed in an ovary or carpel 638

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anisophyllous = having two opposite leaves that are of unequal size or shape annual = completing the life cycle within a single growing season annulus = a ring of thickened cells that aids in opening of a fern sporangium (Fig. 13) Antarctic = growing in cold climates at higher latitudes of the Southern Hemisphere anther = the part of the stamen containing the pollen (Figs. 29, 32) antheridium = male sexual organ in a gametophyte anthesis = the time when a flower opens or is available for pollination (more correctly: time when flower is fertilised) apetalous = without petals apex = tip APG = Angiosperm Phylogeny Group aphlebia = a leaflet different in form from normal leaflets, formed at the base of adult leaves aphyllous = without leaves apiculate = ending abruptly in a short point (Fig. 25i) apocarpous = with a simple fruit consisting of a single carpel or a multiple fruit with free carpels apomict = a taxon reproducing asexually either vegetatively or by producing seeds without fertilisation apomorphic = derived apopetalous = with free petals aposepalous = with free sepals appendage = an attached secondary part appressed = placed close and flat to another structure aquatic = living in water arachnoid = cobweb-like arborescent = tree-like archegonium = female sexual organs in a gametophyte Arctic = growing in cold climates, at higher latitudes of the Northern Hemisphere arcuate = curved like a bow

areole = a more or less circular area that is divided from similar areas by a line such as a vein, often used in description of venation types. The term is also used to describe the spine-bearing areas in cacti. aril = appendage partially or completely enclosing the seed article = an individual segment (of a fruit) articulate = jointed by nodes ascending = curving upwards asexual = without gender or sexless asymmetrical = with the two sides of an organ unequal in size attenuate = gradually narrowed (Fig. 23a) auriculate = with ear-like structures, usually at the base of a leaf or petal (Fig. 23g) autapomorphy = a character or trait unique to a group autotrophic = obtaining food by photosynthesis from carbon dioxide, water and inorganic matter awn = a fine bristle terminating an organ (usually used in Poaceae) axil = the area on the stem immediately above the point of attachment of a leaf axile placentation = a form of placentation (seed positions) for an ovary in which the ovules are attached to the central axis of the ovary (Fig. 31b) axillary = arising in an axil B baccate = berry-like balsamiferous = balsam producing banded = marked with differently coloured stripes bark = outermost layer of stems and roots in woody plants basal = at or near the base. In phylogenetics often used erroneously for a species-poor taxon or clade that is sister to a speciesrich one, but not meaning primitive. base = the point of attachment of an organ basifixed = used for anthers that are fixed to the filaments by the base (Fig. 32)

GLOSSARY

sorus

blade

leaf/frond

rachis pinna

petiole/stipe

sporangium annulus indusium

Figure 13. General fern terminology

sorus scale rhizome/stem roots

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GLOSSARY basiscopic = pointing towards the base beaded = with regular narrowing and widening, making it look like a string of beads berry = an indehiscent soft fruit with many seeds (rarely one) immersed in a fleshy pulp (Fig. 35b) bicrenate = doubly crenate, with scalloped edges of which the lobes are again scalloped biennial = taking two years or two growing seasons to reach maturity bifid = divided at the tip in two parts bifurcate = forked or divided into two sharp branches bilabiate = two-lipped, used when the calyx or corolla forms two separate projections as in many Lamiaceae bilateral = arranged on opposite sides bilateral symmetry = symmetrical on a central line or axis, dividing the organ in two mirrored halves bilobed = with two lobes bilocular = with two compartments, usually of the ovary bipartite = divided into two parts at the apex bipinnate = doubly pinnate with the pinnae divided into pinnules (Fig. 23j) bipinnate-pinnatifid = doubly pinnate with the pinnae partially divided into pinnules bisexual = having both sexes in the same flower or on the same plant biternate = compound ternate, with the blade divided into threes and the divisions themselves also divided into threes (Fig. 22f) bladder = a hollow, swollen appendage blade = the expanded part of a leaf (or petal) brachyblast = short shoot of limited growth usually borne on a long main axis bract = a modified leaf in the inflorescence, standing below peduncles and pedicels of flowers bracteate = bearing many small leaf-like structures or bracts bracteole = a secondary bract, usually smaller than the bracts and always borne above these branch = a lateral division of the growth axis bud = a meristem in early development or in resting stages of a plant, usually with protective coverings (Fig. 15) bulb = underground storage organ, the bud enclosed by fleshy scale-like leaves or leaf bases, as in Allium or Tulipa (Fig. 16d) bulbil = a small, usually axillary bulb or tuber capable of developing into a new plant, in some cases produced on a leaf or an inflorescence axis bullate = with the surface of the leaf raised in blisters between the veins

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bundle = a strand of specialised tissue bush = woody plant with many stems and smaller than a tree (equivalent to a shrub) C C3 photosynthesis = a metabolic pathway during photosynthesis in which carbon is fixed by converting a 5-carbon sugar (ribulose biphosphate) and CO2 into 3-phosphoglycerate C4 photosynthesis = a metabolic pathway during photosynthesis in which CO2 is incorporated into a 4-carbon organic acid, which has the ability to regenerate CO2 in chloroplasts aggregated in bundle sheath cells that then utilise this CO2 to generate carbohydrates the conventional way, but saving water in the process caespitose = growing in tight clumps calceolate = slipper-shaped calyculate = having bracts around the calyx or with an involucre resembling an outer calyx calyptra = a cap covering a flower or fruit calyx = the outermost whorl of floral organs, often divided into free or fused sepals CAM photosynthesis = crassulacean acid metabolism is a carbon fixation pathway specific to some plants growing under arid conditions. These plants collect CO2 only during the night which is stored as a 4-carbon malate; during the day, when stomata are closed, this is transported to chloroplasts where it is converted to CO2 and then used for conventional photosynthesis (C3). cambium = layer of growing tissue that produces new cells canopy = uppermost layer of vegetation of woodland or forest capitate = head-like capitulum = a compact cluster of more or less sessile flowers (Fig. 28j) capsule = a dry dehiscent fruit composed of two or more united carpels, opening by valves, slits or pores (Fig. 35m-s) carnivorous = trapping animals and deriving some or most of the nutrients for the plant by digesting them caruncle = an outgrowth of the seed, usually fleshy and associated with animal (especially ant) dispersal catkin = a slender, pendent cylindrical raceme or spike composed of crowded, sessile apetalous flowers (Fig. 28l) caudate = with an apex abruptly ending in a long tail-like tip or appendage, excessively acuminate (Fig. 25n) caulescent = with an evident stem above ground

cauliflorous = with flowers and fruits emerging from the stem or trunk (Fig. 14) cauline = arising from the stem centripetalous = developing from the margin towards the middle cereal = a grass of which seeds are used for human consumption cernuous = nodding chamaephyte = a plant for which the growing point survives adverse seasons with a resting bud at or near the ground character = single technical difference used to distinguish between taxa; a feature of a plant chlorophyll = the green pigment in plant cells that facilitates photosynthesis ciliate = bearing a fringe of hairs along the margins (Fig. 26c) cincinnus = inflorescence with flowers appearing alternately to the right and left on one side of the axis, a scorpioid cyme, or a situation in which each successive flower arises in the axil of a bract on the preceding flower stalk circinnate = coiled inwards upon itself circular = round circumscissile = opening by a slit running around the circumference with the upper part coming off as a lid (Fig. 35s)

Figure 14. Cauliflorous

GLOSSARY

Figure 15. General flowering plant morphology

flower fruit

pedicel

inflorescence

peduncle

vein leaf

blade

petiole

bud

infructescence

node

internode

stem

rhizome

roots

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GLOSSARY Figure 16. Underground structures. [a] rhizome. [b] tuber. [c] corm. [d] bulb.

a cirrhose = with tendrils (Figs. 18g, 25o) clade = an exclusive group of plants in which each member is more closely related to the others than any member is to a member of another group cladode = node or internode of stem or branch that is flattened and expanded to serve the functions of a leaf classification = ordering of taxa in specialised categories based on perceived relationships clathrate = pierced with holes, like a lattice (usually used for fern scales) clavate = club-shaped claw = the narrow, stem-like proximal part of a petal cleft = divided almost to the middle (Fig. 25a) cleistogamous = self-fertilising in unopened flowers clone = a plant resulting from vegetative reproduction that is genetically identical to the parent coalescent = partially and irregularly fused cochleate = spiral, like a snail shell coenosorus = a group of fern sori that have coalesced to look like a single large sorus columella = persistent central axis around which fruit locules are arranged column = the adnate styles and stamens forming a solid central body in Orchidaceae or the tube of anther filaments in Malvaceae coma = a tuft of long hairs on a seed that aids in wind dispersal comose = bearing a tuft of hairs complanate = flattened compound = composed of several similar parts, in fruits derived from several flowers, of inflorescences and leaves when there are several orders of branching cone = an inflorescence or fruit with the scales overlapping, like the fruit

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b

c

of Magnoliaceae or the seed-bearing structure in a gymnosperm (Fig. 35l) confluent = coming together and merging congested = crowded conglomerated = clustered conical = cone-shaped connate = fused or united with other surfaces of the same type (as opposed to adnate) connective = the part of a stamen between the anther cells distinct from the filament and forming conspicuous structures in some families conocarpium = a multiple fruit composed of many fruits on a single receptacle (like a strawberry) (Fig. 35f) conspicuous = standing out, remarkable contiguous = without interruption, adjacent and touching contorted = of a flower bud when each petal overlaps its neighbour on one side and is overlapped by its neighbour on the other side convolute = of a flower bud when petals are rolled in along their length and overlap, each segment enveloping the next cordate = heart-shaped; when used for a leaf base, it is deeply notched so that the base has a heart-shape (Figs. 21n, 23f) corm = short underground swollen stem (not the leaf bases) for storage (Fig. 16c) cornute = horned corolla = the second whorl of floral organs inside or above the calyx and below the stamens, consisting of free or fused petals corona = series of appendages on the corolla or stamens or at the junction of the corolla tube and corolla lobes (as in Narcissus or Passiflora) corymb = a flat-topped, racemose inflorescence in which the branches or the pedicels start from different points but all reach roughly the same level

d cosmopolitan = occurring (nearly) all over the world costa = a rib, often of a leaf or leaflet, sometimes used for the midrib costapalmate = a palmate leaf in which the petiole extends into the lamina forming a midrib (only used in Arecaceae) cotyledon = seed-leaf, the first-formed leaf that is often of a different shape from later leaves coumarin = an organic chemical (a benzopyrone) that smells of freshly cut grass creeper = plant with stems running along the ground that may root at various points along their length crenate = notched with regular rounded (scalloped) teeth (Fig. 26e, g) crenulate = with small teeth crested = with an elevated irregular ridge crispate = curled or ruffled (Fig. 26l) cristate = with a crest or a band of stiff hairs cryptogam = a plant without stamens, pistils and seeds but reproducing sexually; this term previously included ferns, lycophytes, mosses, algae, fungi and even some marine animals cryptophyte = plant with a growing point that survives adverse seasons as a resting bud below the ground or submersed in water cultigen = a species known only from cultivation cultivar = a cultivated variety of a species cuneate = wedge-shaped, tapering gradually (Fig. 23b) cupule = a cup-like structure at the base of fruits, formed by the dry enlarged floral envelope, as in Lauraceae and Quercus (Fagaceae) cuspidate = abruptly terminating with a sharp rigid point (Fig. 25l) cuticle = outer waxy layer on the epidermis

GLOSSARY composed of a fatty water-repellent substance cyathium = a cup-shaped structure, in ferns an indusium completely surrounding the receptacle, in Euphorbiaceae an involucre on which the flowers sit, formed from fused bracts cylindrical = like a cylinder: a long and narrow shape with a circular cross-section cyme = a sympodial inflorescence in which the terminal flower opens first with axillary flowers following; this can be simple or a compound dichasium (Fig. 28g-j) D deciduous = with parts falling off seasonally, in the case of leaves, not evergreen decumbent = lying on the ground but with the tip upright decurrent = extended downwards, such as leaves that grow down along the stem decussate = with organs on opposite sides of a stem or other axis in which the pairs above each other are at right angles definite = with finite growth or fixed in number dehisce = to open when mature dehiscence = the mode of opening deltate = shaped like a triangle with equal sides dentate = toothed, with the projections sharp and the sinuses rounded (Fig. 26h) denticulate = finely toothed depauperate = poorly developed depressed = more or less flattened or indented at the top derived from = evolved from and somewhat different from one or the other taxon descending = growing downwards determinate = with finite growth dichasium = a peduncle with a terminal flower and two stalked flowers in its axils (Fig. 28i) dichotomous = forking, each branch dividing into two equal branches dicot = flowering plant in which the embryos have two seed leaves didynamous = with two pairs of stamens of different lengths differentiation = development into several forms digitate = finger-like, for instance when leaflets emerge from a single point dilated = expanded dimerous = with parts in pairs dimorphic = of two shapes or forms dioecious = having unisexual flowers with the sexes on different plants disc floret = the radially symmetrical flower of an Asteraceae inflorescence, different

from the bilaterally symmetrical ray florets discoid = orbicular, rounded or partially circular discolorous = with two colours discrete = separate disjunct = with widely separated distributions disk = a flat, round organ, often an enlarged part of an ovary that excretes nectar dispersal = the mode of distribution of propagation units dissected = divided into segments dissimilar = unlike distant = not overlapping, free distichous = organised in two opposite rows, usually in a single plane (Fig. 20d) distribution = the geographical occurrence of an organism diurnal = flowering during the day divaricate = widely spreading or branching divergence = gradual separation, used usually in an evolutionary context divided = of a structure that is split into two or more units domatium = a small cavity in the axils of leaf veins or on stems, shoots or roots that is linked to harbouring arthopods, such as ants or mites dominant = the most common or prominent plant in a vegetation type or region dormant = sleeping, not active dorsal = the back side, referring usually to the upper side of structures dorsifixed anthers = where the filament is attached between the base and the apex of the anther (Fig. 32) dorsiventral = with two surfaces, upper and lower drepanium = a sickle-shaped (scorpioid) cyme, branching always to the same side, the axis often forming a coil (Fig. 28h) drip-tip = a tip of a leaf that is curved downwards and from which water can drip; common in tropical rainy climates dropper = a shoot from a bulb, corm or tuber that grows down and forms a new bulb, corm or tuber at its end drupe = a stone fruit, a fleshy fruit with usually a single seed (but sometimes more) enclosed in a hard endocarp (Fig. 35a) dye = a colouring substance derived from plants E ebracteate = without bracts echinate = prickly, with small sharp projections ecology = the study of the interaction between organisms and the organism with its environment edaphic = related to soil conditions

edentate = without teeth effuse = loosely spreading efoliate = without leaves eglandular = without glands ejaculator = a structure that actively shoots pollen or seeds away ellipsoid = of a three-dimensional oval shape elliptic = oval shaped: broadest in the middle with two equal rounded ends (Fig. 21g) elongate = long, stretched emarginate = with a sharp notch (Fig. 25c) embryo = an undeveloped plant still inside the seed emergent = arising from endangered = in danger of extinction endemic = occuring only in, restricted to or unique to a certain area endocarp = the innermost layer of a fruit wall endosperm = the food storage tissue in a seed ensiform = sword-shaped: long, narrow and ending in a sharp tip (Fig. 21d) entire = undivided and without teeth, serrations etc. (Fig. 26a) environment = the conditions that surround and may influence a plant ephemeral = a short-lived plant or structure of a plant epicalyx = a whorl of bracts below the flower that resembles a calyx epicarp = the outermost layer of a fruit wall epidermis = the outermost layer of cells on an organ of a plant epigynous = atop the ovary, when the other flower parts are placed above the ovary (e.g. a flower with an inferior ovary) epilithic = growing on rocks epipetalous = fused with the petals epiphyllous = of an organ growing atop a leaf, as in the inflorescence of Helwingiaceae and Phyllonomaceae

a

b

Figure 17. Defence structures. [a] thorn. [b] spine.

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GLOSSARY

a

c

b

d e

f

h g

i

Figure 18. Means by which some plants climb. [a] by winding stems. [b] by adventitious roots. [c] by winding rachises and petioles (Clematis). [d] by tendril opposite a leaf (Vitaceae). [e] by tendrils 90° to the petiole (Cucurbitaceae). [f] by tendrils from the leaf axils (Passifloraceae). [g] by cirrhose leaf tips. [h] by tendrillate leaf tips. [i] by petiolar tendrils (Smilacaceae).

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GLOSSARY epiphyte = a plant growing on (and usually attached to) another plant only for support and not deriving nourishment from the host episepalous = fused with the sepals epistemonous = fused with the stamens epithet = the second part of a scientific species name equilateral = equal-sided equitant = with the base of the leaf clasping the stem or the leaf above it and which in turn clasps the next leaf above; not referring to a leaf that is unifacial or laterally flattened erect = upright erose = irregularly toothed (Fig. 21zd) ethnobotany = the study of the uses of plants in human culture etiolated = with pale leaves and elongate internodes due to lack of light eudicots = one of the major clades of flowering plants composed of the majority of the classical dicots; those with pollen of three pores (tricolpate) or derived from this condition eutrophic = rich in minerals evanescent = slowly disappearing evergreen = retaining leaves throughout the year and during adverse seasons evolute = unfolded or turned back evolution = the cumulative change in characters of a group of organisms over time excavated = hollowed out excentric = off-centre, not in the middle excurrent = with the main part in the middle of the other, much smaller or less welldeveloped parts surrounding it excurved = curved away from the middle exindusiate = without an indusium, in ferns exine = the outer wall of a pollen grain exstipulate = without stipules F falcate = curved like a sickle or scythe (Fig. 21t) false indusium = the reflexed leaf margin covering the sorus in ferns false vein = a line of elongate cells across the leaf surface giving the appearance of a vein but not connected to the vascular tissue of the leaf (Fig. 27d) family = a higher taxonomic unit composed of a clade with one or more genera, usually distinct from other families in morphological characters farinose = covered in flour-like powder fasciated = abnormally flattened and widened, often the result of abnormal fusion of parts fascicle = a cluster of similar organs arising from one point

fastigiate = bundled, erect and closely parallel structures emerging from one point female flower = a flower with only female parts, lacking or having sterile rudimentary male parts (staminodes) ferns = major clade of vascular plants, composed of plants lacking flowers and seeds but instead having simple or complex leaves that bear spores, which germinate to produce sexual plants (usually minute) that produce male and female gametes and produce fernlings after fertilisation fern allies = colloquial term used to describe other seedless vascular plants that are not ferns, generally now considered simply as ferns or lycopods fertile = bearing flowers, fruits or spores fertilisation = the result of a male gamete reaching a female gamete, in seed plants a pollen tube reaching an ovule, leading to the fusion of gametes and production of a new individual fiddlehead = the coiled immature leaf with the apex in the centre, usually of an emerging fern leaf filament = a stalk that bears the anther (Fig. 29) filamentous = thread-like filiform = thread-like, slender fimbriate = fringed, bordered by hair-like processes fissure = deep and narrow split fistulose = long, round, hollow and closed at both ends (Fig. 21b) flabellate = fan-shaped (Figs. 21u, 27g) flaccid = limp flagellate = whip-like flange = a ring-like projection around a cylinder or globe flavones, flavonoids = natural yellowpigmented ketone-containing compounds in plants and fungi fleshy = swollen, succulent flexible = easily bending but readily springing back to the original position flexuose = bent alternatively in opposite directions, e.g. zigzag flora = all plants occurring in a certain area floret = a small flower, a flower in a flower head or a flower with its bracts floriferous = bearing flowers flower = an axis bearing one or more pistils, one or more stamens or both stamens and pistils and often with parts to attract pollinators such as petals, sepals and nectaries (Fig. 15) foetid = stinking foliaceous = leaf-like foliage = leaves

foliole = a leaflet: a division of a compound leaf follicle = a seed pod formed from a single carpel that opens along an inner suture to which the seeds are attached (Fig. 35m) foraminate = pitted with small holes forked = separating into two (or more) parts from a common base form = a slight variant, usually with a minor genetic basis foveolate = minutely pitted with small depressions fractiflex = zigzag free = not attached to other parts, not fused frond = a leaf, usually compound, often used for ferns and palms (Fig. 13) fruit = the seed-bearing organ that can also function as a seed-dispersing unit (Figs. 15, 35) fruitlet = a part of the fruit that functions as a separate seed-dispersing unit frutescent = becoming shrubby fruticose = shrubby funicle = the stalk of the ovule or seed funnel-shaped = of a tubular shape that widens abruptly to a wider distal part furcate = forked with sharp tips furrowed = grooved, with longitudinal channels fused = joined together fusiform = spindle-shaped, thick in the middle but tapering abruptly to both ends G galbule = a cone of a conifer that becomes fleshy when it matures as in Ephedraceae and Cupressaceae gamete = unisexual body that when fused with a gamete of the other sex, then together produce a zygote, which then gives rise to a new organism gametophyte = the stage of a plant that forms the sexual organs in seed plants, ferns, lycopods and mosses; in seed plants, this stage is dependent on and enclosed by the sporophyte; in ferns and lycopods, the two stages live independently from each other; and in mosses, the sporophyte is dependent on the gametophyte gemma = adventitious bud on a frond that can develop into a new plant genera = plural of genus generation = age group, or in plants, a sexual or asexual phase geniculate = bent like a knee or with a kneelike structure genus = a group of similar and related species, usually all more closely related to each other than any of the species is to a member of another genus

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GLOSSARY geocarpic = with fruits that mature underground, like a peanut (Arachis, Fabaceae) geophyte = a plant with the growing point surviving adverse seasons as a resting bud on an underground storage organ, like a bulb, corm, tuber, rhizome or root germination = the process in which a seed develops into a seedling or spore of a fern or related species develops into a gametophyte gibbous = pouched, inflated on one side, or one side more convex than the other glabrescent = nearly glabrous glabrous = without hairs or scales gland = an area or structure that secretes a substance, sometimes terminating a hair glandular = covered with glands, bearing glands glaucous = covered with a waxy blue-grey material that rubs off easily (Fig. 19) globose = spherical, round glochid = a barbed bristle glomerate = compactly clustered glomerule = a dense cluster of flowers glume = a bract usually occurring in pairs at the base of a grass or sedge spikelet (inflorescence) gnarled = twisted Gondwana = the supercontinent in the Southern Hemisphere formed after the breakup of Pangaea 200 million years ago; it persisted until 100 million years ago, when it split and produced the land masses now called South America, Africa, Arabia, India, Madagascar, Australia, New Zealand, New Caledonia and Antarctica grade = a group of plants that do not form a monophyletic group, but branch off the main lineage sequentially and independently

Figure 19. Glaucous

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grain = a small rounded body about the shape and size of a rice grain, more generally used for the fruit of a cereal granular = covered with small grains grooved = with long narrow indented lines growing point = the apex of a growing stem or the place where cell division takes place growth ring = an annual ring of growth that can be seen in wood guttation = secretion of water from a plant through glands (hydathodes) on leaves gymnosperm = major clade of seed plants in which the developing ovules and seeds are produced in a cone-like structure and not enclosed in an ovary but rather are produced nakedly gynandrium = structure in which the stamens are fused to the pistil gynobasic style = the situation in which the style arises in the centre at the base of the ovary as in many Lamiaceae, Boraginaceae and Allium (Amaryllidaceae) gynodioecious = dioecious plants, with some plants bearing bisexual flowers and others female flowers gynoecium = the female element of a flower, the pistil, consisting of an ovary, style and stigmas gynomonoecious = a plant with both female and bisexual flowers gynophore = a stalk carrying the ovary gynostemium = a structure formed by the fusion of the androecium and gynoecium in Orchidaceae H habit = general appearance of a plant, especially the vegetative aspects habitat = the environment or vegetation type in which a species is usually found hair = an outgrowth of the epidermis consisting of one or more elongate cells halophyte = a plant that lives in saline habitats hamate = with a hook at the tip (often referring to a hair) hapaxanthic = a plant that dies after a single period of flowering and fruiting hardwood = wood from non-coniferous trees hardy = able to withstand unfavourable conditions or seasons, usually with reference to winter conditions hastate = with two triangular lobes pointing outwards (Fig. 21p) hastula = a flange of tissue where a palmate or costapalmate leaf joins the petiole in Arecaceae haustorium = a structure by which a parasitic plant anchors itself to and extracts nutrition from its host head = a short dense inflorescence

helicoid = spiralling in three dimensions heliophyte = plants with adaptations to grow in full sun heliotropic = turning towards the sun helophyte = herb with basal parts in the water or mud and upper parts above the water, or a plant that survives adverse seasons as a resting stage in swampy ground hemicryptophyte = a plant that survives adverse seasons as a resting bud at or near the soil level or as rosettes or tussocks hemi-epiphyte = a plant that is epiphytic during part of its life cycle but is rooted in the soil for part of its life hemispherical = shaped like half a globe herb = a plant without a persistent woody stem above ground herbaceous = with the texture of a herb or a plant with non-woody structures herbarium = a collection of dried plants that can be used for taxonomic and other studies hermaphrodite = bisexual plant hesperidium = a fleshy berry with a leathery rind, the fleshy part divided into segments, as in Citrus (Rutaceae) heterocarpous = with fruit of more than one kind heterogamous = with flowers of more than one kind heterogeneous = not uniform heterophyllous = with leaves of more than one kind heterosporous = with spores of two kinds heterostylous = with styles of more than one kind in a flower or in different flowers on one plant, a phenomenon called heterostyly heterotrophic = a plant that obtains starch from sources other than photosynthesis hilum = the scar left on the seed from its attachment to the placenta hip = a false fruit developed from the hypanthium (floral) in Rosa (Rosaceae) (Fig. 35d) hippocrepiform = horseshoe-shaped hirsute = with coarse stiff hairs hispid = with long stiff hairs, sharply bristly holophyte = a plant that produces its own food through photosynthesis homogamous = having flowers of the same kind homogeneous = uniform homologous = with a shared evolutionary origin, the structure of two things does not have to be similar in form to be homologous (e.g. the forelimbs of a horse and the wings of a bat) homophyllous = with leaves of one kind homosporous = with spores of one kind homostylous = with styles of one kind in a flower

GLOSSARY

a c

b

f

d

e

i

h

g

Figure 20. Leaf attachment. [a] alternate. [b] opposite. [c] whorled (verticillate). [d] distichous (in a plane). [e] resupinate (up-side down). [f] amplexicaul. [g] peltate. [h] perfoliate. [i] ligulate

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GLOSSARY honeyguide = a set of coloured streaks and/ or blotches on the perianth of a flower that point to the nectary host = the plant on which a parasite or epiphyte grows humifuse = ground covering humus = layer of organic matter resulting from decomposition of organisms and their parts hyaline = nearly transparent hybrid = a plant that is the product of a cross between two (or more) species hybridisation = the process of crossing between two (or more) species hydathodes = glands that exude water, often seen as broadened veins, a term frequently used in ferns hydrochory = distribution of propagules, pollen or gametes by water hydrogeophyte = a plant surviving adverse seasons with its resting buds under water hydrophyte = a plant surviving adverse seasons with resting buds under water and underground in mud hygrophilous = moisture-loving hypanthium = cup-shaped enlargement of the flower base surrounding the ovary and carrying the perianth above or part-way above the ovary hypocotyl = the part of a seedling below the cotyledons and above the roots hypodermis = layer of cells below the epidermis hypogeal = below the surface of the soil hypogynous = with the perianth placed below the ovary, the ovary thus being superior I idioblast = a cell differing from surrounding cells in shape or function imbricate = the parts overlapping like roof tiles immersed = fully sunken into imparipinnate = oddly pinnate, a pinnate leaf with a single terminal leaflet (Fig. 22g, h) imperfect flower = a flower with one of the whorls missing, often the stamens or carpels thus making the flower unisexual inaperturate = of pollen, without an opening incertae sedis = literally, of uncertain seating: a group of uncertain taxonomic placement incised = deeply cut included = not emergent from the surrounding organ included veinlet = small vein that terminates in an areole formed by other veins incomplete = missing a part inconspicuous = not clearly visible or easily perceived

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incrassate = thickened incurved = bent inwards indefinite = continuous or numerous indehiscent = not splitting open indented = dimpled, marked with a dent indeterminate = continuous, not clearly defined indument, indumentum = the covering of plant parts, usually hairs or scales induplicate = curved inwards but not overlapping or v-shaped in cross-section, usually with reference to margins indurate = hardened indusium = a thin flap of tissue covering a sorus, present in some ferns (Fig. 13) inequilateral = with two uneven sides inferior ovary = an ovary that is placed below the perianth insertion (Fig. 33c) inflated = swollen as a balloon inflexed = curved inwards inflorescence = part of the plant that bears the flowers including bracts and peduncles (Figs. 15, 28) infrageneric taxa = taxonomic categories below the rank of genus (e.g. subgenus, section, species, subspecies, variety) infrapetiolar = between the petioles on the stem infraspecific taxa = taxonomic categories below the rank of species (e.g. subspecies, variety, forma) infructescence = part of the plant that bears the fruit/seeds (Fig. 15) infundibular = funnel-shaped inrolled = rolled toward the centre of a structure, usually in reference to margins insectivorous = obtaining part of the required nutrition by catching and digesting insects inserted = placed within integument = a layer covering an organ interaxillary = between axils interfoliar = between two opposite leaves intergeneric hybrid = a hybrid between species that belong to different genera internodal = between two nodes internode = the stem between two nodes interpetiolar = placed between the two petioles of a pair of opposite leaves interrupted = with a break in continuity interspersed = scattered among intrafloral = within the flower intramarginal = near the margin intrapetiolar = between the petiole and the stem intricate = tangled introrse = opening inwards invaginated = enclosed by a sheath involucrate = with a cluster of bracts involucre = a series of closely placed or fused bracts below an inflorescence iridescent = with an oily or metallic gloss,

referring to the rainbow colouring it may produce isomerous = with equal numbers of parts in a whorl isophyllous = with leaves of one kind J joint = an articulation juvenile = young, immature K keel = a narrow ridge protruding from a round surface, more commonly used for the two (partially) united lowest petals in Fabaceae that conceal the stamens and pistil keiki = a shoot that can grow roots while still attached to the stem and become independent of the mother plant, a term exclusively used in Orchidaceae Kew = village in Richmond, United Kingdom, where the Royal Botanic Gardens are situated, where the authors of this book work and where many of the photographs in this book were taken knee root = a root that protrudes above the ground, usually occurring in species that grow in waterlogged soils L labellum = lip, the lowest and usually largest petal in Orchidaceae or the largest of the three petaloid stamens in Cannaceae labiate = with lips, usually when in zygomorphic flowers the corolla is divided into an upper and a lower part lacerate = torn, irregularly lobed laciniate = dissected into slender lobes or elongate teeth lactiferous = milk-producing, with milky sap laevigate = polished, smooth lamella = thin structure, like a plate lamina = expanded part of a leaf blade or perianth lanate = woolly lanceolate = narrowly ovate, widest towards the base and tapering to a pointed tip (Fig. 21e) lanose = woolly lateral = on the side latex = sticky milky sap laticiferous = latex producing Laurasia = the supercontinent of the Northen Hemisphere combining North America and Eurasia, formed after the breakup of Pangaea 200 million years ago when it separated from Gondwana lax = loose, not close together leaf = chlorophyll-containing organ growing out from the stem, subtending an axillary bud (although in some cases chlorophyll is secondarily lost)

GLOSSARY leaflet = a part of a compound leaf leaf scar = mark on the stem where the leaf was attached before falling off leaf sheath = part of the leaf that envelops the stem legume = the fruit of species of Fabaceae (Fig. 35n) lemma = the outermost bract enclosing a grass flower lenticel = corky structure on bark that allows exchange of gasses lenticular = in the shape of a lens, a three dimensional form that is circular from one side but convex towards the margins lepidote = clothed with scales leptosporangiate = with sporangia developed from a single cell, as in many ferns liana = woody climber lianescent = becoming climbing ligneous = woody ligule = projection of a leaf sheath, commonly used in the identification of Poaceae. In lycopods it is a small outgrowth at the base of a fertile leaf (Fig. 20i) linear = like a line, narrow and much longer than wide and with the margins running parallel (Fig. 21c) lingulate = tongue-shaped lip = one of the two parts of a zygomorphic bilabiate flower, or the labellum in Orchidaceae litter-trapper = plant with special structures to catch litter and gain nutrients lithophyte = plant growing on rocks littoral = growing along a shore of the sea, a lake or a river lobe = a part of a leaf with a deep sinus separating it from other parts of the leaf lobule = a small lobe, often on a lobe locule = a compartment, the cavity of the carpel in which the ovules are formed or the compartment of the anther in which the pollen is formed loculicidal = a capsule that splits into independent locules (Fig. 35p) lomentum = a flat fruit that is constricted between each seed (Fig. 35o) lunate = shaped like a crescent moon lustrous = glossy lycophyte = colloquial term used sometimes incorrectly instead of lycopod lycopod = major clade of seedless vascular plants sister to all other vascular plants (ferns+gymnosperms+flowering plants). The clade is distinguished in having non-specialised leaves called microphylls and simple vascular structures and consists of Isoëtaceae, Lycopodiaceae and Selaginellaceae. lyrate = lyre-shaped, with a large terminal lobe and smaller lateral lobes at the base

M maculate = spotted Malpighian hair = a T-shaped hair, with the base of the stalk attached near the middle mammillate = with nipple-like or breast-like processes mammiform = breast-like, conical with a rounded tip mangrove = coastal swamp in the tropics that is regularly inundated with tidal salt water, usually with trees and other plants adapted to growing in such conditions manicate = with a dense covering of hair that can be peeled off in its entirety margin = edge or boundary of a leaf or other organ matted = with hairs that are intertwined maturation = ripening medifixed = attached at or near the middle megaphyll = a large leaf, usually used for the complex leaves of ferns and seed plants that are not microphylls megasporangium = a sporangium in which megaspores develop megaspore = the larger size of spores in heterosporous plants megasporophyll = one of the ovule-bearing scales in the female cone of a gymnosperm membrane = a thin layer mericarp = part of a fruit that contains seeds and is formed of a carpel, dehiscing independently of others in the same flower when ripe mesocarp = the middle layer of a fruit mesophyte = a plant growing under conditions that are neither wet nor dry microclimate = a local climatic condition in which a plant may thrive, despite a different climate in the surrounding area microphyll = leaf of a lycopod, which has a single unbranched vein and usually simple structure micropyle = an opening in the integuments surrounding the ovule through which the pollen tube may enter the ovule and through which the root emerges upon germination of the seed microsporangium = a sporangium that produces microspores, but is not necessarily smaller than the megasporangium midrib = the main vein of a leaf minute = at the lower end of the range of sizes that can be seen with the naked eye molecular study = DNA phylogenetics, the analysis of DNA sequences to produce a phylogenetic tree and estimate evolutionary relationships of taxa monadelphous = arranged in a bundle fused by their filaments, as for stamens

monandrous = with a single stamen moniliform = like a string of beads, a structure constricted at regular intervals forming regular globose shapes monilophyte = colloquial term often used for the fern clade including Equisetaceae and Psilotaceae, to distinguish them from pteridophyte, which typically includes the lycopods; the correct term for this clade is ‘the ferns’ monocarpic = flowering and fruiting only once and then dying monochasium = an inflorescence type with a single terminal flower and a single lateral flower emerging from a bracteole. A compound monochasium, where the laterals branch further in the same fashion is called a cincinnus. monocot (or monocotyledon) = flowering plant in which the embryos have one seed leaf, often recognised by their trimerous flowers and parallel veins although there are many exceptions monodynamous = with one stamen much larger than the others monoecious = with all flowers on a plant bisexual or with separate male and female flowers on the same plant monogeneric = a taxon (or group) with a single genus monolete = a spore wall with a single linear aperture monomorphic = of one kind only monophyletic = of a group in which all taxa are more closely related to each other than any one is to a member of another group monopodial = with a main stem that grows indefinitely at the tip with the shoots formed laterally monospecific = of a taxon (or group) consisting of a single species monosulcate = with a single furrow-like aperture, as in pollen monothecal = with a single anther cell monotypic = consisting of a single species montane = growing in the mountains or pertaining to mountainous regions morphological = of shape and form morphology = shape and form of organisms, both internal and external motile = actively moving mouth = part where a tubular organ or structure opens mucilage = slime or jelly exuded by a plant, composed of polysaccharides mucro = a short sharp tip mucronate = ending abruptly in a short sharp tip (Fig. 25j) multicellular = composed of two or more cells multicolpate = of pollen with many linear apertures

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GLOSSARY

a

c

b

g

k

h

e

d

i

l

f

j

m

n

Figure 21. Leaf shapes. [a] acerose (needle-like). [b] subulate/fistulose (round and hollow). [c] linear. [d] ensiform (sword-like). [e] lanceolate. [f] oblanceolate. [g] elliptic. [h] oval. [i] ovate. [j] obovate. [k] oblong (rectangular). [l] orbicular (round). [m] spatulate (spoon shaped). [n] cordate (heart-shaped). [o] sagittate (arrow-shaped). [p] hastate. [q] praemorse. [r] obliquely cordate. [s] peltate. [t] falcate (sickle-shaped). [u] flabellate (fan-shaped). [v] palmate. [w] tendrillate. [x] pinnatisect/pinnatifid. [y] pinnately lobed. [z] palmatisect/palmatifid. [za] palmately lobed. [zb] sinuately lobed. [zc] spinose. [zd] erose. [ze] branch with imbricate, scale-like leaves.

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GLOSSARY

p

o

s

r

u

t

x

zb

q

y

zc

v

z

w

za

zd

ze

Plants of the World

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GLOSSARY multifid = split into numerous lobes multiseriate = arranged into many rows muricate = bearing short hard protuberances mutation = a spontaneous change in genetic material, often resulting in a change of characteristics mutualism = an association between two organisms that benefits both mycoheterotrophic = extracting nutrition from a fungus and hence indirectly from other organic matter, rather than producing energy through photosynthesis mycorrhiza = symbiotic or mutualistic fungus associated with or inside the roots of plants mycotrophic = obtaining some or all nutrition via an interaction with mycorrhizal fungi myrmecochorous = dispersed by ants myrmecodomatia = domatia (hollows) inhabited or visited by ants myrmecophyte = a plant associated with ants often with specialised structures to feed or house them N naked = without other organs (leaves, scales, hairs, indusium or perianth) napiform = turnip-shaped, used usually for roots that are broadly ovoid and taper to a point naturalised = non-native, introduced by man and becoming established such that the plants reproduce freely in their new habitat navicular = boat-shaped Nearctic = the northern part of North America nectar = sugary fluid excreted by glands to attract pollinators nectar guides = coloured streaks or blotches on the perianth that point to the nectary nectary = an organ in which nectar is formed neophyte = a plant newly introduced to an area neoteny = the condition in which plants become fertile when still vegetatively in a juvenile stage Neotropics = the tropical area of the Americas from North America (Mexico and southern Florida) and the Caribbean south to Bolivia, Paraguay, northern Argentina and southern Brazil nested within = of a taxon or clade placed within another clade in a phylogenetic tree nest leaves = leaves that collect litter net-veined = with smaller veins connected forming a mesh-like network nitrogen fixation = the process in which bacteria (or other organisms) take

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atmospheric nitrogen and convert it into compounds that can be taken up by plants. This process often takes place in specialised nodules. nocturnal = night-flowering nodding = bending downwards node = the area of a stem where a leaf is or would have been attached (Fig. 15) nodule = a knob on a root or leaf that contains (nitrogen-fixing) bacteria nomenclature = the application of names to organisms notched = with a nick in an otherwise entire margin nothogenus = a hybrid genus, formed when two species belonging to different genera hybridise nothospecies = a hybrid species nucellus = the central part of the ovule enclosed by the integuments numerous = many, usually more than ten nut = a one-seeded indehiscent fruit with a hard dry outer shell (Fig. 35h) nutlet = a small nut, often used in Cyperaceae for their often small hardened fruits O obcordate = heart-shaped but with the narrow end at the base and the lobed end at the tip (Fig. 25b) oblanceolate = inverted lanceolate, with the widest part towards the tip (Fig. 21f) oblate = globose with the poles flattened obligate = restricted to a certain habitat or life history strategy oblique = unequal, when the two sides are of a different size or when the organ is held at an angle to the symmetrical plane (Fig. 21r, 23i) oblong = a shape longer than broad with parallel sides and usually a rounded end (Fig. 21k) obovate = inverted egg-shaped in two dimensions, with the broadest part near the tip (Fig. 21j) obovoid = inverted egg-shaped in three dimensions, with the broadest part near the tip obsolete = rudimentary or no longer in use obtriangular = inverted triangular, with the narrow end at the base obtuse = blunt, not pointed (Fig. 25d) ocellate = with an eye-like spot ochrea = a sheath, in leaves an extension along the stem beyond the petiole, in Polygonaceae associated with their fused sheathing stipules odd-pinnate = a pinnate leaf with an uneven number of leaflets odorous = smelling, usually not pleasant

oligomerous = with fewer than the usual number of parts oligotrophic = poor in nutrients ontogeny = embryonic development of an individual opaque = occluded, not transparent operculate = with a lid operculum = a lid covering a flower, bud or fruit opposite = when two organs are placed on either side of a stem emerging from the same node, or when the organ is placed in front of another one (Fig. 20b) orbicular = globular (three dimensions), or circular (two dimensions) (Fig. 21l) order = a higher taxonomic rank composed of one to several families organ = a part of a plant serving a function orientation = place or position ornamental = cultivated for decoration rather than for food or manufacturing ornamentation = sculpturing, used for pollen or seed surfaces that can bear spines, granules, warts, tubercles, reticulations, flanges, etc. ornithophily = bird pollination orthographic error = an unintentional misspelling of a scientific name in the scientific description of a taxon osmophore = a structure producing an odour ovary = the part of the pistil that encloses the ovules and will develop into a fruit (Figs. 29, 33) ovate = egg-shaped (two dimensional), with the widest part below the middle (Fig. 21i) ovoid = egg-shaped (three dimensional), with the broadest part at the base ovule = the small body that contains the female germ cell of a plant and becomes a seed after fertilisation P pachycaul = a thick-stemmed and sparsely branched tree or bush with a bottle-shaped trunk pachyphyllous = with thick and fleshy leaves paedomorphic = retaining juvenile characteristics when mature Palaeotropics = the tropics of the Old World: Africa, Asia, Australia and the Pacific palate = a projection on the lower lip enclosing the mouth as found in Lentibulariaceae and some Plantaginaceae palea = a chaffy scale, in particular in Poaceae used for the inner of the two bracts enclosing a floret palmate = lobed or compound with all lobes or leaflets originating from a central point (Figs. 21v, 22e, 27f) palmately veined = when the main veins of a leaf orginate from a central point (Fig. 27f)

GLOSSARY

c

b

a

d

g

f

e

k

i h

j

Figure 22. Leaf division. [a] simple. [b] bifoliate. [c] trifoliate/ternate. [d] quadrifoliate. [e] palmate. [f ] biternate. [g] (impari-)pinnate (few pinnae). [h] (impari-)pinnate (many pinnae). [i] paripinnate. [j] (pari-)bipinnate. [k] (impari-)tripinnate.

Plants of the World

653

GLOSSARY

a

c

b

e

d

f g

h

i j

Figure 23. Leaf bases. [a] attenuate. [b] cuneate. [c] acute. [d] rounded. [e] truncate. [f ] cordate. [g] auriculate. [h] subpeltate. [i] oblique. [j] clasping.

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GLOSSARY palmatifid = with a simple leaf cut into a palmately lobed shape (Fig. 21z) palynology = the study of pollen grains and spores panboreal = throughout the far northern temperate zones of the world pandurate = fiddle-shaped, oblong or elliptic, but constricted in the middle and usually widest above the middle Pangaea = supercontinent that united all continents, which started breaking up roughly 300 million years ago panicle = an inflorescence with several lateral branches, each of which is branched, sometimes in which the main and the lateral branches are indeterminate (Fig. 28d) pantropical = throughout the tropical zones of the world papilionaceous = shaped like a pea flower with two keels, two wings and a standard, or belonging to Fabaceae papillae = small protuberances papillose = bearing many small nipple-like lumps pappus = a series of bristles or scales surrounding the corolla and persistent on the fruit in Asteraceae parallel = running in the same direction beside each other (Fig. 27a) paraphyletic = a group in which the closest relatives of at least one member are excluded and placed in another group paraphysis = hairs that are formed among sporangia in a sorus of a fern parasitic = connected to and deriving nutrients from another plant or fungus parenchyma = soft tissue composed of cells with thin walls parietal placentation = a form of placentation (seed positions) for an ovary in which the ovules are attached to the inner surface of the outer wall of a singlecelled syncarpous ovary (Fig. 34a) paripinnate = a pinnate leaf terminated by a pair of leaflets (Fig, 22i) parthenocarpy = the process when a fruit develops without fertilisation of the ovules, often the fruit seedless parthenogenesis = seed or plantlet development without fertilisation taking place partite = cleft or divided patent = spreading, held at right angles to the main axis paxillate = of a pinnately veined leaf with close parallel venation running from the midrib to the margin at an angle pectinate = comb-like, with close, narrow, parallel lobes pedate = similar to palmate, but the basal

lobes divided further pedicel = the stalk of an individual flower (Figs. 15, 29) peduncle = the stalk of an inflorescence or the stalk that supports the cone of gymnosperms (Fig. 15) pellucid = translucent, but not transparent peloric = unusually regular (actinomorphic) when the normal condition is irregular (zygomorphic) peltate = shield-like, rounded with the attachment near the middle of the blade (Figs. 20g, 21s) pendant = a type of jewelry that is worn around the neck, often mistakenly used for pendent (hanging) pendent = hanging penicillate = brush-shaped, with a tuft of hairs on the end of a stalk pentagonal = five-angled pepo = berry-like fruit with a thick exocarp and parietal placentation, as in Cucurbitaceae perennial = living for several years perfect flower = flower that has both male and female parts perfoliate = with the leaf surrounding the stem at the node (Fig. 20h) perforate = punctured by numerous little holes perianth = collective term for calyx (sepals) and corolla (petals) perianth tube = the lower united part of a perianth pericarp = a wall of a ripe ovary, a fruit wall or a fleshy layer surrounding a stony endocarp perigon = a perianth without differentiated whorls (used in monocots) perigynium = a membrane or sac enclosing a female flower and fruit in Cyperaceae perigynous = with perianth and stamens carried above the ovary on a floral tube (hypanthium) perpendicular = at right angles to the main axis persistent = remaining in place, not falling off petal = a single, free (or fused) unit of the second floral whorl (Figs. 29, 30) petaloid = shaped and/or coloured like a petal petalostemonous = a flower in which the stamens are fused with the corolla tube petiolate = having a leaf stalk petiole = a leaf stalk, the basal usually narrow and terete part of the leaf that carries the vascular bundles from the stem into the leaf blade (Fig. 15) petiolule = a stalk of an individual leaflet in a compound leaf phalange = a bundle of fused structures, like

stamen filaments (e.g. in Malvaceae) or fruit clusters (in Pandanaceae) phanerogam = term used for seed plants or for flowering plants phanerophyte = a plant surviving a resting period with a growing point or a bud well above the ground phenetics = the classification of taxa on the basis of overall morphological similarity phenology = study of the timing of natural phenomena phloem = the main vascular tissue in plants undertaking food transport phorophyte = a plant (host) that supports an epiphyte phyllary = one of the bracts surrounding a capitulum forming the involucre in Asteraceae and other plants with capitulate inflorescences phylloclade = a photosynthetic portion of a stem that is flattened and expanded to serve the function of a leaf (Fig. 24) phyllode = a laterally flattened photosynthetic blade, for instance the expanded petiole in some Acacia (Fabaceae) phyllopodium = a small outgrowth of the stem or rhizome to which the leaf is attached as in the fern Oleandra (Polypodiaceae) phyllotaxy = the arrangement of leaves along a stem phylogenetic = related to evolutionary history phylogeny = the actual and ultimately unknowable evolutionary history of a group that includes, among many things, derivation, time and amount and type of change over time; it is not the output of a computer algorithm, which is merely an estimate, or tree produced from morphological, chemical and DNA sequence data

Figure 24. Phylloclade

Plants of the World

655

GLOSSARY

a

b

c

e

h

l

f

i

m

d

g

j

k

n

o

Figure 25. Leaf apices. [a] cleft. [b] obcordate. [c] emarginate. [d] obtuse. [e] retuse. [f ] truncate. [g] rounded. [h] acute. [i] apiculate. [j] mucronate. [k] acuminate. [l] cuspidate. [m] aristate. [n] caudate. [o] cirrhose.

656

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GLOSSARY

a

b

e

i

c

g

f

j

d

h

l

k

Figure 26. Leaf margins. [a] entire. [b] undulate. [c] ciliate. [d] sinuate. [e] crenately lobed. [f ] glandular dentate. [g] crenate. [h] dentate. [i] spinose. [j] serrate. [k] biserrate. [l] crispate.

Plants of the World

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GLOSSARY phylum = a taxonomic rank consisting of several classes phytotelma = a water body produced by terrestrial plants, often a pitcher (Cephalotaceae, Nepenthaceae, Sarraceniaceae), bamboo internodes or tree hollows, formed by leaves that hold water, as in some Bromeliaceae, Heliconiaceae, Musaceae and Dipsacus (Caprifoliaceae) pigmented = with pigment, i.e. coloured pilose = hairy with short thin hairs pinna = leaflet of a compound, usually pinnate leaf (Fig. 13) pinnate = feathered, a leaf divided into a central axis with several lateral leaflets (Figs. 22g-i, 27b) pinnatifid = simple leaf that has the margin pinnately lobed (Fig. 20x) pinnule = a leaflet of a leaflet in a bipinnate leaf pinnulule = a leaflet of a multiply pinnate leaf pistil = the female organ of a flower, often consisting of a gynoecium or the entire ovary, style and a stigma (Fig. 29) pistillate flower = functionally female flower pitcher plant = carnivorous plant with a trapping mechanism of a hollow tube-like leaf that is filled partially with liquid including (usually) digestive enzymes, as in Cephalotaceae, Nepenthaceae and Sarraceniaceae pith = spongy tissue in the centre of a stem pitted = with small depressions placenta = the part of an ovary to which the ovules or seeds are attached placentation = the arrangement of the ovules in the ovary (Fig. 34) plantlet = a small plant formed on the leaf or stem of a mother plant pleated = with parallel folds pleiocarpic = flowering and fruiting more than once in a lifetime pleiochasium = a cymose inflorescence in which each main axis of a cyme produces more than two branches plesiomorphic = ancestral plicate = with parallel folds ploidy = the number of complete chromosome sets plumule = the shoot bud of an embryo in the seed plurilocular = with many locules pneumathode = an aerating tissue pneumatophore = erect root in mangroves or Taxodium of unknown function, or small tubes along the petioles and rachises of some Aspleniaceae pod = a dry dehiscent fruit with a firm outer layer enclosing a hollow centre with one

658

Christenhusz, Fay & Chase

or more seeds, often used for Fabaceae fruits (Fig. 35n) poikilohydrous = becoming desiccated during dormancy in the dry season but capable of rehydration when water becomes available again polar = relating to the poles of a globe (such as a pollen grain) or of the World pollen = powder-like male fertilising agent produced by the anthers of seed plants pollen mass = pollen grains clustering into a single body pollinarium = the set of pollinia produced by one or more anthers of a flower in some Apocynaceae or the pollen masses plus accessory structures in Orchidaceae pollination = the transfer of pollen to the stigma pollinator = the agent effecting pollination pollinium = a tightly-bound cluster of pollen grains that is distributed as one unit, as in some Apocynaceae and Orchidaceae polychasium = a cymose inflorescence in which each axis produces more than two lateral branches polygamous = bearing male, female and bisexual flowers on the same plant polymorphic = variable, with several forms or shapes polyphyletic = a group of mixed evolutionary origin derived from more than one ancestor polyploidy = the state of having three or more sets of chromosomes polystichous = with leaves arranged in several rows pome = an indehiscent fruit in which the receptacle or hypanthium enlarges to enclose the ripe ovary, like an apple or pear (Fig. 35e) population = all individuals of a species in a given area porate = opening by pores or having lots of small pores pore = a small hole praemorse = a tip with an abrupt and ragged ending prehensile = clasping, twining prickle = a sharp outgrowth of the epidermis, detachable without damaging the stem, as in a rose probract = small, usually glandular, leaf-like structures at the base of the peduncle in Cucurbitaceae process = a small projecting part or tissue progeny = offspring prolate = a globose shape drawn out at the ends like a lemon proliferation = production of vegetative shoots used for propagation or spreading abundantly

proliferous buds = adventitious buds on leaves, flowers, stems, peduncles or roots that are capable of rooting and forming independently living plants prominent = expanded beyond another organ propagation = multiplication by means of spores, seeds and cuttings or other vegetative means propagule = vegetatively or sexually formed body that can give rise to a new plant prophyll = one or two of the first leaves of a branch that are different from the rest of the leaves in shape or size, also used for the bract of a developing inflorescence in Arecaceae and for leaflets in the inflorescences of Poaceae and Cyperaceae prop root = root growing from the lower stem or branches that curves down and roots in the soil, as in Rhizophoraceae, Pandanaceae and many Ficus (Moraceae) prostrate = lying flat on the ground protandrous = a flower that sheds the pollen before the stigma of that same flower is receptive proteranthy = flowers developing when the plant is leafless prothallus = the gametophyte of a fern, resulting from germination of a spore and bearing the sexual organs protocorm = a nearly uniform structure formed after germination of an Orchidaceae seed that makes an association with a fungus via penetration of specific cells and from which the first true shoot and root differentiate protogynous = a flower that has the stigma receptive before the anthers open proximal = nearest to the point of attachment psammophyte = a plant growing on sandy soil pseudanthium = an inflorescence that consists of several reduced flowers together functioning as a single flower pseudobulb = an aerial storage organ resembling but not homologous to a bulb, used in Orchidaceae for swollen internodes of a stem pseudocopulation = the act of an insect attempting to mate with a specialised flower that resembles a member of the opposite sex and thereby effecting pollination pseudostem = a false stem consisting of leaf sheaths or leaf bases emerging vertically from a horizontal stem as found in Musaceae and Strelitziaceae pseudowhorl = a stem with the leaves singly per node, but the internodes being so short that it resembles a whorl

GLOSSARY

a

b

c

f

d

e

g

Figure 27. Leaf venation. [a] parallel. [b] pinnate. [c] reticulate. [d] pinnate-parallel forking with false veins. [e] anastomosing. [f ] palmate. [g] flabellate

Plants of the World

659

GLOSSARY Figure 28. Inflorescence types.

a b

c

d

e

f

Racemose inflorescences: [a] simple raceme. [b] spike. [c] spadix. [d] panicle. [e] simple umbel. [f ] compound umbel.

660

Christenhusz, Fay & Chase

GLOSSARY

i

h

g j

Cymose inflorescences: [g] simple cyme. [h] drepanium. [i] dichasium. [j] capitulate cyme.

l k m

Mixed inflorescences: [k] thyrse. [l] catkins. [m] verticillaster.

Plants of the World

661

GLOSSARY Figure 29. General flower terminology

petal

anther

stigma

stamen filament

sepal

pedicel

Figure 30. Petals free (Linaceae) and petals fused (Campanulaceae).

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Christenhusz, Fay & Chase

style

ovary (carpels)

pistil

GLOSSARY Figure 31. Flower symmetry. [a] actinomorphic (regular). [b] zygomorphic (irregular).

a

pteridophyte = colloquial term used for all seedless vascular plants, e.g. lycopods and ferns, a polyphyletic assemblage pterocarpous = with winged fruit pterospermous = with winged seed puberulent = minutely pubescent with hairs barely visible with the naked eye puberulous = with a dense covering of short soft hairs pubescence = hairiness pubescent = densely covered with fine soft hairs pulp = juicy or fleshy tissue of a fruit pulverulent = with a fine dusting of powder or flour-like material pulvinus = swelling on the petiole that aids in the movement of leaves punctate = marked with dots or glands pungent = with a tip ending in a sharp point or prickle purpurescent = turning or tinged purple pustulate = pimpled, with minute elevations putative = probable, but still debatable pyrene = the stone, a seed with a hard outer layer, in a fruit pyriform = pear-shaped pyrophyte = plant adapted to grow in areas that regularly burn pyxis = a capsule with a circumscissile dehiscence, with the top falling off like a lid Q quadrangular = square, with four edges quaternate = divided in fours quinate = divided in fives quincuncial aestivation = folding in a flower bud with two parts outside, two parts inside and with one margin inside and the other margin outside the adjacent petal

b

R raceme = an inflorescence in which the flowers are borne on pedicels along a central axis, with the terminal flower being the last to open; spikes and spadices are specialised types of racemes in which the flowers are sessile (Fig. 28a-f) racemose = resembling a raceme (Fig. 28a) racemule = a part of a compound raceme rachilla = the axis of a spikelet in Poaceae rachis = part of the main axis of a compound leaf above the petiole that bears leaflets, or the main axis in an inflorescence above the peduncle that bears flowers radial symmetry = symmetry around a central point radiate = arranged around or spreading from a central point radiation (in evolutionary biology) = the development of several species from an initial colonising event, a form of speciation in which three or more species are formed, often quickly radiatisect = cut in a radiate way, the lobes spreading like a wheel as in some species of Schizaeaceae radical leaves = leaves sprouting from the stem close to the root as in a rosette radicle = the root emerging from a germinating seed rambling = climbing in other vegetation in a free manner, not having specialised structures with which to climb ramification = branching ramiflorous = with flowers on the branches away from the leaves range = the geographical distribution of a species raphides = bundles of needle-shaped crystals of calcium oxalate

ray = one of the branches of an umbel ray floret = the bilaterally symmetrical flower of an inflorescence in some Asteraceae, different from the radially symmetrical disc florets receptacle = the expanded part of the flower stalk on which the organs of a flower are inserted; receptacles are fused in capitulate inflorescences and then the word is often used to mean the entire broadened stalk (head) on which the individual flowers develop reclinate = bending downwards recurved = bent backward reduced = less than normal in size or number reduplicate aestivation = folding in a flower bud in which the petals are touching with the edges valvate and reflexed reflexed = curved backward refuge = an area where the climate was relatively stable at times when the climate was changing elsewhere, allowing species that died out elsewhere to survive regma = capsule with three or more cells that breaks open or falls apart when ripe especially used in Euphorbiaceae, Putranjivaceae and Phyllanthaceae relictual distribution = currently restricted distribution of a taxon that used to be more widespread in the past reniform = kidney-shaped repand = with an uneven, wavy or shallowly lobed margin replum = the persistent partition between the locules of the capsules in Brassicaceae resin = hardened sap from a wounded stem that is not soluble in water resin thread = elastic, sticky threads that appear when a leaf is broken and pulled into two parts, e.g. in Cornaceae

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663

GLOSSARY resupinate = upside down (Fig. 20f), often in reference to flowers of Orchidaceae in which the lip is lowermost, achieved by twisting or bending of the ovary or the inflorescence being pendent reticulate = with a network of ridges and empty spaces in between, also used for net-venation (Fig. 27c) retrorse = abruptly pointing downwards or backwards, especially used for prickly leaf margins and barbs retuse = notched, with a rounded indentation in an otherwise entire margin (Fig. 25e) revolute = rolled or curved backwards rheophyte = a plant adapted to growing in rapidly flowing water, such as Podostemaceae rhipidium = cymose inflorescence with the branches and peduncles growing in a single plane, as in some Sambucus (Adoxaceae) and Iridaceae rhizobium = soil bacterium that fixes nitrogen and has an association with a plant rhizoid = a rootlet, a small root-like organ or a hair functioning as a root rhizomatous = with an underground stem rhizome = an underground or superficial stem, differing from a root in having nodes, growth buds or leaves. In ferns, all stem are called rhizomes, even if they form a tree-like trunk. (Fig. 16a) rhizophore = a specialised stem that bears rhizoids or roots as in Selaginellaceae and Rhizophoraceae rhombic = lozenge-shaped, the shape of an equilateral parallelogram ridged = with an elevated line rigid = stiff riparian = of river banks and lake shores robust = strong, vigorous, clear (as in certain) root tuber = thickened part of a root used for storage (Fig. 16b) rootstock = a rhizome or underground stem from which roots and shoots emerge

a

b

Figure 32. Anther attachment. [a] dorsifixed. [b] basifixed.

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rosette = a circle of closely spaced leaves, usually at ground level rostellum = a slender projection like a bird’s beak, in Orchidaceae used for a specialised portion of the stigma on the column rostrate = beaked rosulate = with the leaves at the base of the stem or having a rosette rotund = round, nearly circular rounded = curved without abrupt angles (Figs. 21l, 23d, 25g) ruderal = growing in waste places rudimentary = small and non-functional organ rugose = wrinkled ruminate endosperm = intruded endosperm in a seed, formed by the infolding of the inner layer of the seed coat, as in nutmeg runcinate = pinnatifid leaf with the lobes pointed towards the base as in Taraxacum (Asteraceae) runner = a lateral shoot that can give rise to a new individual at its tip rupiculous = growing on or among rocks rupturing = bursting irregularly S saccate = pouch-shaped sagittate = triangular in shape with two sharp basal lobes, like an arrowhead (Fig. 21o) salient = projecting forward saline = salty salverform = with a slender tube and an abruptly widening limb, like a trumpet samara = a dry indehiscent fruit with a wing that is longer than the seed, for instance, in many Malpighiaceae, Dipterocarpaceae and Acer (Sapindaceae) (Fig. 35k) saprophyte = colloquial term for a plant that obtains some or all of its nutrients from non-living tissue, but in fact these are provided by a fungus, mostly from decomposing plant tissue in the soil; the correct term is ‘mycoheterotroph’ sarcotesta = fleshy layer surrounding a seed, developed from the outer seed coat sarmentose = with long thin runners or long-creeping rhizomes or with whip-like branches savanna = dry grassland area with isolated (groups of) trees saxicolous = growing on rocks scabrous = rough to the touch, usually because of small stiff hairs scalariform = ladder-like scale = a small peltate disc, formed of a reduced leaf or an expanded hair. Also used for the parts of a cone of a gymnosperm and the degree of magnification in illustrations. scalloped = notched with regular, rounded teeth

scandent = climbing, usually without specialised structures scape = a leafless flower or inflorescence stalk scar = mark left on the stem by a fallen leaf, stem or peduncle or a seed when it is separated from the fruit schizocarp = a dry fruit splitting into its carpels or in one-seeded portions schopfbaum = a tree or large shrub that forms rosettes at the branch tips, like some woody Senecio, Lobelia, Lecythidaceae, some woody Asparagaceae, some branching or clumping palms and some tree ferns. Their leaves often collect fallen litter (see litter-trapper). sciaphyte = a plant adapted to shady environments sclerenchyma = thick-walled woody cells scleromorphic = of plants that have thick, fibrous leaves, adapted to survive dry climatic conditions sclerophyll = a thick leathery leaf scorpioid = cymose and one-sided, coiled to resemble a scorpion’s tail, as in Boraginaceae scrambler = a plant growing upwards needing support, but not attaching itself scrobiculate = minutely pitted or grooved scurfy = covered with many small scales scutate = shield-shaped sebaceous = fatty secreting = producing by glands section = taxonomic rank between subgenus and species secund = one-sided seed = the structure produced from a fertilised ovule, in which the embryo is protected and can function as a propagule for dispersal of the plant seed coat = the outermost layer of a seed seedling = juvenile plant, produced by germination of a seed segment = the smallest division of a compound leaf segregate = split off self-fertilising = of an ovary fertilised by pollen of the same flower or from another flower on the same plant semi-inferior = with the ovary only partly below the level of the perianth attachment (Fig. 33b) semperflorous = flowering throughout the year senescent = aging, slowly dying sensitive = reacting to touch by moving, as in Biophytum sensitivum (Oxalidaceae), Dionaea muscipula (Droseraceae) and Mimosa pudica (Fabaceae) sepal = one part of the outermost whorl of a flower, part of the calyx, often green and protecting the flower in bud (Fig. 29)

GLOSSARY Figure 33. Ovary position. [a] superior. [b] semi-inferior. [c] inferior.

a

sepaloid = resembling a sepal septicidal = splitting along the junction line of a carpel, the valves remaining attached septifragal = splitting along the junction line of a carpel, the valves falling off septum = a partition in a fruit or ovary sericeous = with closely appressed soft, silky hairs serotiny = a type of dehiscence in which a cone or fruit only releases the seeds after fire serrate = saw-toothed, with sharp tips and sinuses (Fig. 26j) serrulate = finely saw-toothed sessile = not stalked, attached directly to the stem seta = a bristle or stiff hair setose = clothed with bristles shaggy = covered with long rough coarse hairs sheath = a tubular or half-tubular organ enveloping or clasping another organ sheathing = enveloping and enclosing shoot = an elongate stem shrub = a self-supporting woody plant branching near the ground with several stems from the base silica bodies = crystals of silica found inside cells silique = a fruit divided into two cells by a thin partition and opening by two valves that fall away from the centre on which the seeds are borne. It is common in

c

b

Brassicaceae and some Papaveraceae. (Fig. 35r) simple = undivided, with only one order of branching or resulting from a single ovary (Fig. 22a) sinuate = wavy, with a margin that curves in and out with rounded lobes and sinuses (Figs. 21zb, 26d) sinus = the area between two teeth or two lobes sister groups = two monophyletic groups of taxa that are each other’s closest relatives slender = long and thin smooth = not rough or hairy solobiferous = clump-forming, making shoots at or just below ground level solenostele = a vascular system with a central core surrounded by rings of phloem, xylem and another ring of phloem sorus = a structure bearing sporangia in ferns (Fig. 13) spadix = a racemose inflorescence with a fleshy axis in which the flowers are partially sunken as in Araceae spathe = a sheathing bract surrounding the inflorescence, as in Araceae, Arecaceae, Amaryllidaceae and Commelinaceae (Fig. 28c) spathella = a closed thin sac that envelops an undeveloped flower in Podostemaceae spatheole = a bladeless sheath surrounding an inflorescence of Poaceae

Figure 34. Placentation. [a] parietal. [b] axile.

a

b

Plants of the World

665

GLOSSARY

b

c

a

e

d f

g

h

i

Figure 35. Fruit types. [a] drupe. [b] berry. [c] aggregation of drupes. [d] hip. [e] pome. [f ] pseudocarp/conocarpium. [g] syconium. [h] nut. [i] achene. [j] cypsela. [k] samara. [l] cone. [m] follicle. [n] legume or pod. [o] lomentum. [p] loculicidal capsule. [q] porose capsule. [r] silique. [s] circumcissile capsule.

666

Christenhusz, Fay & Chase

GLOSSARY

j

k

l

m

o

p n

q

r

s

Plants of the World

667

GLOSSARY spatulate = spoon-shaped, oblong with an extended basal part (Fig. 21m) speciation = process that results in formation of new species species = unit of classification, consisting of a group of populations with similar morphology and constant distinctive characters, usually capable of interbreeding and producing offspring (although there are many other definitions) species-rich = a genus or geographic area that includes many species specimen = a part of a plant or an entire plant, either dried in a herbarium or living in a botanical garden speciose = beautiful, often mistakenly used to mean species-rich spermatophyte = colloquial term for seed plant, angiosperms + gymnosperms and several other extinct lineages of seedbearing plants spheroidal = shaped like a globe spicate = with a spike, an unbranched raceme with flowers directly borne on the axis spicule = a small needle spike = an unbranched raceme with flowers borne on the axis, sessile and alternate (Fig. 28b) spikelet = a structure of two sterile bracts (glumes) with a short axis and a number of florets (each consisting of lemma, palea and flower) in Poaceae and Cyperaceae spine = a sharp-tipped, hardened structure derived from a vascular part of a plant, leaf, stipule, root or shoot (Fig. 17b) spinose = a dentate margin in which the lobes have sharp tips (Figs. 21zc, 26i) spinulose = with small spines spirally arranged = with the parts alternating and arranged in a spiral along a central axis spongiose = spongy sporangiophore = a modified part of a fertile leaf or a peltate organ (in Equisetaceae) that bears the sporangia sporangium = a receptacle in which spores are formed in lycopods and ferns (Fig. 13) spore = a cell capable of developing into a gametophyte, found in lycopods, ferns, mosses and algae sporocarp = a hardened case in which the sporangia are placed, found in heterosporous ferns (Marsileaceae and Salviniaceae) sporophyte = the plant generation that produces spores spur = a short tapering projection often associated with the flower and typically containing nectar (Fig. 36) squamate = scaly, with small scales or bracts

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squamose = covered with scales squamule = a small scale squarrose = rough, with the tips of the hairs or scales projecting outwards or a shrub with the branches at right angles to the main stem stalk = a structure that attaches one organ to another by a narrow terete part stamen = often considered to be the male organ of a flower (although it is the pollen that actually produces the sperm cells), consisting of a filament and an anther, which are connected to each other by the connective; anthers contain the pollen (Fig. 29) staminate = male, bearing male organs or male flowers only staminode = a sterile or aborted stamen standard = the large upper petal of a flower in Fabaceae or one of the three upward pointing tepals in Iris (Iridaceae) stele = part of the plant axis that is made up of the vascular system and associated tissue stellate = star-shaped, usually used for hairs that have numerous arms radiating from a central point stem = the main axis of a plant, usually bearing roots and leaves sterile = not fertile, lacking flowers, fruits or sporangia stigma = the pollen receptor that typically sits on top of the gynoecium (Fig. 29) stinging hair = a tubular hair filled with liquid that upon breaking injects the liquid into the skin causing irritation; these are common in, for instance, Loasaceae and Urticaceae stipe = a petiole of a fern, a single trunk of a clump-forming palm, a stalk inside a flower that supports the carpels, a stalk in the pollinarium in orchids or a short, narrow extension at the base of a nutlet in Cyperaceae; the term is used in a variety

spur

Figure 36.

of contexts, which makes it important to know to which family it is being applied stipel = stipule-like outgrowth occurring at the base of a leaflet in compound leaves stipitate = supported by a specialised stalk stipular = related to a stipule stipulate = with stipules stipule = leaf-like, spine-like, scale-like or gland-like appendages of the leaf, usually in pairs at the base of the petiole; in some ferns (Marattiaceae, Osmundaceae) outgrowths of the rhizome on either side of the petiole are referred to as stipules, although these are not associated with the leaf stolon = a vegetative shoot spreading along the ground and rooting at the node, giving rise to new plantlets stoloniferous = bearing stolons stomata = pores in the leaf epidermis used for transpiration stone = a hard seed of a drupe straggling = growing untidily stramineous = straw-like or straw-coloured striate = with parallel grooves strigose = covered with sharp stiff hairs lying flat on the surface strobilate = cone-like, of an inflorescence or fruit that resembles a gymnosperm cone by being covered with imbricate scales strobilus = cone, an inflorescence or sporebearing axis covered in overlapping scales or reduced leaves stunted = of less than normal stature style = part of the gynoecium between the ovary and stigma, often slender and sometimes branching, sometimes absent (Fig. 29) stylopodium = a structure above the ovary from which the styles emerge suberose = corky subfamily = a rank below the level of family subgenus = a rank below the level of genus subopposite = nearly opposite, but not precisely so subsessile = nearly sessile, but with a short petiole or pedicel subshrub = a small shrub, especially one that is woody only basally subspecies = a rank below the level of species, generally geographically disjunct from other subspecies of the same species substrate = material in or on which a plant grows or to which it is attached subtending = standing below and surrounding subulate = awl-shaped, like a needle, tapering to a fine point (Fig. 21b) succulent = juicy, pulpy, fleshy, often used for plants with thick, fleshy, swollen parts sucker = a shoot arising below ground from the roots of a plant, sometimes some

GLOSSARY distance from the main stem suffrutescent = becoming like a subshrub; a herb that becomes somewhat woody at base sulcate = furrowed, grooved superior ovary = an ovary situated above the perianth insertion (Fig. 33a) suture = a juction or seam of union, used for the line along which a carpel opens for instance syconium = a compound hollow fruit with flowers and seeds inside, like a fig (Ficus, Moraceae) (Fig. 35g) symbiont = an individual of one organism living together with a different organism, usually to their mutual advantage symbiosis = a relationship between dissimilar organisms, living together to their mutual advantage symmetrical = able to be divided into two matching halves sympetalous = with united petals synandrium = an androecium with the anthers cohering synangium = compound structure with several locules that produce spores in Marattiaceae, sometimes used for compound fruits in seed plants synapomorphy = a cladistic term used for a shared derived character state that is shared by a clade syncarp = a fruit composed of the fusion of fruits from several flowers, as in Morus synoecious = with male and female flowers in the same inflorescence synonym = a scientific name for a taxon that already has a valid name syntepalum = a tubular structure formed by the fusion of sepals and petals (and split along one side like in Musaceae) synsepalum = a structure formed by two or more fused sepals systematics = the science of classification of organisms, based on evolutionary relationships or the study of variation in, and evolution of, taxa T taproot = the primary root that grows straight down in some species, like a carrot taxon = a term denoting a named group of organisms at any rank taxonomy = the science of classification and ordering of organisms into groups teeth = small, sharp protuberances tegumen = the inner coat of the seed tendril = a coiling structure derived from a branch, leaf or inflorescence, which the plant uses to climb (Figs. 18, 21w) tenuous = thin, narrow tepal = a division of the perianth, used when

there is poor differentiation into petals and sepals or when it is unclear which is which terete = an elongate solid structure that is circular in cross-section, sometimes confused with cylindrical (hollow) terminal = ending an apex or a main axis ternate = arranged in threes, usually in a whorl of three, sometimes used for trifoliate leaves (Fig. 22c) terrestrial = on or in the ground tesselate = of a surface, with markings in squares or rectangles, like venation of Aponogetonaceae or petals of some species of Fritillaria (Liliaceae) testa = the outer coat of the seed tetrad = a cluster of four pollen grains released from the anther as a single unit tetradynamous = a flower with four long stamens and two short ones tetragonal = four-angled in cross-section tetramerous = with parts in fours thallus = vegetative structure not clearly divided into stem and leaf, as in Lemna (Araceae) theca = the locule of an anther therophyte = a plant surviving adverse seasons in the form of seeds thorn = a spine derived from a reduced branch (Fig. 17a) throat = in tubular flowers the part where the corolla tube widens thyrse = a mixed inflorescence type with the main axis racemose and the lateral branches cymose (Fig. 28k) timber = wood used in construction and carpentry tissue = material formed by cells with a similar character and origin tomentose = densely covered in short soft somewhat matted hairs tomentum = a felt-like covering of soft hairs torose = cylindrical with spaced contractions, forming a string of beads tortuous = twisted in various directions trailing = spreading over the surface of the ground but not rooting translucent = letting light through but not transparent transparent = being able to be seen through transverse = at right angles to another organ tree = woody plant with a clear main trunk, usually with secondary thickening and being at least two or three metres tall, although this definition is fluid and merges with the definition of a large shrub triad = a structure occurring in groups of three triandrous = with three stamens tribe = a taxonomic category between subfamily and genus

trichome = an outgrowth of the epidermis, which can have various forms, but has no vascular tissue; a hair, bristle, prickle or scale trichotomous = forked into threes tridentate = with three teeth trifid = split into threes trifoliate = with three leaves, often used erroneously instead of trifoliolate (Fig. 22c) trifoliolate = ternate, a compound leaf divided into three leaflets (Fig. 22f) trilocular = of a gynoecium with three chambers trimerous = with all parts in threes trimonoecious = bearing male, female and bisexual flowers trioecious = with male, female and bisexual flowers on different plants triploid = a plant with cells that have three complete sets of chromosomes tristichous = in three planes, where leaves are arranged in three rows around the stem tristylous = with flowers on different plants of the same species having three lengths of styles trullate = shaped like a trowel, with the widest point below the middle, the base rounded and the apex sharply pointed truncate = with a base or apex that abruptly ends in a straight line as if cut off (Figs. 23e, 25f) trunk = the main axis of a tree or tree-like plant (like a palm or tree fern), from its roots to the crown tuber = an underground swollen root or stem serving as a storage organ (Fig. 16b) tuberculate = covered with wart-like or globe-like outgrowths tubular = cylindrical and hollow tufted = growing in tight clusters tumble-weed = a plant that when in fruit breaks off at the base and is blown around by the wind distributing its seeds, most common in xeric or coastal habitats tundra = flat treeless area in the Arctic tussock = compact clump of grass-like plants twig = a small, usually thin, terminal stem on a woody plant twiner = a climbing plant thats supports itself by having branches that coil around other plants or structures U umbel = an inflorescence with all branches arising (more or less) from the same point on a common peduncle; compound umbels are an umbel on which each branch bears another umbel (Fig. 28e, f)

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GLOSSARY umbilicate = navel-like, with a small central hollow or depression umbo = a small blunt protuberance unarmed = without spines, prickles or other sharp structures to deter herbivory uncinate = hooked at the tip understorey = a layer of vegetation below the canopy of a forest unguiculate = clawed, narrowed into a stalklike base uniflorous = with a single flower unijugate = with a single pair of leaflets making up a compound leaf unilateral = secund, one-sided unilocular = with a single cavity unisexual = having only male or female parts urceolate = urn-shaped, a swollen base and contracted apex, with the rim slightly expanded urticating hairs = stinging hairs utricular = bladder-shaped V vaginate = sheathed valvate = meeting at the margins, not overlapping variable = not constant in appearance variegated = irregularly coloured or striped with two or more colours (Fig. 37) variety = taxonomic rank below the level of subspecies and above the level of form, not usually geographically disjunct from other varieties of the same species vascular = possessing vessels, referring to the xylem or phloem of plants vascular bundle = specialised tissue that transports water, nutrients and carbohydrates within a plant

vascular plant = plant that possesses vessels for the transport of water and nutrients, i.e. lycopods, ferns and seed plants vegetative = associated with roots, stems or leaves, not sexual vein = strand of vascular tissue in a flat organ like a leaf, bract, petal or sepal veinlet = small vein velutinous = velvety hairy venation = the arrangement of veins in a leaf ventral = lower surface vernacular name = name of a plant in any language, not the scientific name vernation = folding of leaves in bud verrucose = warty verticillate = in a whorl, with several structures arising at the same node (Fig. 20c) vesicle = a bladder or small cavity vessels = water-conducting cells vestigial = remnant, of an organ that is reduced in size and has lost its original function vicariant = with a disjunct distribution, usually occupying the same niche vigorous = strong villous = hairy with long, soft, weak hairs vine = climbing herbaceous or woody plant viscous = sticky viviparous = live-bearing, used for when seeds germinate on the parent plant, or where plantlets are formed from vegetative parts of a plant volute = rolled up W wart-like = shaped like a small irregular dome

wax = a fatty substance sometimes covering leaves, fruits or stems weed = plant of disturbed habitats and perceived by the observer to be in the wrong place, like a dandelion, or a rose in a cabbage patch whorl = a set of similar organs arranged in a plane around a node wild = spontaneous and not introduced or cultivated wilting = becoming limp due to too little water in the cells wing = a flattened extension of any organ, used in particular for the lateral petals in a Fabaceae flower winter bud = hibernating shoot that is protected by scales woody = made of wood or wood-like tissue, often due to secondary growth X xeric = of dry areas xerophyte = plant adapted to growing and reproducing in areas with little water availability xylem = the woody element of a vascular bundle with the function of water transport Z zigzag = with short sections arranged in alternating directions from side to side zygomorphic = with bilateral symmetry, each side the mirror image of the other (Fig. 31b)

awn palea

palea

stigma

lemma

lemma

anther

ovary filament peduncle

Figure 37. Variegated

670

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Figure 38. Spikelet, the Poaceae inflorescence (Arrhenatherum elatius) [139]

ACKNOWLEDGEMENTS We thank the curators of the many botanical gardens, plant nurseries, and other organisations for maintaining collections of plants that we have studied. These include: Royal Botanic Gardens, Kew; Chelsea Physic Garden; Hampton Court Palace Garden; Trebah Garden; Caerhays Estate; Burncoose Nurseries; Royal Horticultural Society Garden, Wisley; Beth Chatto Gardens; East Bergholt Place Arboretum; Royal Botanic Garden Edinburgh; Glasgow Botanic Gardens; Bodnant Gardens; Crûg Farm Plants; National Botanic Gardens, Glasnevin; Kilmacurragh Botanic Gardens; Hortus botanicus, University of Leiden; Hortus Bulborum, Limmen; Kwekerij de Border, Delden; Arboretum Poortbulten, De Lutte; Twickel Estate gardens, Hof van Twente; Natura Docet, Denekamp; Finnish Museum of Natural History and Botanical Gardens, Helsinki; University of Turku Botanical Gardens, Ruissalo; Bergius Botanical Garden, Stockholm; Gothenburg Botanical Garden; Copenhagen Botanical Garden; Botanic Garden Meise; Jardin des Plantes, Paris; Gibraltar Botanical Garden; Lisbon Botanical Garden; Archeological Museum of Thessaloniki; Botanical Garden, Berlin-Dahlem; Neues Museum, Berlin; Bucharest Botanical Garden; Beijing Natural History Museum; Chan Shan Botanical Garden, Shanghai; Kunming Botanical Garden; Hong Kong Zoological and Botanical Gardens; Singapore Botanic Garden; Gardens by the Bay, Singapore; Morrison Smith Botanical Garden, Tahiti; Bogor Botanical Garden; National Museums of Kenya Botanical Garden, Nairobi; Conservatoire botanique national de Mascarin, La Réunion; Kings Park and Botanic Garden, Perth; Australian National Botanic Gardens, Canberra; Royal Botanic Gardens, Sydney; Royal Botanic Gardens, Victoria; Adelaide Botanic Garden; Australian Arid Lands Botanic Garden, Port Augusta; University of British Columbia Botanical Garden, Vancouver; Brooklyn Botanic Garden; New York Botanic Garden; Matthaei Botanical

Garden, Ann Arbor; Nichols Arboretum, Ann Arbor; Ann Arbor Natural History Museum; Arrowhead Alpines, Lansing; University of Wisconsin Botanical Garden, Madison; Sarah Duke Botanical Garden, Durham; Marie Selby Botanical Garden, Sarasota; Fairchild Tropical Botanical Garden; Montgomery Botanical Center, Miami; San Antonio Botanical Garden; Rancho Santa Anna Botanical Garden; Santa Barbara Botanical Garden; Santa Barbara Orchid Estate; University of California Botanical Garden, Berkeley; California Academy of Sciences, San Francisco; San Francisco Botanical Garden; The Living Desert, Palm Desert; CICY Botanical Garden, Merida; Parc Zoologique Guadeloupe; Roseau Botanical Garden, Dominica; Trésor Voluntary Reserve, French Guiana; Ecuagenera, Gualaceo; Institute of Botany, São Paulo; Recife Botanical Garden; Rio de Janeiro Botanical Garden; Native Garden and Zoo, Santiago de Chile; Institute of Jamaica, Kingston; Botanical Garden of Bogotá “José Celestino Mutis” and Botanical Garden of Cartagena “Guillermo Piñeres”. We also thank Sandra Knapp for her contribution on Solanaceae, Maria Vorontsova for discussing grasses, and many colleagues and friends, including Jim Ackerman, Maria do Carmo Amaral, William and Christianne Anderson, Pieter Baas, Michelle van der Bank, Matt and Russell Barrett, Clemens Bayer, Cássio van den Berg, Sarah Bollendorf, Kåre and Birgitta Bremer, Jeremy Bruhl, Dick Brummitt, James Byng, Ken Cameron, Keron Campbell, Glenda Gabriela Cárdenas Ramírez, Sherwin Carlquist, Laure Civeyrel, Tracy Commock, John Conran, Helena Cotrim, Salvatore Cozzolino, Benito Crespo, Brian Dennis, Kingsley Dixon, Wayne England, Friedrich Ehrendorfer, Hajo Esser, Leonardo Pessoa Felix, Manfred Fischer, Félix Forest, Mike Frohlich, Cheng-xin Fu, Favio González, Peter Goldblatt, Luis Diego Gómez, A. R. A. Görts-van Rijn, Renée Grayer,

Tony Hall, Sun Hang, Mikael Hedrén, Rebecca Hilgenhof, Eric Holterman, Steve Hopper, Chad Husby, Peris Kamau, Machiel Kiel, Joo-Hwan Kim, Harold Koopowitz, Kathy Kron, Klaus Kubitzki, Samuli Lehtonen, Ilia Leitch, Peter Linder, Lola Lledó, Jana Leong-Škorničková, Viviana Londoño Lemos, Gerard Leussink, Alicia Lourteig, David Mabberley, Santiago Madriñan Restrepo, Carlos Magdalena, John Manning, Peter McCarthy, Rogers McVaugh, Claire Micheneau, Julio Morales Can, Jérôme Munzinger, David Negrotto, Dick Olmstead, Mikko Paajanen, Thierry Pailler, Jeff Palmer, Ovidiu Paun, Terry Pennington, Toby Pennington, Yohan Pillon, João Pinto, Jefferson Prado, Ghillean Prance, George Proctor, André van Proosdij, Hassan Rankou, James Reveal, Martyn Rix, Paula Rudall, Kalle Ruokolainen, Gerardo Salazar, Rose Samuel, Vincent Savolainen, Harald Schneider, Pedro Schwartsburd, Alexander Sennikov, Pam and Doug Soltis, Lassi Suominen, Jorge Carlos Trejo-Torres, Hanna Tuomisto, Mats Thulin, Frédéric Tronchet, Gerda van Uffelen, Henry Väre, Herb Wagner, Ben Erik van Wyk, Nelson Zamora, Wendy Zomlefer and Scott Zona. This book would not have happened without all the discussions, field assistance and support over many years. Many colleagues and friends provided photographs, which we acknowledge separately. Last, but not least , we thank the teams at Kew Publishing (Gina Fullerlove, Andrew Illes Georgie Hills, Lydia White, Ruth Linklater, Sharon Whitehead, Nicola Thompson, Chris Beard) and the University of Chicago Press (Christie Henry, Nicholas Lilly, Miranda Martin, Jill Shimabukuro) for agreeing to take on this major project and seeing it through from manuscript submission to publication. Special thanks to Gina for believing that it was worth taking on and to Andrew and Lydia for their patience and hard work. It’s been a long haul, but we got there in the end.

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PHOTOGRAPHY CREDITS The majority of the photographs were taken by the first author, Maarten Christenhusz, but obviously it was not possible to travel everywhere and photograph every plant in person. Therefore, we asked around and found many skilled photographers who wanted to contribute photographs to this project. We are very grateful that they gave us permission to use these, helping us to illustrate all 451 plant families described in this book. We thank them and hope to be able to work with them on future projects. List of contributors: HP = Homer Edward Price

OO = Olga Martha Monties

AP = André van Proosdij

HR = Hassan Rankou

PE = Pablo Endemico

AS = Alexey Shipunov

HS = Hang Sun

AD = Andy de Wet

(Wikimedia Commons)

AK = Augustin Konda ku Mbuta

(Wikimedia Commons)

BH = Barry Hammel BP = Bob Peterson

(Wikimedia Commons)

BX = Bo Xu BY = Bin Yang CC = Cheng-Wei Chen

(Census and classification of plant resources in the Solomon Islands)

HW = Horngyih Wong IP

= Ghillean Prance

IV = Inao Vasquez

(Wikimedia Commons)

JA = John Manning JB = Jeremy Bruhl JC = John Conran

CD = Christopher Davidson

JD = João de Deus Medeiros

CF = Christian Fischer

JG = John Game

(www.floraoftheworld.org) (Wikimedia Commons)

DA = Daniela Cano

(Wikimedia Commons)

(Wikimedia Commons)

JH = Jason Hollinger

(Wikimedia Commons)

(Wikimedia Commons)

PH = Paula Havas-Matilainen PM = Paul Maas PW = Peter Woodard

(Wikimedia Commons)

QY = Qing-jun Yuan RB = Robert von Blittersdorff

(www.africanplants.senckenberg.de)

RK = Ralph Knapp RW = Rainer Wendt

(www.africanplants.senckenberg.de)

SC = Sue Carnahan SK = Sandra Knapp SP = Stefan Porembski

(www.africanplants.senckenberg.de)

JL = Johannes Lundberg

SS = Stan Shebs

DK = Doris McKellar

JM = Jérôme Munzinger

DM = David Merritt

KC = Kevin Chien-Wen Chen

SZ = Scott Zona

DN = D. H. Hansen

KD = Kingsley Dixon

DH = Dan Heims

DS = Dan Skean DV = Dinesh Valke DZ = Russ Kleinman

(Dale A. Zimmerman Herbarium, Western New Mexico University)

FF = Félix Forest FT = Frédéric Tronchet GB = Günther Baumann

(www.africanplants.senckenberg.de)

GP = Gideon Pisanty

(Wikimedia Commons)

HB = Hans B.

(Wikimedia Commons)

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HV = Henry Väre

(www.floraoftheworld.org)

Christenhusz, Fay & Chase

(Swedish Museum of Natural History)

KH = Kwan Han

(www.natureloveyou.sg)

LW = Lubbert Th. Westra

(Wikimedia Commons)

TD = Tony Dold TF = Thassilo Franke

(Wikimedia Commons)

TM = Tony Miller TW = Tim Waters

MF = Mateo Fernández-Lucero and

UM = Ulf Mehlig

MN = Matti Niisalo

VS = Victoria Sosa

MT = Marcia Stefani

VZ = Vojtěch Zavadil

Santiago Madriñán-Restrepo

(Wikimedia Commons)

(Wikimedia Commons)

(Wikimedia Commons)

MV = Maria Vorontsova

WA = Wee Foong Ang

MW = Maximilian Weigend

WJ = Walter Judd

NH = Nick Helme

YN = Yang Niu

OM = Olivier Maurin

ZZ = Zhuo Zhou

FURTHER READING 1. LYCOPODIACEAE CLUBMOSS FAMILY Bruce JG. 1976. Gametophytes and subgeneric concepts in Lycopodium. American Journal of Botany 63: 919–924. Nessel H. 1929. Die Bärlappgewächse. Gustav Fischer, Jena. Øllgaard B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard B. 2012. New combinations in Neotropical Lycopodiaceae. Phytotaxa 57: 10–22. Wilce JH 1972. Lycopod spores I. General spore patterns and the generic segregates of Lycopodium. American Fern Journal 62: 65–79. Wikström N, Kenrick P. 1997. Phylogeny of Lycopodiaceae (Lycopsida) and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. International Journal of Plant Sciences 158: 862–871. Wikström N, Kenrick P, Chase MW. 1999. Epiphytism and terrestrialisation in tropical Huperzia (Lycopodiaceae). Plant Systematics and Evolution 218: 221–243. Wikstöm N, Kenrick P. 2001. Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by use of nonparametric rate smoothing. Molecular Phylogenetics and Evolution 19: 177–186. Yatsentyuk SP, Valiejo-Roman KM, Samigullin TH, Wikström N, Troitsky AV. 2001. Evolution of Lycopodiaceae inferred from spacer sequencing of chloroplast rRNA genes. Russian Journal of Genetics 37: 1068–1073. 2. ISOËTACEAE QUILLWORT FAMILY Amstutz E. 1957. Stylites, a new genus of Isoëtaceae. Annals of the Missouri Botanical Garden 44: 121–123. Fuchs HP. 1962. Nomenklatur, taxonomie und Systematik der Gattung Isoëtes Linnaeus in geschichtlicher Betrachtung. Beihefte zur Nova Hedwigia 3: 1–103. Hickey RJ. 1986. Isoëtes megaspore surface morphology: nomenclature, variation and systematic importance. American Fern Journal 76: 1–16. Hoot SB, Taylor WC, Napier NS. 2006. Phylogeny and biogeography of Isoëtes (Isoëtaceae) based on nuclear and chloroplast DNA sequence data. Systematic Botany 31: 449–460. Pfeiffer NE. 1922. Monograph of the Isoëtaceae. Annals of the Missouri Botanical Garden 9: 79–233. Retallack GJ. 1997. Earliest Triassic origin of Isoëtes and quillwort evolutionary radiation. Journal of Paleontology 71: 500–521. Rydin C, Wikström N. 2002. Phylogeny of Isoëtes (Lycopsida): resolving basal relationships using rbcL sequences. Taxon 51: 83–89. Weber U. 1922. Zur Anatomie und Systematik des Gattung Isoëtes L. Hedwigia 63: 219–262. 3. SELAGINELLACEAE SPIKEMOSS FAMILY Baker JG. 1883. A synopsis of the genus Selaginella, pt. 1. Journal of Botany (London) 21: 1–5. Banks JA, et al. (103 authors). 2011. The Selaginella genome identifies genetic changes associated with the evolution of vascular plants. Science 332: 960–963. Bienfait A, Waterkeyn L. 1974. Contribution à l’étude

systématique des Selaginella: spécificité des formations callosiques foliaires observées en fluorescence. Bulletin du Jardin Botanique National de Belgique 44: 295–302. Hebant C, Lee DW. 1984. Ultrastructural basis and developmental control of blue iridescence in Selaginella leaves. American Journal of Botany 71: 216–219. Korall P, Kenrick P, Therrien JP. 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. International Journal of Plant Sciences 160: 585–594. Korall P, Kenrick P. 2004. The phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: extreme substitution rates and rate heterogeneity. Molecular Phylogenetics and Evolution 31: 852–864. Minaki M. 1984. Microspore morphology and taxonomy of Selaginella (Selaginellaceae). Pollen et Spores 26: 421–480. Spring A, 1850. Monographie de la famille Lycopodiacées: 2. Selaginella. Mémoires de l’Académie Royale des Sciences, Lettres et Beaux Arts de Belgique 24: 52–264. Wang J-K, Noel JP. 2013. Chemodiversity in Selaginella: a reference system for parallel and convergent metabolic evolution in terrestrial plants. Frontiers in Plant Science 4: 119. 4. EQUISETACEAE HORSETAIL FAMILY Bennert W, Lubienski M, Körner S, Steinberg M. 2005. Triploidy in Equisetum subgenus Hippochaete (Equisetaceae, Pteridophyta). Annals of Botany 95: 807–815. Des Marais DL, Smith AR, Britton DM, Pryer KM. 2003. Phylogenetic relationships and evolution of extant horsetails, Equisetum, based on chloroplast DNA sequence data (rbcL and trnL-F). International Journal of Plant Science 164: 737–751. Guillon JM. 2004. Phylogeny of horsetails (Equisetum) based on the chloroplast rps4 gene and adjacent noncoding sequences. Systematic Botany 29: 251–259. Hauke RL. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Beihefte Nova Hedwigia 8: 1–123. Hauke RL. 1977. Experimental studies on growth and sexual determination in Equisetum gametophytes. American Fern Journal 67: 18–31. Hauke RL. 1978. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedwigia 30: 385–455. Soltis DE. 1986. Genetic evidence for diploidy in Equisetum. American Journal of Botany 73: 908–913. Stanich NA, Rothwell GW, Stockey RA. 2009. Phylogenetic diversification of Equisetum (Equisetales) as inferred from Lower Cretaceous species of British Columbia, Canada. American Journal of Botany 96: 1289–1299. 5. OPHIOGLOSSACEAE ADDER’S-TONGUE FAMILY Clausen RT. 1938. A monograph of the Ophioglossaceae. Memoirs of the Torrey Botanical Club 19: 1–177. Dauphin B, Vieu J, Grant JR. 2014. Molecular phylogenetics supports widespread cryptic species in

moonworts (Botrychium s.s., Ophioglossaceae). American Journal of Botany 101: 128–140. Hauk WD, Parks CR, Chase MW. 2003. Phylogenetic studies of Ophioglossaceae: evidence from rbcL and trnL-F plastid DNA sequences and morphology. Molecular Phylogenetics and Evolution 28: 131–151. Hauk WD, Kennedy L, Hawke HM. 2012. A phylogenetic investigation of Botrychium s. s. (Ophioglossaceae): evidence from three plastid DNA sequence datasets. Systematic Botany 37: 320–330. Kato M. 1987. A phylogenetic classification of the Ophioglossaceae. Gardens’ Bulletin (Singapore) 40: 1–14. Winther JL, Friedman WE. 2007. Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae). American Journal of Botany 94: 1248–1255. 6. PSILOTACEAE WHISK-FERN FAMILY Bierhorst DW. 1977. The systematic position of Tmesipteris and Psilotum. Brittonia 29: 3–13. Hidalgo O, Pellicer J, Christenhusz MJM, Schneider H, Leitch IJ. 2017. Genomic gigantism in the whisk-fern family (Psilotaceae): Tmesipteris obliqua challenges the record holder Paris japonica. Botanical Journal of the Linnean Society 183: 509–514. Jonker FP. 1973. The taxonomic position of the Psilotales in the light of our knowledge of Devonian plant life. Palaeobotanist 20: 33–38. Renzaglia KS, Johnson TH, Gates HD, Whittier DP. 2001. Architecture of the sperm cell of Psilotum. American Journal of Botany 88: 1151–1163. Schneider EL, Carlquist S. 2000. SEM studies on vessels in ferns. 17. Psilotaceae. American Journal of Botany 87: 176–181. Sykes MG. 1908. The anatomy and morphology of Tmesipteris. Annals of Botany o.s. 22: 63–89. Wagner WH. 1977. Systematic implications of the Psilotaceae. Brittonia 29: 54–63. Winther JL, Friedman WE. 2009. Phylogenetic affinity of arbuscular mycorrhizal symbionts in Psilotum nudum. Journal of Plant Research 122: 485–496. 7. MARATTIACEAE KING-FERN FAMILY Brebner G. 1902. On the anatomy of Danaea and other Marattiaceae. Annals of Botany (Oxford) 16: 517–552. Christenhusz MJM. 2007. Evolutionary history and taxonomy of Neotropical marattioid ferns: studies of an ancient lineage of plants. Annales Universitatis Turkuensis ser. AII 216: 1–134. Christenhusz MJM, Toivonen TK. 2008. Giants invading the tropics: the oriental vessel fern, Angiopteris evecta (Marattiaceae). Biological Invasions 10: 1215–1228. Christenhusz MJM, Tuomisto H, Metzgar JS, Pryer KM. 2008. Evolutionary relationships within the Neotropical, eusporangiate fern genus Danaea (Marattiaceae). Molecular Phylogenetics and Evolution 46: 34–48. Christenhusz MJM. 2010. Danaea (Marattiaceae) revisited: biodiversity, a new classification and ten new species of a Neotropical fern genus. Botanical Journal of the Linnean Society 163: 360–385.

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FURTHER READING Christenhusz MJM. 2010. Revision of the Neotropical fern genus Eupodium (Marattiaceae). Kew Bulletin 65: 1–7. De Vriese WH, Harting P. 1853. Monographie des Maratiacées. Arnz Co., Leiden, Düsseldorf. Hill CR, Camus JM. 1986. Evolutionary cladistics of marattialean ferns. Bulletin of the British Museum (Natural History), Botany 14: 219–300. Holttum RE. 1978. The morphology and taxonomy of Angiopteris (Marattiaceae) with description of a new species. Kew Bulletin 32: 587–594. Lavalle MC. 2003. Taxonomía de las especies neotropicales de Marattia. Darwiniana 41: 61–86. Murdock AG. 2008. Phylogeny of marattioid ferns (Marattiaceae): inferring a root in the absence of a closely related outgroup. American Journal of Botany 95: 626–641. Murdock AG. 2008. A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new genus Ptisana. Taxon 57: 737–755. Rolleri CH. 2003 Caracteres diagnósticos y taxonomía en el género Angiopteris Hoffm. (Marattiaceae Bercht, J.S.Presl) II. Sinopsis de las especies. Revista del Museo de La Plata, Botánica 16: 1–23. West C. 1917. A contribution to the study of the Marattiaceae. Annals of Botany (Oxford) 31: 361–414. 8. OSMUNDACEAE ROYAL-FERN FAMILY Bobrov AE. 1967. The family Osmundaceae (R.Br.) Kaulf., its taxonomy and geography. Botanicheskii Zhurnal 52: 1600–1610. Hewitson W. 1962. Comparative morphology of the Osmundaceae. Annals of the Missouri Botanical Garden 49: 57–93. Magrini S, Scoppola A. 2011. First results from conservation studies of chlorophyllous spores of the royal fern (Osmunda regalis, Osmundaceae). Cryobiology 64: 65–69. Metzgar JS, Skog JE, Zimmer EA, Pryer KM. 2008. The paraphyly of Osmunda is confirmed by phylogenetic analyses of seven plastid loci. Systematic Botany 33: 31–36. Stokey AG, Atkinson LR. 1956. The gametophyte of the Osmundaceae. Phytomorphology 6: 19–40. Tian N, Wang Y-D, Jiang Z-K. 2008. Permineralized rhizomes of the Osmundaceae (Filicales): diversity and tempo-spatial distribution pattern. Palaeoworld 17: 183–200. Yatabe Y, Nishida H, Murakami N. 1999. Phylogeny of Osmundaceae inferred from rbcL nucleotide sequences and comparison to the fossil evidences. Journal of Plant Research 112: 397–404. 9. HYMENOPHYLLACEAE FILMY-FERN FAMILY Copeland EB. 1938. Genera hymenophyllacearum. Philippine Journal of Science 67: 1–110. Ebihara A, Dubuisson J-Y, Iwatsuki K, Hennequin S, Ito M. 2006. A taxonomic revision of Hymenophyllaceae. Blumea 51: 221–280. Ebihara A, Iwatsuki K, Ito M, Hennequin S, Dubuisson J-Y. 2007. A global molecular phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem anatomy. Botanical Journal of the Linnean Society 155: 1–27. Farrar DR. 1967. Gametophytes of four tropical fern genera reproducing independently of their sporophytes in the southern Appalachians. Science 155: 1266–1267. Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson

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J-Y. 2003. Molecular systematics of the fern genus Hymenophyllum s.l. (Hymenophyllaceae) based on chloroplastic coding and noncoding regions. Molecular Phylogenetics and Evolution 27: 283–301 Morton CV. 1968. The genera, subgenera and sections of the Hymenophyllaceae. Contributions from the United States National Herbarium 38: 153–214. Pryer KM, Smith AR, Hunt JS, Dubuisson J-Y. 2001. rbcL data reveal two monophyletic groups of filmy ferns (Filicopsida: Hymenophyllaceae). American Journal of Botany 88: 1118–1130. Stokey AG. 1948. Reproductive structures of the gametophytes of Hymenophyllum and Trichomanes. Botanical Gazette 109: 363–380. 10. GLEICHENIACEAE FORKING-FERN FAMILY Bierhorst DW. 1969. On Stromatopteris and its illdefined organs. American Journal of Botany 56: 160–174. Gonzales R. J, Kessler M. 2011. A synopsis of the Neotropical species of Sticherus (Gleicheniaceae), with descriptions of nine new species. Phytotaxa 31: 1–54. Hagemann W, Schulz U. 1978. Wedelanlegung und Rhizomverzweigung bei einigen Gleicheniaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 99: 380–399. Holttum RE. 1957. Morphology, growth habit and classification in the family Gleicheniaceae. Phytomorphology 7: 168–184. Nakai T. 1950. A new classification of Gleicheniales. Bulletin of the National Science Museum (Tokyo) 29: 1–71. Ohl C, Bussmann R. 2004. Recolonisation of natural landslides in tropical mountain forests of southern Ecuador. Feddes Repertorium 115: 248–264. Perrie LR, Bayly MJ, Lehnebach CA, Brownsey PJ. 2007. Molecular phylogenetic and molecular dating of the New Zealand Gleicheniaceae. Brittonia 59: 129–141. Shaw SW, Ranker TA. 2011. New and improved leaf terminology for Gleicheniaceae. American Fern Journal 101: 117–124. Stokey AG. 1950. The gametophyte of the Gleicheniaceae. Bulletin of the Torrey Botanical Club 77: 323–339. Wallace JW, Pozner, RS, Gómez LD. 1983. A phytochemical approach to the Gleicheniaceae. American Journal of Botany 70: 207–211. 11. DIPTERIDACEAE DOUBLE-FERN FAMILY Kato M, Yatabe Y, Sahashi N, Murakami N. 2001. Taxonomic studies of Cheiropleuria (Dipteridaceae). Blumea 46: 513–525. Posthumus O. 1928. Einige Eigentümlichkeiten der Blattform bei Dipteris und bei anderen noch lebenden oder fossilen Pf lanzen. Recueil des Travaux Botaniques Néerlandais 25: 244–249. Stockey RA, Rothwell GW, Little SA. 2006. Relationships among fossil and living Dipteridaceae: anatomically preserved Hausmannia from the Lower Cretaceous of Vancouver Island. International Journal of Plant Sciences 167: 649–663. Stokey AG. 1945. The gametophyte of Dipteris conjugata. Botanical Gazette 106: 402–411. Stokey AG, Atkinson LR. 1954. The gametophyte of Cheiropleuria bicuspis (Bl.) Presl. Phytomorphology 4: 192–201.

12. MATONIACEAE UMBRELLA-FERN FAMILY Kato M. 1993. A taxonomic study of the genus Matonia (Matoniaceae). Blumea 38: 167–172. Kato M, Iwatsuki I. 1985. Juvenile leaves and leaf ramification in Phanerosorus major (Matoniaceae). Acta Phytotaxonomica et Geobotanica 36: 139–147. Klavins SD, Taylor TN, Taylor EL. 2004. Matoniaceous ferns (Gleicheniales) from the Triassic of Antarctica. Journal of Paleontology 78: 211–217. Stokey AG, Atkinson LR. 1952. The gametophytes and young sporophyte of Matonia pectinata R. Br. Phytomorphology 2: 138–150. Tansley MA, Lulham RBJ. 1905. A study of the vascular system of Matonia pectinata. Annals of Botany (Oxford) 19: 475–519. 13. SCHIZAEACEAE FAN-FERN FAMILY Bierhorst DW. 1966. The fleshy cylindrical subterranean gametophyte of Schizaea melanesica. American Journal of Botany 53: 123–133. Bierhorst DW. 1968. Observations on Schizaea and Actinostachys spp., including A. oligostachys sp. nov. American Journal of Botany 55: 87–108. Bierhorst DW. 1971. Morphology and anatomy of new species of Schizaea and Actinostachys. American Journal of Botany 58: 634–648. Brown K. (ed.) 2006. [articles by various authors dedicated to invasive Lygodium in Florida]. Wildland Weeds 9 (2), Spring 2006. Corrêa Pinto S, Guimarães Leitão G, Ribeiro de Oliveira D, Ribeiro Bizzo H, Fernandes Ramos D, Silveira Coelho T, Silva PEA, Lourenço MCS, Guimarães Leitão S. 2009. Chemical composition and antimycobacterial activity of the essential oil from Anemia tomentosa var. anthriscifolia. Natural Product Communications 4: 1675–1678. Davidson C, Prusinkiewicz P, von Aderkas P. 2008. Description of a novel organ in the gametophyte of the fern Schizaea pusilla and its contribution to overall plant architecture. Botany 86: 1217–1223. Duek JJ. 1978. A taxonomic revision of Lygodium (Filicinae) in America. Feddes Repertorium 91: 69–87. Duek JJ. 1980. A taxonomical monograph of Anemia subgenus Anemiorrhiza (Filicinae). Feddes Repertorium 91: 69–87. Lott MS, Volin JC, Pemberton RW, Austin DF. 2003. The reproductive biology of the invasive ferns Lygodium microphyllum and L. japonicum (Schizaeaceae): implications for invasive potential. American Journal of Botany 90: 1144–1152. Madeira PT, Pemberton RW, Center TD. 2008. A molecular phylogeny of the genus Lygodium (Schizaeaceae) with special reference to the biological control and host range testing of Lygodium microphyllum. Biological Control 45: 308–318. Mickel JT. 1967. A monographic study of the fern genus Anemia subgenus Coptophyllum. Iowa State Journal of Science 36: 349–482. Mickel JT. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33: 413–429. Richter A. 1914. Phylogenetisch-taxonomische und physiologisch-anatomische Studien über Schizaea. Mathematische und Naturwissenschaftlische Berichte aus Ungarn 30: 213–297. Skog, JE, Zimmer EA, Mickel JT. 2002. Additional suppor t for t wo subgenera of Anemia (Schizaeaceae) from data for the chloroplast

FURTHER READING intergenic spacer region trnL-F and morphology. American Fern Journal 92: 119–130. Wikström N, Kenrick P, Vogel JC. 2002. Schizaeaceae: a phylogenetic approach. Review of Palaeobotany and Palynology 119: 35–50. 14. MARSILEACEAE PILLWORT FAMILY Bhardwaja TN. 2008. Marsileaceae: an amphibious heterosporous group of ferns. The persisting enigma. Asian Journal of Experimental Science 22: 189–191. Johnson DM. 1985. New records of longevity for Marsilea sporocarps. American Fern Journal 75: 30–31. Mahlberg PG, Baldwin M. 1975. Experimental studies on megaspore viability, parthenogenesis and sporophyte formation in Marsilea, Pilularia and Regnellidium. Botanical Gazette 136: 269–273. Nagalingum NS, Schneider H, Pryer KM. 2007. Molecular phylogenetic relationships and morphological evolution in the heterosporous fern genus Marsilea. Systematic Botany 32:16–25. Nagalingum NS, Novak MD, Pryer KM. 2008. Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia. Botanical Journal of the Linnean Society 157: 673–685. Pryer KM. 1999. Phylogeny of marsileaceous ferns and relationships of the fossil Hydropteris pinnata reconsidered. International Journal of Plant Sciences 160: 931–954. Schneider H, Pryer KM. 2002. Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. International Journal of Plant Sciences 163: 485–505. Schneider H, Pryer KM. 2002. Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. International Journal of Plant Sciences 163: 485–505. Yamada T, Kato M. 2002. Regnellites nagashimae gen. et sp. nov., the oldest macrofossil of Marsileaceae, from the Upper Jurassic to Lower Cretaceous of western Japan. International Journal of Plant Sciences 163: 715–723. 15. SALVINIACEAE WATER-FERN FAMILY Barthlott W, Schimmel T, Wiersch S, Koch K, Brede M, Barczewski M, Walheim S, Weis A, Kaltenmaier A, Leder A, Bohn HF. 2010. The Salvinia paradox: superhydrophobic surfaces with hydrophilic pins for air retention under water. Advanced Materials 22: 2325–2328. Cohen-Shoel N, Barkay Z, Ilzycer D, Gilath I, Tel-Or E. 2002. Biofiltration of toxic elements by Azolla biomass. Water, Air, and Soil Pollution 135: 93–104. Duckett JG. Toth R, Soni SL. 1975. An ultrastructural study of the Azolla-Anabaena azollae relationship. New Phytologist 75: 111–118. Evrard C, Van Hove C. 2004. Taxonomy of the American Azolla species (Azollaceae): a critical review. Systematics and Geography of Plants 74: 301–318. Follieri M. 1977. Classification and phylogeny of living and fossil water ferns of the genus “Azolla”. Webbia 31: 97–104. Hall JW. 1974. Cretaceous Salviniaceae. Annals of the Missouri Botanical Garden 61: 354–367. Metzgar JS, Schneider H, Pryer KM. 2007. Phylogeny and divergence time estimates for the fern genus Azolla (Salviniaceae). International Journal of Plant Sciences 168: 1045–1053.

Nagalingum NS, Novak MD, Pryer KM. 2008. Assessing phylogenetic relationships in extant heterosporous ferns (Salviniales), with a focus on Pilularia and Salvinia. Botanical Journal of the Linnean Society 157: 673–685. Olguín EJ, Hernández E, Ramos I. 2002. The effect of both different light conditions and the pH value on the capacity of Salvinia minima Baker for removing cadmium, lead and chromium. Acta Biotechnologica 22: 121–131. Prasanna R, Tripathi U, Dominic TK, Singh AK, Yadav AK, Singh PK. 2003. An improvized technique for measurement of nitrogen fixation by blue green algae and Azolla using moist soil cores from rice fields. Experimental Agriculture 39: 145–150. Van Hove1 C, Lejeune A. 2002. The Azolla–Anabaena symbiosis. Biology, Environment: Proceedings of the Royal Irish Academy 102: 23–26. Yasui K. 1911. On the life-history of Salvinia natans. Annals of Botany (London) 25: 469–483. 16. CYATHEACEAE TREE-FERN FAMILY Becker R. 1984. The identification of Hawaiian tree ferns of the genus Cibotium. American Fern Journal 74: 97–100. Berry EW. 1911. A revision of the fossil ferns from the Potomac group which have been referred to the genera Cladophlebis and Thyrsopteris. Proceedings of the United States National Museum 41: 307–332. Cao J-G, Dai X-L, Wang Q-X. 2011. Archegonial development and oogenesis of the fern Plagiogyria euphlebia and their phylogenetic significance. American Fern Journal 101: 231–240. Chandra S. 1970. Vascular organisation of the rhizome of Cibotium barometz. American Fern Journal 60: 68–72. Christenhusz MJM. 2009. New combinations and an overview of Cyathea subg. Hymenophyllopsis. Phytotaxa 1: 37–42 Conant DS. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. Journal of the Arnold Arboretum 64: 333–382. Domin K. 1930. The species of the genus Cyathea J. E. Sm. Acta Botanica Bohemica 9: 85–174. Eleutério AA, Pérez-Salicrup D. 2006. Management of tree ferns (Cyathea spp.) for handicraft production in Cuetzalan, Mexico. Economic Botany 60: 182–186. Fernandes I. 2000. Taxonomia dos representantes de Dicksoniaceae no Brasil. Pesquisas, Botânica 50: 5–26. Gastony GJ. 1973. A revision of the fern genus Nephelea. Contributions from the Gray Herbarium 203: 81–148. Holttum RE. 1957. The scales of Cyatheaceae. Kew Bulletin 1957: 41–45. Holttum RE, Sen U. 1961. Morphology and classification of the tree ferns. Phytomorphology 11: 406–420. Korall P, Conant DS, Metzgar JS, Schneider H, Pryer KM. 2007. A molecular phylogeny of scaly tree ferns (Cyatheaceae). American Journal of Botany 94: 873–886. Korall P, Pryer KM, Metzgar JS, Schneider H, Conant DS. 2006. Tree ferns: monophyletic groups and their relationships as revealed by four proteincoding plastid loci. Molecular Phylogenetics and Evolution 39: 830–845. Lee H. 1887. The vegetable lamb of Tartary; a curious fable of the cotton plant. To which is added a sketch of the history of cotton and the cotton trade.

Sampson Low, Marston, Searle and Rivington, London Lehnert M. 2011. The Cyatheaceae (Polypodiopsida) of Peru. Brittonia 63: 11–45. Lehnert M. 2012. A synopsis of the species of Cyathea (Cyatheaceae-Polypodiopsida) with pinnate to pinnate-pinnatifid fronds. Phytotaxa 61: 17–36. Lehnert M, Mönnich M, Pleines T, Schmidt-Lebuhn A, Kessler M. 2001. The relictual fern genus Loxsomopsis. American Fern Journal 91: 13–24. Markham KR, Given DR. 1979. The flavonoids of ferns in the isolated genera Loxsoma cunninghamii and Loxsomopsis costaricensis. Biochemical Systematics and Ecology 7: 91–93. Maxon WR. 1912. The American species of Cibotium. Contribution from the United States National Herbarium 16: 54–58. Maxon WR. 1922. The genus Culcita. Journal of the Washington Academy of Science 12: 454–460. Mukherjee AK, Sen T. 1986. Gametophytes of some tree ferns and their impact on phylogenetic relationships. Indian Fern Journal 3: 70–81. Perrie L, Brownsey P. 2007. Molecular evidence for long-distance dispersal in the New Zealand pteridophyte flora. Journal of Biogeography 34: 2028–2038. Qiu Y-J, White RA, Turner MD. 1995. The developmental anatomy of Metaxya rostrata (Filicales: Metaxyaceae). American Journal of Botany 82: 969–981. Ramírez-Barahona S, Luna-Vega I, Tejero-Díez D. 2010. Species richness, endemism, and conservation of American tree ferns (Cyatheales). Biodiversity and Conservation 20: 59–72. Roy SK, Holttum RE. 1965. Cytological and morphological observations on Metaxya rostrata (H. B. K.) Presl. American Fern Journal 55: 158–164. Sen T, Rahaman S. 1999. Anatomy of Thyrsopteris elegans Kunze. Indian Fern Journal 16: 123–129. Sen U. 1968. Anatomy of Culcita macrocarpa. Canadian Journal of Botany 46: 43–46. Smith AR, Tuomisto H, Pryer KM, Hunt JS, Wolf PG. 2001. Metaxya lanosa, a second species in the genus and fern family Metaxyaceae. Systematic Botany 26: 480–486. Stokey AG, Atkinson LR. 1956. The gametophyte of Loxsoma cunninghamii R. Br. and Loxsomopsis costaricensis Christ. Phytomorphology 6: 249–261. Stolze RG. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana, Botany 37: 1–98. Tryon R. 1970. The classification of the Cyatheaceae. Contributions from the Gray Herbarium 200: 1–53. Tryon RM. 1976. A revision of the genus Cyathea. Contributions from the Gray Herbarium 206: 19–98. Tryon RM, Gastony GJ. 1975. The biogeography of endemism in the Cyatheaceae. Fern Gazette 11: 73–79. Virtbauer J, Krenn L, Kählig H, Hüfner A, Donath O, Marian B. 2008. Chemical and pharmacological investigations of Metaxya rostrata. Zeitschrift für Naturforschung 63c: 469–475. White RA, Turner MD. 1988. Calochlaena, a new genus of dicksonioid ferns. American Fern Journal 78: 86–95. Windisch PG. 1977. Synopsis of the genus Sphaeropteris (Cyatheaceae) with a revision of the Neotropical exindusiate species. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 98: 176–198.

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FURTHER READING Wolf PG, Sipes S, White M, Martines M, Pryer KM, Smith AR, Ueda K. 1999. Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: evidence from rbcL nucleotide sequences. Plant Systematics and Evolution 219: 263–270. Zhang XC, Nooteboom HP. 1998. A taxonomic revision of Plagiogyriaceae (Pteridophyta). Blumea 43: 401–469. 17. LONCHITIDACEAE VELVET-FERN FAMILY Lehtonen S, Walhberg N, Christenhusz MJM. 2012. Diversification of lindsaeoid ferns and phylogenetic uncertainty of early polypod relationships. Botanical Journal of the Linnean Society 170: 489–503. Lellinger DB. 1977. The identity of Lonchitis aurita and the generic names Anisosorus and Lonchitis. Taxon 26: 578–580. 18. SACCOLOMATACEAE POUCH-FERN FAMILY Nair GB. 1979. Peltate scales in Saccoloma. Fern Gazette 12: 53–55. Nair GB. 1992. The fern genus Saccoloma Kaulf.: a taxonomic study. Journal of Economic and Taxonomic Botany 16: 637–646. 19. CYSTODIACEAE ROWAN-FERN FAMILY Atkinson LR. 1965. The gametophyte of Cystodium. American Fern Journal 55: 32–35 Croft JR. 1986. The stipe and rachis vasculature of the dicksonioid fern, Cystodium sorbifolium (Cystodiaceae). Kew Bulletin 41: 789–803. Korall P, Conant DS, Schneider H, Ueda K, Nishida H, Pryer KM. 2006. On the phylogenetic position of Cystodium: it’s not a treefern — it’s a polypod! American Fern Journal 96: 45–53. 20. LINDSAEACEAE LACE-FERN FAMILY Kramer KU. 1957. A revision of the genus Lindsaea in the New World with notes on allied genera. Acta Botanica Neerlandica 6: 97–290. Kramer KU. 1968. The lindsaeoid ferns of the Old World II. A revision of Tapeinidium. Blumea 15: 545–556. Lehtonen S, Tuomisto H, Rouhan G, Christenhusz MJM. 2010. Phylogenetics and classification of the pantropical fern family Lindsaeaceae. Botanical Journal of the Linnean Society 163: 305–359. Lehtonen S, Tuomisto H, Rouhan G, Christenhusz MJM. 2013. A revision of the fern genus Osmolindsaea (Lindsaeaceae). Systematic Botany 38: 887–900. Maxon WR. 1913. The genus Odontosoria. Contributions of the United States National Herbarium 17: 157–168. Schneider H, Kenrick P. 2001. An Early Cretaceous root-climbing epiphyte (Lindsaeaceae) and its significance for calibrating the diversification of polypodiaceous ferns. Review of Palaeobotany and Palynology 115: 33–41. 21. DENNSTAEDTIACEAE BRACKEN FAMILY: Brownsey PJ. 1983. Polyploidy and aneuploidy in Hypolepis, and the evolution of the Dennstaedtiales. American Fern Journal 73: 97–108. Camino A, Johns T. 1988. Laki-laki (Dennstaedtia glauca, Polypodiaceae): a green manure used in traditional Andean agriculture. Economic Botany 42: 45–54. Keating RC. 1968. Trends of specialisation in the

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stipe anatomy of Dennstaedtia and related genera. American Fern Journal 58: 126–140. Mickel JT. 1973. The classification and phylogenetic position of the Dennstaedtiaceae. Botanical Journal of the Linnean Society 67 Suppl. 1: 135–144. Perring FH, Gardiner BG (eds). 1976. The biology of bracken. Botanical Journal of the Linnean Society 73: 1–302. Schwartsburd PB. 2012. Three new taxa of Hypolepis (Dennstaedtiaceae) from the Brazilian Atlantic Forest, and a key to the Brazilian taxa. Kew Bulletin 67: 815–825. Taylor JA, Smith RT (eds). 2000 Bracken fern: toxicity, biology and control. Special Publication 4. Aberystwyth: International Bracken Group xiii; 218p. [Comprehensive study made up of 40 papers]. Thomson JA. 2000. Morphological and genomic diversity in the genus Pteridium (Dennstaedtiaceae). Annals of Botany 85: 77–99. Thomson JA. 2004. Towards a taxonomic revision of Pteridium (Dennstaedtiaceae). Telopea 10: 793–803. Tryon RM. 1960. A review of the genus Dennstaedtia in America. Contributions from the Gray Herbarium 191: 91–107. Wolf PG, Soltis PD, Soltis DE. 1994. Phylogenetic relationships of dennstaedtioid ferns: evidence from rbcL sequences. Molecular Phylogenetics and Evolution 3: 383–392. Wolf PG. 1995. Phylogenetic analysis of rbcL and nuclear ribosomal RNA sequences in Dennstaedtiaceae. American Fern Journal 85: 306–327. 22. PTERIDACEAE RIBBON-FERN FAMILY: Crane EH. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic Botany 22: 509–517. Crane EH, Farrar DR, Wendel JF. 1996. Phylogeny of the Vittariaceae: convergent simplification leads to a polyphyletic Vittaria. American Fern Journal 85: 283–305. Eiserhardt WL, Rohwer JG, Russell SJ, Yesilyurt JC, Schneider H. 2011. Evidence for radiations of cheilanthoid ferns in the Greater Cape Floristic Region. Taxon 60: 1269–1283. Gastony GJ, Rollo DR. 1995. Phylogeny and generic circumscriptions of cheilanthoid ferns (Pteridaceae: Cheilanthoideae) inferred from rbcL nucleotide sequences. American Fern Journal 85: 341–360. Kirkpatrick REB. 2007. Investigating the monophyly of Pellaea (Pteridaceae) in the context of a phylogenetic analysis of cheilanthoid ferns. Systematic Botany 32: 504–518. Knobloch IW, Tai W, Adangappuram T. 1975. Chromosome counts in Cheilanthes and Aspidotis with a conspectus of the cytology of the Sinopteridaceae. American Journal of Botany 62: 649–654. Kuhn M. 1881. Übersicht über die Arten der Gattung Adiantum. Jahrbuch des Königlichen Botanischen Gartens und des Botanischen Museums zu Berlin 1: 337–351. Nakazato T, Gastony GJ. 2003. Molecular phylogenetics of Anogramma species and related genera (Pteridaceae: Taenitidoideae). Systematic Botany 28: 490–502. Prado J, Del Nero Rodrigues C, Salatino A, Salatino MLF. 2007. Phylogenetic relationships among Pteridaceae, including Brazilian species, inferred from rbcL sequences. Taxon 56: 355–368. Prado J, Windisch PG. 2000. The genus Pteris L. (Pteridaceae) in Brazil. Boletim do Instituto de

Botânica, São Paulo 13: 103–199. Sánchez-Baracaldo P. 2004. Phylogenetics and biogeography of the Neotropical fern genera Jamesonia and Eriosorus (Pteridaceae). American Journal of Botany 91: 274–284. Sánchez-Baracaldo P. 2004. Phylogenetic relationships of the subfamily Taenitoideae, Pteridaceae. American Fern Journal 94: 126–142. Schneider H, He L, Hennequin S, Zhang X-C. 2013. Towards a natural classification of Pteridaceae: inferring the relationships of enigmatic pteridoid fern species occurring in the Sino-Himalaya and Afro-Madagascar. Phytotaxa 77: 49–60. Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM. 2007. A molecular phylogeny of the fern family Pteridaceae: assessing overall relationships and the affinities of previously unsampled genera. Molecular Phylogenetics and Evolution 44: 1172–1185. Singh M, Singh N, Khareb PB, Rawata AKS. 2008. Antimicrobial activity of some important Adiantum species used traditionally in indigenous systems of medicine. Journal of Ethnopharmacology 115: 327–329. Sundue M. 2009. Silica bodies and their systematic implications in Pteridaceae (Pteridophyta). Botanical Journal of the Linnean Society 161: 422–435. Tryon AF. 1957. A revision of the fern genus Pellaea sect. Pellaea. Annals of the Missouri Botanical Garden 44: 125–192. Tryon AF. 1964. Platyzoma — a Queensland fern with incipient heterospory. American Journal of Botany 51: 939–942. Williams S. 1927. A critical examination of the Vittarieae with a view to their systematic comparison. Transactions of the Royal Society of Edinburgh 55: 173–217. Windham MD, Yatskievych G. 2003. Chromosome studies of cheilanthoid ferns (Pteridaceae: Cheilanthoideae) from the western United States and Mexico. American Journal of Botany 90: 1788–1800. Wolf PG, Rowe CA, Sinclair RB, Hasebe M. 2003. Complete nucleotide sequence of the chloroplast genome from a leptosporangiate fern, Adiantum capillus-veneris L. DNA Research 10: 59–65. Zhang G, Zhang X, Chen Z. 2005. Phylogeny of cryptogrammoid ferns and related taxa based on rbcL sequences. Nordic Journal of Botany 2: 485–493. 23. ASPLENIACEAE SPLEENWORT FAMILY Adjie B, Takamiya M, Ohto M, Ohsawa TA, Watano Y. 2008. Molecular phylogeny of the lady fern genus Athyrium in Japan based on chloroplast rbcL and trnL-trnF sequences. Acta Phytotaxonomica et Geobotanica 59: 79–95. Askerov AM. 1986. Gimenocist — endemničnyi rod flory Kavkaza [Hymenocystis — an endemic genus of the flora of the Caucasus]. Izvestiya Akademii Nauk Azerbajdzhanskoj SSR. Ser. Biologiceskaja 1986(3): 52–55. Bergeron ME, Lapointe L. 2001. Impact of one year crozier removal on long-term frond production in Matteuccia struthiopteris. Canadian Journal of Plant Science 81: 155–163. Bir SS. 1969. Observations on the morphology and anatomy of Diplaziopsis javanica. Plant Science 1: 109–118. Blasdell RF. 1963. A monographic study of the fern genus Cystopteris. Memoirs of the Torrey Botanical Club 21: 1–102.

FURTHER READING Brown D. 1964. A monographic study of the fern genus Woodsia. Beihefte zur Nova Hedwigia 16: 1–154. Ching R-C. 1964. On the genus Diplaziopsis C.Chr. Acta Phytotaxonomica Sinica 9: 31–38. Christenhusz MJM, Zhang X-C, Schneider H. 2011. A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. Christenhusz MJM, Schneider H. 2011. Corrections to Phytotaxa 19: linear sequence of lycophytes and ferns. Phytotaxa 28: 50–52. Copeland EB. 1942. Edible ferns. American Fern Journal 32: 121–126. DeLong JM, Prange RK. 2008. Fiddlehead fronds: nutrient rich delicacy. Chronica Horticulturae 48: 12–15. Gastony GJ, Ungerer MC. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae). American Journal of Botany 84: 840–849. He L-J, Zhang X-C. 2012. Exploring generic delimitation within the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution 65: 757–764. Holttum RE. 1971. Studies in the family Thelypteridaceae III: a new system of genera in the Old World. Blumea 19: 17–52. Holttum RE. 1971. The genus Stenochlaena J. Smith with descriptions of a new species. American Fern Journal 61: 119–123. Holttum RE. 1973. The family Thelypteridaceae in the Old World. Botanical Journal of the Linnean Society 67, Suppl. 1: 173–189. Iwatsuki K. 1962. The trichomes of the thelypteroid ferns. Memoirs of the College of Science, Kyoto Imperial University ser. B 31: 125–197. Kato M. 1975. On the systematic position of Athyrium mesosorum. Acta Phytotaxonomica et Geobotanica 27: 56–60. Kato M. 1979. Taxonomic study of the genus Cornopteris (Athyriaceae). Acta Phytotaxonomica et Geobotanica 30: 101–118. Kato M. 1984. A taxonomic study of the athyrioid fern genus Deparia with main reference to Pacific species. Journal of the Faculty of Science, University of Tokyo, Section III. Botany 13: 375–430. Kato M, Darnaedi D. 1988. Taxonomic and phytogeographic relationships of Diplazium flavoviride, D. pycnocarpon and Diplaziopsis. American Fern Journal 78: 77–85. Li C, Lu S, Sun X, Yang Q. 2011. Phylogenetic positions of the enigmatic Asiatic fern genera Diplaziopsis and Rhachidosorus from analyses of four plastid genes. American Fern Journal 101: 142–155. Liu Y-C, Chiou W-L, Kato M. 2011. Molecular phylogeny and taxonomy of the fern genus Anisocampium (Athyriaceae). Taxon 60: 824–830. Lovis JD. 1973. A biosystematic approach to phylogenetic problems and its application to the Aspleniaceae. Botanical Journal of the Linnean Society 67 Suppl. 1: 211–228. Meyer DE. 1952. Untersuchungen über Bastardierung in der Gattung Asplenium. Bibliotheca Botanica (Stuttgart) 123: 1–34. Miyazawa M, Horiuchi E, Kawata J. 2007. Components of the essential oil from Matteuccia struthiopteris. Journal of Oleo Science 56: 457–461. Muñiz-Díaz de León ME, Pérez-García B, MárquezGuzmán J, Mendoza-Ruiz A. 2008. Developmental gametophyte morphology of seven species of

Thelypteris subg. Cyclosorus (Thelypteridaceae). Micron 39: 1351–1362. Murakami N. 1995. Systematics and evolutionary biology of the fern genus Hymenasplenium (Aspleniaceae). Journal of Plant Research 108: 257–268. Murakami N, Nogami S, Watanabe M, Iwatsuki K. 1999. Phylogeny of Aspleniaceae inferred from rbcL nuleotide sequences. American Fern Journal 89: 232–243. Pearman RW. 1976. A scanning electron microscopic investigation of the spores of the genus Cystopteris. Fern Gazette 11: 221–230. Pinter I, Bakker F, Barrett J, Cox C, Gibby M, Henderson S, Morgan-Richards M, Rumsey F, Russell S, Trewick S, Schneider H, Vogel J. 2002. Phylogenetic and biosystematic relationships in four highly disjunct polyploid complexes in the subgenera Ceterach and Phyllitis in Asplenium (Aspleniaceae). Organisms Diversity, Evolution 2: 299–311. Rothfels CJ, Larsson A, Kuo L-Y, Korall P, Chiou W-L, Pryer KM. 2012. Overcoming deep roots, fast rates, and short internodes to resolve the ancient rapid radiation of eupolypod II ferns. Systematic Biology 16: 490–509. Rothfels CJ, Sundue MA, Kuo L-Y, Larsson A, Kato M, Schuettpelz E, Pryer KM. 2012. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61: 515–533. Sano R. Takamiya M, Ito M, Kurita S, Hasebe M. 2000. Phylogeny of the lady fern group, tribe Physematieae (Dryopteridaceae), based on chloroplast rbcL gene sequences. Molecular Phylogenetics and Evolution 15: 403–413. Sarvela J. 1978. A synopsis of the fern genus Gymnocarpium. Annales Botanici Fennici 15: 101–106. Schneider H, Russel S, Cox C, Bakker F, Henderson S, Rumsey F, Barrett J, Gibby M, Vogel, J. 2004. Chloroplast phylogeny of asplenioid ferns based on rbcL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its implications for biogeography. Systematic Botany 29: 260–274. Smith AR, Cranfill RB. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae). American Fern Journal 92: 131–149. Smith AR. 1971. Systematics of the Neotropical species of Thelypteris section Cyclosorus. University of California Publications in Botany 59: 1–143. Smith AR. 1974. A revised classification of Thelypteris subgenus Amauropelta. American Fern Journal 64: 83–95. Van den Heede CJ, Viane RLL, Chase MW. 2003. Phylogenetic analysis of Asplenium subgenus Ceterach (Pteridophyta: Aspleniaceae) based on plastid and nuclear ribosomal ITS DNA sequences. American Journal of Botany 90: 481–495. Von Aderkas P. 1984. Economic history of ostrich fern, Matteuccia struthiopteris, the edible fiddlehead. Economic Botany 38: 14–23. Wagner WH Jr. 1951. The fern genus Diellia. Its structure, affinities and taxonomy. University of California Publications in Botany 26: 1–212. Wang M-L, Chen Z-D, Zhang X-C, Lu S-G, Zhao G-F. 2003. Phylogeny of the Athyriaceae: evidence from chloroplast trnL-F region sequences. Acta Phytotaxonomica Sinica 41: 416–426. Wei R, Zhang X-C, Qi X-P. 2010. Phylogeny of Diplaziopsis and Homalosorus based on two chloroplast DNA sequences: rbcL and rps4+rps4-trnS IGS. Acta Botanica Yunnanica 17: 46–54.

Wood CC. 1973. Spore variation in the Thelypteridaceae. Botanical Journal of the Linnean Society 67 Suppl. 1: 191–202. 24. POLYPODIACEAE POLYPODY FAMILY Chang YF, Wang RX, Lu SG. 2006. Morphological and anatomical studies of subfam. Polypodioideae (Polypodiaceae). Acta Botanica Yunnanica 28: 587–592. Christenhusz MJM, Jones M, Lehtonen S. 2013. Phylogenetic placement of the enigmatic fern genus Dracoglossum. American Fern Journal 103: 131–138. Fraser-Jenkins CR. 1986. A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bulletin of the British Museum (Natural History) Botany 14: 183–218. Geiger JMO, Ranker TA. 2005. Molecular phylogenetics and historical biogeography of Hawaiian Dryopteris (Dryopteridaceae). Molecular Phylogenetics and Evolution 34: 392–407. Hennequin S, Hovenkamp P, Christenhusz MJM, Schneider H. 2010. Phylogenetics and biogeography of Nephrolepis — a tale of old settlers and young tramps. Botanical Journal of the Linnean Society 164: 113–127. Hennipman E, Roos MC. 1982. A monograph of the fern genus Platycerium (Polypodiaceae). NorthHolland Publishing Co., Amsterdam. Holttum RE. 1986. Studies on the fern genera allied to Tectaria I. A commentary on recent schemes of classification. Fern Gazette 12: 313–319. Hovenkamp PH. 1986. A monograph of the fern genus Pyrrosia (Polypodiaceae). Leiden Botanical Series 9. E. J. Brill, Leiden. Hovenkamp PH, Ho B-C. 2012. A revision of the fern genus Oleandra (Oleandraceae) in Asia. PhytoKeys 11: 1–37. Hovenkamp PH, Miyamoto F. 2005. A conspectus of the native and naturalized species of Nephrolepis (Nephrolepidaceae) in the World. Blumea 50: 279–322. Janssen T, Schneider H. 2005. Exploring the evolution of humus collecting leaves in drynarioid ferns (Polypodiaceae, Polypodiidae) based on phylogenetic evidence. Plant Systematics and Evolution 252: 175–197. Jarrett FM. 1980. Studies on the classifiction of the leptoporangiate ferns: I. The affinities of the Polypodiaceae sensu stricto and the Grammitidaceae. Kew Bulletin 34: 825–833. Kato M, Mitsuta S. 1980. Stelar organisation in davallioid ferns. Phytomorphology 29: 362–369. Kreier HP, Schneider H. 2006. Phylogeny and biogeography of the staghorn fern genus Platycerium (Polypodiaceae, Polypodiidae). American Journal of Botany 93: 217–225. Li F-W, Tan BC, Buchbender V, Moran RC, Rouhan G, Wang CN, Quandt D. 2009. Identifying a mysterious aquatic fern gametophyte. Plant Systematics and Evolution 281: 77–86. Little DP, Barrington DS. 2003. Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). American Journal of Botany 90: 508–514. Liu H-M, Zhang X-C, Chen Z-D, Dong S-Y, Qiu Y-L. 2007. Polyphyly of the fern family Tectariaceae sensu Ching: insights from cpDNA sequence data. Science in China Series C: Life Sciences 50: 789–798. Liu H-M, Zhang X-C, Wang W, Qiu Y-L, Chen Z-D. 2007. Molecular phylogeny of the fern family

Plants of the World

677

FURTHER READING Dryopteridaceae inferred from chloroplast rbcL and atpB genes. International Journal of Plant Sciences 168: 1311–1323. Liu H-M, Zhang X-C, Wang W, Zeng H. 2012. Molecular phylogeny of the endemic fern genera Cyrtomidictyum and Cyrtogonellum (Dryopteridaceae) from East Asia. Organisms Diversity, Evolution 10: 57–68. Moran RC. 1987. Monograph of the Neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural History Survey Bulletin 34: 1–138. Moran RC, Labiak P, Sundue M. 2010. Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae). Brittonia 62: 337–356. Moran RC, Labiak P, Sundue M. 2010. Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae). International Journal of Plant Sciences 171: 547–559. Nayar BK, Bajpai N, Chandra S. 1968. Contributions to the morphology of the fern genus Oleandra. Botanical Journal of the Linnean Society 60: 265–282. Pérez-García B, Mendoza A. 2005. Comparative studies of the gametophytes of five New World species of Tectaria (Tectariaceae). American Fern Journal 95: 141–152. Potes A. 2010. Comparative anatomy of the nectaries of Aglaomorpha and Drynaria (Polypodiaceae). American Fern Journal 100: 80–92. Prado J, Moran RC. 2008. Revision of the Neotropical species of Triplophyllum (Tectariaceae). Brittonia 60: 103–130. Rakotondrainibe F. 2009. Le genre Triplophyllum Holt t um (Pter idophy ta, Tectar iaceae) à Madagascar. Adansonia 31: 235–248. Schneider H, Smith AR, Cranfill R, Hildebrand TJ, Haufler CH, Ranker TA. 2004. Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Molecular Phylogenetics and Evolution 31: 1041–1063. Schneider H, Kreier HP, Wilson R, Smith AR. 2006. The Synammia enigma: evidence for a temperate lineage of polygrammoid ferns (Polypodiaceae, Polypodiidae) in southern South America. Systematic Botany 31: 31–41. Sen T, Sen U, Holttum RE. 1972. Morphology and anatomy of the genera Davallia, Araiostegia and Davallodes, with a discussion on their affinities. Kew Bulletin 27: 217–243. Sessa EB, Zimmer EA, Givnish TJ. 2012. Reticulate evolution on a global scale: a nuclear phylogeny for New World Dryopteris (Dryopteridaceae). Molecular Phylogenetics and Evolution 64: 563–581. Shankar R, Khare PK. 1991. Pharmacognostic studies on Hypodematium crenatum. Pharmaceutical Biology 29: 169–175. Skog JE, Mickel JT, Moran RC, Volovsek M, Zimmer EA. 2004. Molecular studies of representative species in the fern genus Elaphoglossum (Dryopteridaceae) based on cpDNA sequences rbcL, trnL-F, and rps4-trnS. International Journal of Plant Sciences 165: 1063–1075. Smith AR. 1986. Revision of the Neotropical fern genus Cyclodium. American Fern Journal 76: 56–98. Stokey AG, Atkinson LR. 1958. The gametophyte of the Grammitidaceae. Phytomorphology 8: 391–403. Sundue M, Islam MB, Ranker TA. 2010. Systematics of grammitid ferns (Polypodiaceae): using

678

Christenhusz, Fay & Chase

morphology and plastid sequence data to resolve the circumscriptions of Melpomene and the polyphyletic genera Lellingeria and Terpsichore. Systematic Botany 35: 701–715. Tindale MD. 1957. A preliminary revision of the genus Lastreopsis Ching. Victorian Naturalist 73: 180–185. Tsutsumi C, Kato M. 2006. Evolution of epiphytes in Davalliaceae and related ferns. Botanical Journal of the Linnean Society 151: 495–510. Von Euw J, Lounasmaa M, Reichstein T, Widén C-J. 1980. Chemotaxonomy in Dryopteris and related genera. Studia Geobotanica 1: 275–311. Walker TG. 1972. The anatomy of Maxonia apiifolia: a climbing fern. British Fern Gazette 10: 241–250. Wang FG, Barratt S, Falcón W, Fay MF, Lehtonen S, Tuomisto H, Xing FW, Christenhusz MJM. 2014. On the monophyly of subfamily Tectarioideae (Polypodiaceae) and the phylogenetic placement of some associated fern genera. Phytotaxa 164: 1–16. Wang L, Qia X-P, Xiang Q-P, Heinrichs J, Schneider H, Zhang X-C. 2010. Phylogeny of the Paleotropical fern genus Lepisorus (Polypodiaceae, Polypodiopsida) inferred from four chloroplast DNA regions. Molecular Phylogenetics and Evolution 54: 211–225. Wei X-P, Zhang Z-C. 2013. Species delimitation in the fern genus Lemmaphyllum (Polypodiaceae) based on multivariate analysis of morphological variation. Journal of Systematics and Evolution 51: 485–496. Zhang L-B, Zhang L. 2012. The inclusion of Acrophorus, Diacalpe, Nothoperanema, and Peranema in Dryopteris: the molecular phylogeny, systematics, and nomenclature of Dryopteris subg. Nothoperanema (Dryopteridaceae). Taxon 61: 1199–1216. 25. CYCADACEAE SAGO FAMILY De Laubenfels DJ, Adema F. 1998. A taxonomic revision of the genera Cycas and Epicycas gen. nov. (Cycadaceae) Blumea 43: 351–400. Hill KD 2008. The genus Cycas (Cycadaceae) in China. Telopea 12: 71–118. Nagalingum NS, Marshall CR, Quental TB, Rai HS, Little DP, Mathews S. 2011. Recent synchronous radiation of a living fossil. Science 334: 796–799. 26. ZAMIACEAE COONTIE FAMILY Arnold CA. 1953. Origin and relationships of the cycads. Phytomorphology 3: 51–65. Dehgan B, Dehgan NB. 1988. Comparative pollen morphology and taxonomic affinities in Cycadales. American Journal of Botany 75: 1501–1516. Delavoryas T. 1982. Perspectives on the origin of cycads and cycaeoids. Review of Palaeobotany and Palynology 37: 115–132. Hill KD, Chase MW, Stevenson DW, Hills HG, Schutzman B. 2003. The families and genera of cycads — a molecular phylogenetic analysis of Cycadophyta based on nuclear and plastid DNA sequences. International Journal of Plant Science 164: 933–948. Moretti A, Caputo P, Cozzolino S, De Luca P, Gaudio L, Gigliano Siniscalco S, Stevenson DW. 1993. A phylogenetic analysis of Dioon (Zamiaceae). American Journal of Botany 80: 204–214. Nagalingum NS, Marshall CR, Quental TB, Rai HS, Little DP, Mathews S. 2011. Recent synchronous radiation of a living fossil. Science 334: 796–799. Passalia MG, Del Fueyo G, Archangelsky S. 2010.

An early Cretaceous zamiaceous cycad of south west Gondwana: Restrepophyllum gen. nov. from Patagonia, Argentina. Review of Palaeobotany and Palynology 161: 137–150. Thieret JW. 1958. Economic botany of the cycads. Economic Botany 12: 3–41. Whitelock LM. 2002. The cycads. Timber Press, Portland. Whiting MG. 1963. Toxicity of cycads. Economic Botany 17: 270–302. Zgurski JM, Rai HS, Fai QM, Bogler DJ, FranciscoOrtega J, Graham SW. 2008. How well do we understand the overall backbone of cycad phylogeny? New insights from a large, multigene plastid data set. Molecular Phylogenetics and Evolution 47: 1232–1237. 27. GINKGOACEAE MAIDENHAIR-TREE FAMILY Critchfield WB. 1970. Shoot growth and heterophylly in Ginkgo biloba. Botanical Gazette 131: 150–162. Del Tredici P, Ling Hsieh, Yang Guang. 1992. The ginkgos of Tian Mu Shan. Conservation Biology 6: 202–209. Franklin AH. 1959. Ginkgo biloba L.: historical summary and bibliography. Virginia Journal of Science, n. s. 10: 131–176. Gunckel JE, Wetmore RH. 1946. Studies of development in long shoots and short shoots of Ginkgo biloba L. American Journal of Botany 33: 285–295, 532–543. Hori T, Ridge RW, Tulecke W, Tremouillaux-Guiller J, Tobe H, Del Tredici P (eds). 1997. Ginkgo biloba — a global treasure. Springer, Tokyo. Kaempfer E. 1712. Amoenitatum exoticarum politicophysico-medicarum. HW Meyer, Lemgo. Karstens WKH. 1945. Variability in the female reproductive organs of Ginkgo biloba L. Blumea 5: 532–553. Seward AC, Gowan J. 1900. The maidenhair tree (Ginkgo biloba L.). Annals of Botany (London) 14: 109–154. Sun C, Dilcher DL, Wang H, Sun G, Ge Y. 2008. A study of Ginkgo leaves from the Middle Jurassic of Inner Mongolia, China. International Journal of Plant Sciences 169: 1128–1139. Tralau H. 1968. Evolutionary trends in the genus Ginkgo. Lethalia 1: 63–101. Van Beek TA (ed.). 2000. Ginkgo biloba. Harwood, Amsterdam. 28. WELWITSCHIACEAE TUMBO FAMILY Bustard L. 1990. The ugliest plant of the world: the story of Welwitschia mirabilis. Kew Magazine 7: 85–90. Carlquist S, Gowans DA. 1995. Secondary growth and wood histology of Welwitschia. Botanical Journal of the Linnean Society 118: 107–121. Dilcher DL, Bernardes-de Oliveira ME, Pons D, Lott TA. 2005. Welwitschiaceae from the Lower Cretaceous of northeastern Brazil. American Journal of Botany 92: 1294–1310. Henschel JR, Seely MR. 2004. Long-term growth patterns of Welwitschia mirabilis, a long-lived plant of the Namib Desert (including a bibliography). Plant Ecology 150: 7–26. Leuenberger BE. 2001. Welwitschia mirabilis (Welwitschiaceae), male cone characters and a new subspecies. Willdenowia 31: 357–381. Martens P. 1959. Études sur les Gnetales III: structure et ontogenèse du cone et de la fleur femelles de Welwitschia mirabilis. Cellule 60: 169–268.

FURTHER READING Martens P. 1977. Welwitschia mirabilis and neoteny. American Journal of Botany 64: 916–920. Rodin RJ. 1953. Distribution of Welwitschia mirabilis. American Journal of Botany 40: 280–285. Rodin RJ. 1958. Leaf anatomy of Welwitschia II. American Journal of Botany 45: 96–103. Steyn EMA, Smith GF. 1999. Welwitschiaceae. Species plantarum: flora of the world, vol. 3. International Organisation for Plant Information, Melbourne. 8 pp. Welwitsch FMJ. 1861. On the botany of Benguela, Mossamedes, etc., in western Africa. Journal of the Linnaean Society (Botany) 5: 182–187. Willert DJ von. 1985. Welwitschia mirabilis — new aspects in the biology of an old plant. Advances in Botanical Research 11: 157–191. 29. GNETACEAE EMPING FAMILY Carlquist S, Robinson AA. 1995. Wood and bark anatomy of the African species of Gnetum. Botanical Journal of the Linnean Society 118: 123–137. Kato M, Inoue T, Nagamitsu T. 1995. Pollination biology of Gnetum (Gnetaceae) in a lowland mixed dipterocarp forest in Sarawak. American Journal of Botany 82: 862–868. Markgraf F. 1929. Monographie der Gattung Gnetum. Bulletin du Jardin Botanique de Buitenzorg 10: 407–507. Muhammad AF, Sattler R. 1982. Vessel structure of Gnetum and the origin of angiosperms. American Journal of Botany 69: 1004–1021. Shiembo PN, Newton AC, Leakey RRB. 1996. Vegetative propagation of Gnetum africanum Welw., a leafy vegetable from West Africa. Journal of Horticultural Science 71: 149–155. Takaso T, Bouman F. 1986. Ovule and seed ontogeny in Gnetum gnemon L. Botanical Magazine (Tokyo) 99: 241–266. Won H, Renner SS. 2006. Dating, dispersal and radiation in the gymnosperm Gnetum (Gnetales) — clock calibration when outgroup relationships are uncertain. Systematic Biology 55: 610–622. 30. EPHEDRACEAE JOINTFIR FAMILY Carlquist S. 1988. Near-vessellessness in Ephedra and its significance. American Journal of Botany 75: 598–601. Carlquist S. 1989. Wood and bark anatomy of the New World species of Ephedra. Aliso 12: 441–483. Carlquist S. 1992. Wood, bark and pith anatomy of Old World species of Ephedra and summary for the genus. Aliso 13: 255–295. Cutler HC. 1939. Monograph of the North American species of the genus Ephedra. Annals of the Missouri Botanical Garden 26: 373–427. Eames AJ. 1952. Relationships of the Ephedrales. Phytomorphology 2: 79–100. Ickert-Bond SM, Wojciechowski MF. 2004. Phylogenetic relationships in Ephedra (Gnetales): evidence from nuclear and chloroplast DNA sequence data. Systematic Botany 29: 834–849. Moussel B. 1978. Double fertilisation in the genus Ephedra. Phytomorphology 28: 336–345. Rydin C, Pedersen KR, Crane PR, Friis EM. 2006. Former diversity of Ephedra (Gnetales): evidence from early Cretaceous seeds from Portugal and North America. Annals of Botany 98: 123–140. Steeves MW, Barghoorn ES. 1959. The pollen of Ephedra. Journal of the Arnold Arboretum 40: 221–259. Tanaka T, Ohba K, Sakai E. 1995. Comparison of

the constituents of Ephedra herbs from various countries — on ephedrine type alkaloids. Natural medicines 49: 418. Zhang JS, Tian SZ, Lou C. 1989. Quality evaluation of twelve species of Chinese Ephedra (ma huang). Acta Pharmica Sinica 24: 865–871. 31. PINACEAE PINE FAMILY Bailey IW. 1909. The structure of wood in the Pineae. Botanical Gazette 48: 47–55. Critchfield WB. 1986. Hybridisation and classification of the white pines (Pinus section Strobus). Taxon 35: 647–656. Farjon A. 1984. Pines: drawings and descriptions of the genus Pinus. Brill, Backhuys, Leiden. Farjon A. 1989. A second revision of the genus Keteleeria Carrière (taxonomic notes on Pinaceae II). Notes from the Royal Botanical Garden Edinburgh 46: 81–99. Farjon A. 1990. Pinaceae: drawings and descriptions of the genera Abies, Cedrus, Pseudolarix, Keteleeria, Nothotsuga, Tsuga, Cathaya, Pseudotsuga, Larix and Picea. Koeltz Scientific Books, Königstein. Farjon A, Styles BT. 1997. Pinus (Pinaceae). Flora Neotropica Monograph 75. The New York Botanical Garden, New York. Frankis MP. 1989. Generic inter-relationships in Pinaceae. Notes from the Royal Botanic Garden Edinburgh 45: 527–548. Gernandt DS, Geada López G, Garcia SO, Liston A. 2005. Phylogeny and classification of Pinus. Taxon 54: 29–42. Gros-Louis M-C, Bousquet J, Pâques LE, Isabel NE. 2005. Species-diagnostic markers in Larix spp. based on RAPDs and nuclear, cpDNA, and mtDNA gene sequences, and their phylogenetic implications. Tree Genetics, Genomes 1: 50–63. Hu YS, Wang FH. 1984. Anatomical studies of Cathaya (Pinaceae). American Journal of Botany 71: 727–735. Jain KK. 1976. Evolution of wood structure in Pinaceae. Israel Journal of Botany 25: 28–33. Liston A, Gernandt DS, Vining TF, Campbell CS, Piñero D. 2003. Molecular phylogeny of Pinaceae and Pinus, in: Mill RR (ed.). Proceedings of the fourth international conifer conference [Acta Horticulturae No. 615]. International Society for Horticultural Science, Leuven, pp. 107–114. Malusa J. 1992. Phylogeny and biogeography of the pinyon pines (Pinus subsect. Cembroides). Systematic Botany 17: 42–66. Miller CN Jr. 1976. Early evolution in the Pinaceae. Review of Palaeobotany and Palynology 21: 101–117. Mirov NT. 1967. The genus Pinus. Ronald, New York. Mirov NT, Hasbrouck J. 1976. The story of pines. Indiana University, Bloomington. Perry JP. 1991. The pines of Mexico and Central America. Timber Press, Portland. Ran J-H, Wei X-X, Wang X-Q. 2006. Molecular phylogeny and biogeography of Picea (Pinaceae): implications for phylogeographical studies using cytoplasmic haplotypes. Molecular Phylogenetics and Evolution 41: 405–419. Sigurgeirsson A, Szmidt AE. 1993. Phylogenetic and biogeographic implications of chloroplast DNA variation in Picea. Nordic Journal of Botany 13: 233–246. Viguie MT, Gaussen H. 1929. Revision du genre Abies. Bulletin de la Société d’Histoire Naturelle de Toulouse 57: 369–434, 58: 245–564. Wei X-X, Wang X-Q. 2004. Recolonisation and

radiation in Larix (Pinaceae): evidence from nuclear ribosomal DNA paralogues. Molecular Ecology 13: 3115–3123. 32. ARAUCARIACEAE KAURI-TREE FAMILY Chambers TC, Drinnan AN, McLoughlin S. 1998. Some morphological features of Wollemi pine (Wollemia nobilis: Araucariaceae) and their comparison to Cretaceous plant fossils. International Journal of Plant Science 159: 160–171. De Laubenfels DJ. 1988. Araucariaceae. Flora Malesiana 10: 419–442. Gilmore S, Hill KD. 1997. Relationships of the Wollemi pine (Wollemia nobilis) and a molecular phylogeny of the Araucariaceae. Telopea 7: 275–291. Haines RJ, Prakash N, Nikles DG. 1984. Pollination in Araucaria Juss. Australian Journal of Botany 32: 583–594. Jones WG, Hill KD, Allen JM. 1995. Wollemia nobilis, a new living Australian genus and species in the Araucariaceae. Telopea 6: 173–176. Kershaw P, Wagstaff B. 2001. The southern conifer family Araucariaceae: history, status, and value for paleoenvironmental reconstruction. Annual Review of Ecology and Systematics 32: 397–414. Kunzmann L. 2007. Araucariaceae (Pinopsida): aspects in palaeobiogeography and palaeobiodiversity in the Mesozoic. Zoologischer Anzeiger 246: 257–277. Manauté J, Jaffré T, Veillon J-M, Kranitz M-L. 2003. Revue des Araucariaceae de Nouvelle-Calédonie. Publication IRD/Province Sud de la NouvelleCalédonie, Nouméa. Setoguchi H, Osawa TA, Pintaud J-C, Jaffré, T, Veillon J-M. 1998. Phylogenetic relationships within Araucariaceae based on rbcL gene sequences. American Journal of Botany 85: 1507–1516. Seward AC, Ford SO. 1906. The Araucariaceae, recent and extinct. Philosophical Transactions of the Royal Society of London, series B 198: 305–411. Stockey RA.1994. Mesozoic Araucariaceae: morphology and systematic relationships. Journal of Plant Research 107: 493–502. Veillon J-M. 1980. Architecture des espèces néocalédoniennes du genre Araucaria. Candollea 35: 609–640. Whitmore TC. 1980. A monograph of Agathis. Plant Systematics and Evolution 135: 41–69. Whitmore TC, Page CN. 1980. Evolutionary implications of the distribution and ecology of the tropical conifer Agathis. New Phytologist 84: 407–416. 33. PODOCARPACEAE YEWPINE FAMILY Buchholz JT. 1941. Embryogeny of the Podocarpaceae. Botanical Gazette 103: 1–37. Buchholz JT, Gray NE. 1948. A taxonomic revision of Podocarpus I. The sections of the genus and their subdivisions with special reference to leaf anatomy. Journal of the Arnold Arboretum 29: 49–63. Conran JG, Woods GM, Martin PG, Dowd JM, Quinn CJ, Gadek PA, Price RA. 2000. Generic relationships within and between the gymnosperm families Podocarpaceae and Phyllocladaceae based on an analysis of the chloroplast gene rbcL. Australian Journal of Botany 48: 715–724. De Laubenfels DJ. 1959. Parasitic conifer found in New Caledonia. Science 130: 97. De Laubenfels DJ. 1969. A revision of the Malesian and Pacific rainforest conifers. 1. Podocarpaceae. Journal of the Arnold Arboretum 50: 274–369. De Laubenfels DJ. 1985. A taxonomic revision of the genus Podocarpus. Blumea 30: 251–211.

Plants of the World

679

FURTHER READING De Laubenfels DJ. 1987. Revision of the genus Nageia. Blumea 32: 209–211. Gray NE. 1969. An interpretation of Podocarpus in time and space. Bulletin of the Georgia Academy of Science 27: 144–147. Hair JB, Beuzenberg EJ. 1958. Chromosomal evolution in the Podocarpaceae. Nature (London) 181: 1584–1586. Herbert J, Hollingsworth PM, Gardner MF, Mill RR, Thomas PI, Jaffré, T. 2002. Conservation genetics and phylogenetics of New Caledonian Retrophyllum (Podocarpaceae) species. New Zealand Journal of Botany 40: 175–188. Kelch DG. 1997. The phylogeny of the Podocarpaceae based on morphological evidence. Systematic Botany 22: 113–131. Kelch DG. 1998. Phylogeny of Podocarpaceae: comparison of evidence from morphology and 18S rDNA. American Journal of Botany 85: 986–996. Keng H. 1963. Taxonomic position of Phyllocladus and the classification of the conifers. Garden’s Bulletin Singapore 20: 123–126. Keng H. 1978. T he genus Phyllocladus (Phyllocladaceae). Journal of the Arnold Arboretum 59: 249–273. Köpke E, Musselman LJ, De Laubenfels DJ. 1981. Studies on the anatomy of Parasitaxus ustus and its root connections. Phytomorphology 31: 85–92. Orr MY. 1943. The leaf anatomy of Podocarpus. Transactions and Proceedings of the Botanical Society of Edinburgh 34: 1–54. Quinn CJ. 1970. Generic boundaries in the Podocarpaceae. Proceedings of the Linnean Society of New South Wales 94: 166–172. Quinn CJ. 1982. Taxonomy of Dacrydium Sol. ex Lamb. emend. de Laub. (Podocarpaceae). Australian Journal of Botany 30: 311–320. Sinclair WT, Mill RR, Gardner MF, Woltz P, Jaffré T, Preston J, Hollingsworth ML, Ponge A, Möller M. 2002. Evolutionary relationships of the New Caledonian heterotrophic conifer Parasitaxus usta (Podocarpaceae), inferred from chloroplast trnL-F intron/spacer and nuclear rDNA ITS2 sequences. Plant Systematics and Evolution 233: 79–104. Turner BJ, Cernusak LA (eds). 2011. Ecology of the Podocarpaceae in tropical forests. Smithsonian Institution Scholarly Press, Washington DC. Young M. 1910. The morphology of the Podocarpineae. Botanical Gazette 50: 81–100. 34. SCIADOPITYACEAE UMBRELLA-PINE FAMILY Dickson A. 1866. On the phylloid shoots of Sciadopitys verticillata Sieb. et Zucc. Journal of Botany 4: 224–225. Farjon A. 2005. A monograph of Cupressaceae and Sciadopitys. Royal Botanic Gardens, Kew, Richmond. Florin R. 1922. On the geological history of the Sciadopitineae. Svensk Botanisk Tidskrift 6: 260–270. Hayata B. 1931. The Sciadopityaceae represented by Sciadopitys verticillata Sieb. et Zucc. an endemic species of Japan. Botanical Magazine of Tokyo 45: 567–569. Roth L. 1962. Histogenese und morphologische Deutung der Doppelnadeln von Sciadopitys. Flora 152: 1–23. Takaso T, Tomlinson PB. 1991. Cone and ovule development in Sciadopitys (Taxodiaceae-Coniferales). American Journal of Botany 78: 417–428.

680

Christenhusz, Fay & Chase

35. CUPRESSACEAE CYPRESS FAMILY Adams RP, Bartel JA, Price RA. 2009. A new genus, Hesperocyparis, for the cypresses of the western hemisphere. Phytologia 91: 160–185. Buchholz JT. 1939. The generic segregation of the sequoias. American Journal of Botany 26: 535–538. Debreczy Z, Musial K, Price RA, Racz I. 2009. Relationships and nomenclatural status of the Nootka cypress (Callitropsis nootkatensis, Cupressaceae). Phytologia 91: 140–159. Eckenwalder JE. 1976. Re-evaluation of Cupressaceae and Taxodiaceae: a proposed merger. Madroño 23: 237–256. Farjon A, Hiep NT, Harder DK, Loc PK, Averyanov L. 2002. A new genus and species in Cupressaceae (Coniferales) from northern Vietnam, Xanthocyparis vietnamensis. Novon 12: 179–189. Farjon A. 2005. A monograph of Cupressaceae and Sciadopitys. Royal Botanic Gardens, Kew, Richmond. Gadek PA, Alpers DL, Heslewood MM, Quinn CJ. 2000. Relationships within Cupressaceae sensu lato: a combined morphological and molecular approach. American Journal of Botany 87: 1044–1057. Hu HH, Cheng WC. 1948. On the new family Metasequoiaceae and on Metasequoia glyptostroboides, a living species of the genus Metasequoia found in Szechuan and Hupeh. Bulletin of the Fan Memorial Institute of Biology II 1: 153–159. Li H-L. 1953. A reclassification of Libocedrus and Cupressaceae. Journal of the Arnold Arboretum 34: 17–34. Little DP, Schwarzbach AE, Adams RP, Hsieh C-F. 2004. The circumscription and phylogenetic relationships of Callitropsis and the newly described genus Xanthocyparis (Cupressaceae). American Journal of Botany 91: 1872–1881. Mao K, Hao G, Liu J, Adams RP, Milne RI. 2010. Diversification and biogeography of Juniperus (Cupressaceae): variable diversification rates and multiple intercontinental dispersals. New Phytologist 188: 254–272. Page CN. 1980. The earliest known find of living Taiwania (Taxodiaceae). Kew Bulletin 34: 527–528. Peirce AS. 1937. Systematic anatomy of the woods of the Cupressaceae. Tropical Woods 49: 5–21. Terry RG, Bartel JA, Adams RP. 2012. Phylogenetic relationships among the New World cypresses (Hesperocyparis; Cupressaceae): evidence from noncoding chloroplast DNA sequences. Plant Systematics and Evolution 298: 1987–2000. Cheng Y, Chaw S, Nicholson R, Tripp K. 2000. Phylogeny of Taxaceae and Cephalotaxaceae genera inferred from chloroplast matK gene and nuclear rDNA ITS region. Molecular Phylogenetics and Evolution 14: 353–365. Ferguson DK. 1978. Some current research on fossil and recent taxads. Review of Palaeobotany and Palynology 26: 213–226. Florin R. 1948. On the morphology and relationship of the Taxaceae. Botanical Gazette 110: 31–39. Hao DC, Xiao PG, Huang BL, Ge GB, Yang L. 2008. Interspecific relationships and origins of Taxaceae and Cephalotaxaceae revealed by partitioned Bayesian analyses of chloroplast and nuclear DNA sequences. Plant Systematics and Evolution 276: 89–104. Keng H. 1969. Aspects of the morphology of Amentotaxus formosana with a note on the taxonomic position of the genus. Journal of the Arnold Arboretum 50: 432–445.

Li J, Davis CC, Donoghue MJ, Kelley S, Del Tredici P. 2001. Phylogenetic relationships of Torreya (Taxaceae) inferred from sequences of nuclear ribosomal DNA ITS region. Harvard Papers in Botany 6: 275–281. Lang X-D, Su J-R, Lu S-G, Zhang Z-J. 2013. A taxonomic revision of the genus Cephalotaxus (Taxaceae). Phytotaxa 84: 1–24. Pilger RKF. 1916. Die Taxales. Mitteilungen der Deutschen Dendrologischen Gesellschaft 25: 1–28. Price RA. 2003. Generic and familial relationships of the Taxaceae from rbcL and matK sequence comparisons, in: Mill RR (ed.). Proceedings of the fourth international conifer conference [Acta Horticultural No. 615]. International Society for Horticultural Science, Leuven. Sahni B. 1920. On certain archaic features of the seed of Taxus baccata, with remarks on the the antiquity of the Taxineae. Annals of Botany (London) 34: 117–133. Tripp KE. 1995. Cephalotaxus: the plum yews. Arnoldia 55: 24–39. 37. AMBORELLACEAE AMBORELLA FAMILY Bailey IW, Swamy BGL. 1948. Amborella trichopoda Baill., a new morphological type of vesselless dicotyledon. Journal of the Arnold Arboretum 29: 245–253. Borsch T, Hilu KW, Wilde V, Neinhuis C, Barthlott W. 2000. Phylogenetic analysis of non-coding chloroplast DNA sequences reveals Amborella as basalmost angiosperm. American Journal of Botany 87 (suppl.): 115. Buzgo M, Soltis PS, Soltis DE. 2004. Floral developmental morphology of Amborella trichopoda (Amborellaceae). International Journal of Plant Sciences 165: 925–947. Endress PK, Igersheim A. 2000. The reproductive structures of the basal angiosperm Amborella trichopoda (Amborellaceae). International Journal of Plant Sciences 161: S237–S248. Goremykin VV, Viola R, Hellwig Fh. 2009. Removal of noisy characters from chloroplast genome-scale data suggests revision of phylogenetic placements of Amborella and Ceratophyllum. Journal of Molecular Evolution 68: 197–204. Matthews S, Donoghue MJ. 1999. The root of angiosperm phylogeny inferred from duplicate phytochrome genes. Science 286: 947–950. Parkinson CL, Adams KL, Palmer JD. 1999. Multigene analysis identify the three earliest lineages of extant f lowering plants. Current Biology 9: 1485–1491. Soltis PS, Soltis DE, Chase MW. 1999. Angiosperm phylogeny inferred from multiple genes as a tool for comparative biology. Nature 402: 402–404. Soltis DE, Albert VA, Leebens-Mack J, Palmer JD, Wing RA, dePamphilis CW, Ma H, Carlson JE, Altman N, Kim S, Kerr Wall P, Zuccolo A, Soltis PS. 2008. The Amborella genome: an evolutionary reference for plant biology. Genome Biology 9: 402. Thien LB, Sage TL, Jaffré T, Bernhardt P, Pontieri V, Weston PH, Malloch D, Azuma H, Graham SW, McPherson MA, Rai HS, Sage RF, Dupre J-L. 2003. The population structure and floral biology of Amborella trichopoda (Amborellaceae). Annals of the Missouri Botanical Garden 90: 466–490. Williams JH. 2009. Amborella trichopoda (Amborellaceae) and the evolutionary developmental origins of the angiosperm progamic phase. American Journal of Botany 96: 144–165.

FURTHER READING 38. HYDATELLACEAE WATERTUFTS FAMILY Cooke DA 1983. The seedling of Trithuria (Hydatellaceae). Victorian Naturalist 100: 68–69. Friedman WE. 2008. Hydatellaceae are water lilies with gymnospermaceous tendencies. Nature 453: 94–97. Hamann U. 1976. Hydatellaceae — a new family of Monocotyledonae. New Zealand Journal of Botany 14: 193–196. Rudall PJ, Sokoloff DD, Remizowa MV, Conran JG, Davis JI, Macfarlane TD, Stevenson DW. 2007. Morphology of Hydatellaceae, an anomalous aquatic family recently recognized as an earlydivergent angiosperm lineage. American Journal of Botany 94: 1073–1092. Saarela JM, Rai HS, Doyle JA, Endress PK, Mathews S, Marchant AD, Briggs BG, Graham SW. 2007. Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree. Nature 446: 312–315. Sokoloff D, Remizowa MV, Macfarlane TD, Rudall PJ. 2008. Classification of the early-divergent angiosperm family Hydatellaceae: one genus instead of two, four new species and sexual dimorphism in dioecious taxa. Taxon 57: 179–200. Taylor ML, Macfarlane TD, Williams JH. 2010. Reproductive ecology of the basal angiosperm Trithuria submersa (Hydatellaceae). Annals of Botany 106: 909–920. Yadav SR, Janarthanam MK. 1995. Trithuria konkanensis (Hydatellaceae), eine neue Art aus Indien. Aqua-Planta 3: 91–101. 39. CABOMBACEAE FANWORT FAMILY Chrysler MA. 1938. The winter buds of Brasenia. Bulletin of the Torrey Botanical Club 65: 277–283. Collinson ME. 1980. Recent and Tertiary seeds of the Nymphaeaceae sensu lato with a revision of Brasenia ovula (Brong.) Reid and Chandler. Annals of Botany 46: 603–632. Fassett NC. 1953. A monograph of Cabomba. Castanea 13: 116–128. Ito M. 1986. Studies on the floral morphology and anatomy of Nymphaeales III. Floral anatomy of Brasenia scheberi Gmel. and Cabomba caroliniana A.Gray. Botanical Magazine (Tokyo) 99: 169–184. Kakuta M, Misaki A. 1979. The polysaccharide of junsai (Brasenia scheberi) mucilage: fragmentation analysis by successive Smith degradations and partial acid hydrolysis. Agricultural Biology and Chemistry 43: 1269–1276. Mohr BAR, Bernardes-de-Oliviera MEC, Taylor DW. 2008. Pluricarpellatia, a nymphaealean angiosperm from the Lower Cretaceous of northern Gondwana (Crato Formation, Brazil). Taxon 57: 1147–1158. Ørgaard M. 1991. The genus Cabomba (Cabombaceae) — a taxonomic study. Nordic Journal of Botany 11: 179–203. Ørgaard M, Van Bruggen HWE, Van der Vlugt PJ. 1992. Cabombaceae. Aqua-Planta Suppl. 3: 1–44. Raciborski M. 1894. Die Morphologie des Cabombeen und Nymphaeaceen. Flora 78: 244–279. Taylor ML, Williams JH. 2009. Consequences of pollination syndrome evolution for postpollination biology in an ancient angiosperm family. International Journal of Plant Sciences 170: 584–598. 40. NYMPHAEACEAE WATERLILY FAMILY Borsch T, Löhne C, Wiersema JH. 2008. Phylogeny and evolutionary patterns in Nymphaeales:

integrating genes, genomes and morphology. Taxon 57: 1052–1081. Borsch T, Löhne C, Mbaye MS, Wiersema JH. 2012. Towards a complete species tree of Nymphaea: shedding further light on subgenus Brachyceras and its relationships to the Australian waterlilies. Telopea 13: 193–217. Conard HS. 1905. The waterlilies. A monograph of the genus Nymphaea. Publications of the Carnegie Institute of Washington 4: 1–292. Emboden WA. 1981. Transcultural use of narcotic water lilies in ancient Egyptian and Maya drug ritual. Journal of Ethnopharmacology 3: 39–83. Friis EM, Pedersen KR, Von Balthazar M, Grimm GW, Crane PR. 2009. Monetianthus mirus gen. et sp. nov., a nymphaealean flower from the Early Cretaceous of Portugal. International Journal of Plant Sciences 170: 1986–1101. Holway T. 2013. The flower of empire: an Amazonian water lily, the quest to make it bloom, and the world it created. Oxford University Press, Oxford. Lawson G. 1851. The royal waterlily of South America, and the waterlilies of our own land: their history and cultivation. James Hogg, Edinburgh. Löhne C, Yoo MJ, Borsch T, Wiersema JH, Wilde V, Bell CD, Barthlott W, Soltis DE, Soltis PS. 2008. Biogeography of Nymphaeales: extant patterns and historical events. Taxon 57: 1123–1146. Magallón SA, Hilu KW, Quandt D. 2013. Land plant evolutionary timeline: gene effects are secondary to fossil constraints in relaxed clock estimation of age and substitution rates. American Journal of Botany 100: 556–573. Padgett DG. 2007. A monograph of Nuphar (Nymphaeaceae). Rhodora 109: 1–95. Schneider EL. 1976. The floral anatomy of Victoria Schomb. (Nymphaeaceae). Journal of the Linnean Society, Botany 72: 115–148. Schneider EL, Tucker SC, Williamson PS. 2003. Floral development in the Nymphaeales. International Journal of Plant Sciences 164: S279–S292. Wiersema JH. 1987. A monograph of Nymphaea subgenus Hydrocallis (Nymphaeaceae). Systematic Botany Monographs 16: 1–112. 41. AUSTROBAILEYACEAE AUSTROBAILEYA FAMILY Bailey IW, Swamy BGL. 1949. The morphology and relationships of Austrobaileya. Journal of the Arnold Arboretum 30: 211–226. Behnke HD. 1986. Sieve element characters and the systematic position of Austrobaileya (Austrobaileyaceae). Plant Systematics and Evolution 152: 101–121. Carlquist S. 2001. Observations on the vegetative anatomy of Austrobaileya: habital, organographic and phylogenetic conclusions. Botanical Journal of the Linnean Society 135: 1–11. Dickison WC, Endress PL. 1983. Ontogeny of the stemnode-leaf vascular continuum of Austrobaileya. American Journal of Botany 70: 906–911. Endress PK. 1980. The reproductive structures and systematic position of the Austrobaileyaceae. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 101: 393–433. Endress PK. 1983. The early floral development of Austrobaileya. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 103: 481–493. Endress PK. 1984. The role of inner staminodes in the floral display of some relic Magnoliales. Plant Systematics and Evolution 146: 269–282.

Endress PK, Honegger R. 1980. The pollen of the Austrobaileyaceae and its phylogenetic significance. Grana 19: 177–182. Rüdenberg L. 1967. The chromosomes of Austrobaileya. Journal of the Arnold Arboretum 48: 241–244. White CT. 1933. Ligneous plants collected for the Arnold Arboretum in North Queensland by S. F. Kajewski in 1929. Contributions from the Arnold Arboretum of Harvard University 4: 1–113. White CT. 1948. A new species of Austrobaileya (Austrobaileyaceae) from Australia. Journal of the Arnold Arboretum 29: 255–256. 42. TRIMENIACEAE BITTERVINE FAMILY Bernhardt P, Sage T, Weston P, Azuma H, Lam M, Thien LB, Bruhl J. 2003. The pollination of Trimenia moorei (Trimeniaceae): floral volatiles, insect/wind pollen vectors and stigmatic selfincompatibility in a basal angiosperm. Annals of Botany 92: 445–458. Endress PK, Sampson FB. 1983. Floral structure and relationships of the Trimeniaceae. Journal of the Arnold Arboretum 64: 447–473. Friedman WE, Bachelier JB. 2013. Seed development in Trimenia (Trimeniaceae) and its bearing on the evolution of embryo-nourishing strategies in early flowering plants. American Journal of Botany 100: 906–915. Gilg E, Schlechter R. 1919. Über zwei pflanzengeographisch interessante Monimiaceen aus Deutsch Neu Guinea. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 55: 195–201. Money LL, Bailey IW, Swamy BGL. 1950. The morphology and relationships of the Monimiaceae. Journal of the Arnold Arboretum 31: 372–404. Rodenburg WF. 1971. A revision of the genus Trimenia (Trimeniaceae). Blumea 19: 3–15. Sampson FB, Endress PK. 1984. Pollen morphology in the Trimeniaceae. Grana 23: 129–137. Yamada T, Nishida H, Umebayashi M, Uemura K, Kato M. 2008. Oldest record of Trimeniaceae from the Early Cretaceous of northern Japan. BMC Evolutionary Biology 8: 135. 43. SCHISANDRACEAE STAR-ANISE FAMILY Bailey IW, Nast CG. 1948. Morphology and relationships of Illicium, Schisandra and Kadsura. I. Stem and leaf. Journal of the Arnold Arboretum 29: 77–89. Carlquist S. 1982. Wood anatomy of Illicium (Illiciaceae): phylogenetic, ecological and functional interpretations. American Journal of Botany 69: 1587–1598. Denk T, Oh IC. 2006. Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record. Plant Systematics and Evolution 256: 113–145. Hao G, Chye ML, Saunders RMK. 2001. A phylogenetic analysis of the Schisandraceae based on morphology and nuclear ribosomal (ITS) sequences. Botanical Journal of the Linnean Society 135: 401–411. Liu Z, Hao G, Luo YB, Thien LB, Rosso SW, Lu AM, Chen ZD. 2006. Phylogeny and androecial evolution in Schisandraceae, inferred from sequences of nuclear robosomal DNA ITS and chloroplast DNA trnL-F regions. International Journal of Plant Sciences 167: 539–550. Morris AB, Bell CD, Clayton JW, Judd WS, Soltis DE, Soltis PS. 2007. Phylogeny and divergence

Plants of the World

681

FURTHER READING time estimation in Illicium with implications for New World biogeography. Systematic Botany 32: 236–249. Oh IC, Denk T, Friis EM. 2003. Evolution of Illicium (Illiciaceae): mapping morphological characters on the molecular tree. Plant Systematics and Evolution 240: 175–209. Roberts ML, Haynes RR. 1983. Ballistic seed dispersal in Illicium (Illiciaceae). Plant Systematics and Evolution 143: 227–232. Romanov MS, Bobrov AVFC, Endress PK. 2013. Structure of the unusual explosive fruits of the early diverging angiosperm Illicium (Schisandraceae s.l., Austrobaileyales). Botanical Journal of the Linnean Society 171: 640–654. Saunders RM. 1998. Monograph of Kadsura (Schisandraceae). Systematic Botany Monographs 54: 1–106. Saunders RM. 2000. Monograph of Schisandra (Schisandraceae). Systematic Botany Monographs 58: 1–146. Smith AC. 1947. The families Illiciaceae and Schisandraceae. Sargentia 7: 1–224. 44. CANELLACEAE CANEEL-BARK FAMILY Bastos JK, Kaplan MAC, Gottleib OR. 1999. Drimane-type sesquiterpenoids as chemosystematic markers of Canellaceae. Journal of the Brazilian Chemical Society 10: 136–139. Bell CD, Soltis DE, Soltis PS. 2010. The age and diversification of the angiosperms re-revisited. American Journal of Botany 97: 1245–1258. Bonnett E. 1876. Essaie d’une monographie des Canellées. Parent, Paris. Feild TS, Hudson PJ, Balun L, Chatelet DS, Patino AA, Sharma CA, McLaren K. 2011. The ecophysiology of xylem hydraulic constraints by “basal” vessels in Canella winterana (Canellaceae). International Journal of Plant Sciences 172: 879–888. Igersheim A, Endress PK. 1997. Gynoecium diversity and systematics of the Magnoliales and winteroids. Botanical Journal of the Linnean Society 124: 213–271. Massoni J, Forest F, Sauquet H. 2014. Increased sampling of both genes and taxa improves resolution of phylogenetic relationships within Magnoliidae, a large and early-diverging clade of angiosperms. Molecular Phylogenetics and Evolution 70: 84–93. Miers J. 1858. On the Canellaceae. Annual Magazine of Natural History III, 1: 349–353. Müller S, Salomo K, Salazar J, Naumann J, Jaramillo AM, Neinhuis C, Field TS, Wanke S. 2015. Intercontinental long-distance dispersal of Canellaceae from the New to the Old World revealed by a nuclear single copy gene and chloroplast loci. Molecular Phylogenetics and Evolution 84: 205–219. Salazar J, Nixon K. 2008. New discoveries in the Canellaceae in the Antilles: how phylogeny can support taxonomy. Botanical Review 74: 103–111. Wilson TK. 1960. The comparative morphology of the Canellaceae. I. Synopsis of genera and wood anatomy. Tropical Woods 112: 1–27. Wilson TK. 1966. The comparative morphology of the Canellaceae. IV. Flora morphology and conclusions. American Journal of Botany 53: 336–343. 45. WINTERACEAE WINTER’S-BARK FAMILY Bailey IW. 1944. The comparative morphology of the Winteraceae. III. Wood. Journal of the Arnold Arboretum 25: 97–103.

682

Christenhusz, Fay & Chase

Carlquist S. 1983. Wood anatomy of Bubbia (Winteraceae), with comments on origin of vessels in dicotyledons. American Journal of Botany 70: 578–590. Deroin T. 2000. Notes on the vascular anatomy of the fruits of Takhtajania (Winteraceae) and its interpretation. Annals of the Missouri Botanical Garden 87: 398–406. Doust AN. 2000. Comparative floral ontogeny in Winteraceae. Annals of the Missouri Botanical Garden 87: 366–379. Doust AN, Drinnan AN. 2004. Floral development and molecular phylogeny support the generic status of Tasmannia (Winteraceae). American Journal of Botany 91: 321–331. Endress PK, Igersheim A, Sampson FB, Schatz GE. 2000. Floral structure of Takhtajania and its systematic position in Winteraceae. Annals of the Missouri Botanical Garden 87: 347–365. Feild TS, Chatelet DS, Balun L, Schilling E, Evans R. 2012. The evolution of angiosperm lianescence without vessals — climbing mode and wood structure-function in Tasmannia cordata (Winteraceae). New Phytologist 193: 229–240. Karol KG, Suh Y, Schatz GE, Zimmer EA. 2000. Molecular evidence for the phylogenetic position of Takhtajania in the Winteraceae: inference from nuclear ribosomal and chloroplast gene spacer sequences. Annals of the Missouri Botanical Garden 87: 414–432. Larsen L, Lorimer SD, Perry NB. 2007. Contrasting chemistry of fruits and leaves of two Pseudowintera species: sesquiterpene dialdehyde cinnamates and prenylated flavonoids. Biochemistry Systematics and Ecology 35: 286–292. Marquínez X, Lohmann LG, Salatino MLF, Salatino A, González F. 2009. Generic relationships and dating of lineages in Winteraceae based on nuclear (ITS) and plastid (rps16 and psbA-trnH) sequence data. Molecular Phylogenetics and Evolution 53: 435–449. Schrank E. 2013. New taxa of winteraceous pollen from the Lower Cretaceous of Israel. Review of Palaeobotany and Palynology 195: 19–25. Suh Y, Thien LB, Reeve HE, Zimmer EA. 1993. Molecular evolution and phylogenetic implications of internal transcribed spacer sequences of ribosomal DNA in Winteraceae. American Journal of Botany 80: 1042–1055. Thien LB, Bernhardt P, Gibbs GW, Pellmyr O, Bergström G, Groth I, McPherson G. 1985. The pollination of Zygogynum (Winteraceae) by a moth, Sabatinca (Micropterigidae): an ancient association? Science 227: 540–543. Thomas N, Bruhl JJ, Ford A, Weston PH. 2014. Molecular dating of Winteraceae reveals a complex biogeographical history involving both ancient Gondwanan vicariance and long-distance dispersal. Journal of Biogeography 41: 894–904. Vink W. 1988. Taxonomy in Winteraceae. Taxon 37: 691–698. 46. SAURURACEAE LIZARD’S-TAIL FAMILY Carlquist S, Dauer K, Nishimra SY. 1995. Wood and stem anatomy of Saururaceae with reference to ecology, phylogeny, and origin of the monocotyledons. IAWA Journal 16: 133–150. Liang HX. Karyomorphology of Gymnotheca and phylogeny of four genera in Saururaceae. Acta Botanica Yunnanica 13: 303–307. Liang HX, Tucker SC. 1990. Comparative studies of the floral vasculature in Saururaceae. American

Journal of Botany 77: 607–623. Meng SW, Chen ZD, Li DZ, Liang HX. 2002. Phylogeny of Saururaceae based on mitochondrial matR gene sequence data. Journal of Plant Research 115: 71–76. Meng SW, Douglas AW, Li DZ, Cheng ZD, Liang HX, Yang JB. 2003. Phylogeny of Saururaceae based on morphology and five regions from three plant genomes. Annals of the Missouri Botanical Garden 90: 592–602. Quibell CH. 1941. Floral anatomy and morphology of Anemopsis californica. Botanical Gazette 102: 749–758. Raju MVS. 1961. Morphology and anatomy of the Saururaceae I. Floral anatomy and embryology. Annals of the Missouri Botanical Garden 48: 107–124. Smith SY, Stockey RA. 2007. Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton chert. American Journal of Botany 94: 1642–1657. Smith SY, Stockey RA. 2007. Pollen morphology and ultrastructure of Saururaceae. Grana 46: 250–267. Takahashi M. 1986. Microsporogenesis in a parthenogenetic species, Houttuynia cordata Thunb. (Saururaceae). Botanical Gazette 147: 47–54. Tanaka H. 1979. Pollination of Saururus chinensis (Lour.) Baill. Journal of Japanese Botany 54: 221–224. Tucker SC. 1981. Inflorescence and floral development in Houttuynia cordata (Saururaceae). American Journal of Botany 68: 1017–1032. 47. PIPERACEAE PEPPER FAMILY Burger WC. 1977. The Piperales and the monocots. Botanical Review 43: 345–393. Görts-van Rÿn ARA. 2007. Piperaceae. Flora of the Guianas 24: 15– 163. Hoffstadt RE. 1916. The vascular anatomy of Piper methysticum. Botanical Gazette 62: 115–132. Homer HT, Wanke S, Samain MS. 2009. Evolution and systematic value of leaf crystal macropatterns in the genus Peperomia (Piperaceae). International Journal of Plant Sciences 170: 343–354. Jaramillo MA, Manos PS. 2001. Phylogeny and patterns of floral diversity in the genus Piper (Piperaceae). American Journal of Botany 88: 706–716. Jaramillo MA, Callejas R, Davidson C, Smith JF, Stevens AC, Tepe EJ. 2008. A phylogeny of the tropical genus Piper using ITS and the chloroplast intron psbJ-petA. Systematic Botany 33: 647–660. Kunth K. 1839. Die Familie des Piperaceen. Linnaea 13: 561–726. Lei LG, Wu ZY, Liang HX. 2002. Embryology of Zippelia begoniaefolia (Piperaceae) and its systematic relationships. Botanical Journal of the Linnean Society 140: 49–64. Miquel FA. 1843–1844. Systema piperacearum. Kramers, Rotterdam. Naumann J, Symmank L, Samain MS, Müller KF, Neinhuis C, dePamphilis CW, Wanke S. 2011. Chasing the hare — evaluating the phylogenetic utility of a nuclear single copy gene region at and below species level within the species-rich group Peperomia (Piperaceae). BMC Evolutionary Biology 11: 357. Risch S, McClure M, Vandermeer J, Waltz S. 1977. Mutualism between three species of tropical Piper (Piperaceae) and their ant inhabitants. American Midland Naturalist 98: 433–444.

FURTHER READING Samain MS, Wanke S, Mathieu G, Neinhuis C, Goetghebeur P. 2008. Verhuellia revisited — unravelling an intricate taxonomical history and a new subfamilial classification of Piperaceae. Taxon 57: 583–587. Samain MS, Vrijdaghs A, Hesse M, Goetghebeur P, Rodriguez FJ, Stoll A, Neinhuis C, Wanke S. 2010. Verhuellia is a segregate lineage in Piperaceae: more evidence from flower, fruit and pollen morphology, anatomy and development. Annals of Botany 105: 677–688. Smith JF, Tepe EJ, Stevens AC, Davidson C. 2008. Placing the origin of two species-rich genera in the late Cretaceous with later species divergence in the Tertiary: a phylogenetic, biogeographic and molecular dating analysis of Piper and Peperomia (Piperaceae). Plant Systematics and Evolution 275: 9–30. Symmank L, Samain MS, Smith JF, Pino G, Goetghebeur P, Neinhuis C, Wanke S. 2011. The extraordinary journey of Peperomia subgenus Tildenia: insights into diversification and colonisation patterns from its cradle in Peru to the Trans-Mexican Volcanic Belt. Journal of Biogeography 38: 2337–2349. Trelease W, Yunder TG. 1950. The Piperaceae of northern South America. University of Illinois Press, Urbana. Wanke S, Samain MS, Vanderschaeve L, Matthieu G, Goetghebeur P, Neinhuis C. 2006. Phylogeny of the genus Peperomia (Piperaceae) inferred from the trnK/matK region (cpDNA). Plant Biology 8: 93–102. Wanke S, Jaramillo MA, Borsch T, Samain MS, Quandt D, Neinhuis C. 2007. Evolution of Piperales — matK gene and trnK intron sequence data reveal lineage specific resolution contrast. Molecular Phylogenetics and Evolution 42: 477–497. Yuncker TG. 1958. The Piperaceae — a family profile. Brittonia 10: 1–7. 48. ARISTOLOCHIACEA BIRTHWORT FAMILY Bernadello G, Anderson GJ, Lopez S. P, Cleland MA, Stuessy TF, Crawford DJ. 1999. Reproductive biology of Lactoris fernandeziana (Lactoridaceae). American Journal of Botany 86: 829–840. Bolin JF, Maass E, Musselman LJ. 2009. Pollination biology of Hydnora africana Thunb. (Hydnoraceae) in Namibia: brood site mimicry with insect imprisonment. International Journal of Plant Sciences 170: 157–163. Bolin JF, Tennakoon KU, Maass E. 2010. Mineral nutrition and heterotrophy in the water conservative holoparasite Hydnora Thunb. (Hydnoraceae). Flora 205: 802–810. Carlquist S. 1993. Wood and bark anatomy of Aristolochiaceae: systematic and habital correlations. IAWA Journal 14: 341–357. Carlquist SJ. 1990. Wood anatomy and relationships of Lactoridaceae. American Journal of Botany 77: 1498–1505. Gamerro JC, Barreda V. 2008. New fossil record of Lactoridaceae in southern South America: a palaeobiogeographical approach. Botanical Journal of the Linnean Society 158: 41–50. González F, Rudall P. 2001. The questionable affinities of Lactoris: evidence from branching pattern, inflorescence morphology, and stipule development. American Journal of Botany 88: 2143–2150. González F. 1999. Inflorescence morphology and the systematics of Aristolochiaceae. Systematics and Geography of Plants 68: 159–172.

González FA, Stevenson DW. 2000 Perianth development and systematics of Aristolochia. Flora 195: 370–391. Huber H. 1985. Die Samenmerkmale und Gliederung der Aristolochiaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 107: 277–320. Kelly LM, González F. 2003. Phylogenetic relationships in Aristolochiaceae. Systematic Botany 28: 236–249. Kelly LM. 1998. Phylogenetic relationships in Asarum (Aristolochiaceae) based on morphology and ITS sequences. American Journal of Botany 84: 1752–1765. Lammers T, Stuessy TF, Silva M. 1986. Systematic relationships of the Lactoridaceae, an endemic family of the Juan Fernandez Islands, Chile. Plant Systematics and Evolution 152: 243–266. Leins P, Erbar C, Van Heel WA. 1988. Note on the floral development of Thottea (Aristolochiaceae) Blumea 33: 357–370. Macphail MK, Partridge AD, Truswell EM. 1999. Fossil pollen records of the problematical primitive angiosperm family Lactoridaceae in Australia. Plant Systematics and Evolution 214: 199–210. Musselman LJ, Visser JH. 1989. Taxonomy and natural history of Hydnora (Hydnoraceae). Aliso 12: 317–326. Nickrent DL, Blarer A, Qiu YL, Soltis DE, Soltis PS, Zanis M. 2002. Molecular data place Hydnoraceae with Aristolochiaceae. American Journal of Botany 89: 1809–1817. Oelschägel B, Wagner S, Salomo K, Pradeep NS, Yao TL, Isnard S, Rowe NP, Neinhuis S, Wanke S. 2011. Implications from molecular phylogenetic data for systematics, biogeography and growth from evolution of Thottea (Aristolochiaceae). Garden’s Bulletin Singapore 63: 259–275. Oelschlägel B, Gorb S, Wanke S, Neinhuis C. 2009. Structure and biomechanics of trapping flower trichomes and their role in pollination biology of Aristolochia plants (Aristolochiaceae). New Phytologist 184: 988–1002. Ohi-Toma T, Sugawara T, Murata H, Wanke S, Neinhuis C, Murata J. 2006. Molecular phylogeny of Aristolochia sensu lato (Aristolochiaceae) based on sequences of rbcL, matK, and phyA genes, with special reference to differentiation in chromosome numbers. Systematic Botany 31: 481–492. Sampson FB. 1995. Pollen morphology of Lactoridaceae — a re-examination. Grana 34: 100–107. Seymour RS, Maass E, Bolin JF. 2009. Floral thermogenesis of three species of Hydnora (Hydnoraceae) in Africa. Annals of Botany 104: 823–832. Stuessy TF, Crawford DJ, Anderson GJ, Jenner RJ. 1998. Systematics, biogeography and conservation of Lactoridaceae. Perspectives in Plant Evolution and Systematics 1–2: 267–290. Wagner ST. Isnard S, Rowe NP, Samain MS, Neinhuis C, Wanke S. 2012. Escaping the lianoid habit: evolution of shrub-like growth forms in Aristolochia subgenus Isotrema (Aristolochiaceae). American Journal of Botany 99: 1609–1629. Wanke S, González F, Neinhuis C. 2006. Systematics of pipevines: combining morphological and fastevolving molecular characters to investigate the relationships within subfamily Aristolochioideae (Aristolochiaceae). International Journal of Plant Sciences 167: 1215–1227. Zavada MS, Benson JM. 1987. First fossil evidence for the primitive angiosperm family Lactoridaceae. American Journal of Botany 74: 1590–1594.

49. MYRISTICACEAE NUTMEG FAMILY Armstrong JE, Drummond BA. 1986. Floral biology of Myristica fragrans Houtt. (Myristicaceae), the nutmeg of commerce. Biotropica 18: 32–38. Armstrong JE, Wilson TK. 1978. Floral morphology of Horsfieldia. American Journal of Botany 65: 441–449. Boureau E. 1950. Étude paléoxylologique du Sahara (IX). Sur un Myristicoxylon princeps n. gen. n. sp., du Danien d’Asselar. Bulletin du Muséum d’Histoire Naturelle. Paris II, 22: 523–528. Garrett GA. 1933. Systematic anatomy of the woods of the Myristicaceae. Tropical Woods 35: 6–48. Gottlieb OR. 1979. Chemical studies on medicinal Myristicaceae from Amazonia. Journal of Ethnopharmacology 1: 309–323. Howe HF, Van de Kerckhove GA. 1980. Nutmeg dispersal by tropical birds. Science 210: 925–927. Koster J, Baas P. 1981. Comparative leaf anatomy of the Asiatic Myristicaceae. Blumea 27: 115–173. Sauquet H. 2003. Androecium diversity and evolution in Myristicaceae (Magnoliales), with a description of a new Malagasy genus, Doyleanthus gen. nov. American Journal of Botany 90: 1293–1305. Sauquet H, Doyle JA, Scharaschkin T, Borsch T, Hilu KW, Chatrou LW, Le Thomas A. 2003. Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution. Botanical Journal of the Linnean Society 142: 125–186. Sauquet H, Le Thomas A. 2003. Pollen diversity and evolution in Myristicaceae (Magnoliales). International Journal of Plant Sciences 164: 613–628. Sauquet H. 2004. Systematic revision of Myristicaceae (Magnoliales) in Madagascar, with four new species of Mauloutchia. Botanical Journal of the Linnean Society 146: 351–368. Warburg O. 1897. Monographie der Myristicaceae. Nova Acta Academiae Caesareae LeopoldinoCarolinae Germanicae Naturae Curiosorum 68: 1–680. 50. MAGNOLIACEAE TULIPTREE FAMILY Agababian VS. 1972. Pollen morphology of Magnoliaceae. Grana 12: 166–176. Azuma H, García-Franco JG, Rico-Gray V, Thien LB. 2001. Molecular phylogeny of the Magnoliaceae: the biogeography of tropical and temperate disjunctions. American Journal of Botany 88: 2275–2285. Dilcher DL, Crane PR. 1984. Archaeanthus: an early angiosperm from the Cenomanian of the western interior North America. Annals of the Missouri Botanical Garden 71: 351–383. Doyle JA, Endress PK. 2010. Integrating Early Cretaceous fossils into the phylogeny of recent angiosperms: Magnoliidae and eudicots. Journal of Systematics and Evolution 48: 1–35. Gardiner J. 2000. Magnolias, a gardener’s guide. Timber Press, Portland. Gottsberger G, Siberbauer-Gottsberger I, Seymour RS, Dötterl S. 2012. Pollination ecology of Magnolia ovata may explain the overall larger flower size of the genus. Flora 207: 107–118. Keng H. 1978. The delimitation of the genus Magnolia. Garden’s Bulletin Singapore 31: 127–131. Leppik EE. 1975. Morphogenic stagnation in the evolution of Magnolia flowers. Phytomorphology 25: 451–464. Nie ZL, Wen J, Azuma H, Qiu YL, Sun H, Meng Y, Sun WB, Zimmer EA. 2008. Phylogeny and biogeographic complexity of Magnoliaceae in the

Plants of the World

683

FURTHER READING Northern Hemisphere inferred from three nuclear data sets. Molecular Phylogenetics and Evolution 48: 1027–1040. Qiu YL, Chase MW,, Parks CR. 1995. A chloroplast DNA phylogenetic study of the eastern Asia-east North America disjunct section Rhytidospermum of Magnolia (Magnoliaceae). American Journal of Botany 82: 1582–1588. Thien LB. 1974. Floral biology of Magnolia. American Journal of Botany 61: 1037–1045. Tiffney BH. 1977. Fruits and seeds of the Brandon lignite: Magnoliaceae. Botanical Journal of the Linnean Society 75: 299–323. Treseder NG. 1978. Magnolias. Faber, Faber, London. Xu FX, Rudall PJ. 2006. Comparative floral anatomy and ontogeny in Magnoliaceae. Plant Systematics and Evolution 258: 1–15. 51. DEGENERIACEAE MASIRATU FAMILY Bailey IW, Smith AC. 1942. Degeneriaceae, a new family of flowering plants from Fiji. Journal of the Arnold Arboretum 23: 356–365. Carlquist S. 1989. Wood and bark anatomy of Degeneria. Aliso 12: 485–495 Dahl AO, Rowley JR. 1965. Pollen of Degeneria vitiensis. Journal of the Arnold Arboretum 46: 308–323. Endress PK. 1984. The role of inner staminodes in the floral display of some relic Magnoliales. Plant Systematics and Evolution 146: 269–282. 52. HIMANTANDRACEAE PIGEONBERRY-ASH FAMILY Bailey IW, Nast CB, Smith AC. 1943. The family Himantandraceae. Journal of the Arnold Arboretum 24: 190–206. Buchheim G. 1962. Beobachtungen über den Bau der Frucht der Himantandraceae. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin N.F. 2: 78–92. Doweld AB, Shevyryova NA. 1997. Carpology, anatomy and taxonomic relationships of Galbulimima (Himantandraceae). Annals of Botany 81: 337–347. Prakash N, Foreman DB, Griffith SJ. 1984. Gametogenesis in Galbulimima belgraveana (Himantandraceae). Australian Journal of Botany 32: 605–612. 53. EUPOMATIACEAE BOLWARRA FAMILY Endress PK. 1984. The flowering process in the Eupomatiaceae (Magnoliales). Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 104: 297–319. Hotschkiss AT. 1959. Pollen and pollination in the Eupomatiaceae. Proceedings of the Linnean Society of New South Wales 83: 86–91. Mohana Rao PR. 1983. Seed and fruit anatomy in Eupomatia laurina with a discussion of the affinities of Eupomatiaceae. Flora 173: 311–319. Rix M, Endress PK. 2007. Eupomatia laurina (Eupomatiaceae). Curtis’s Botanical Magazine 24: 230–234. Taylor WC. 1985. Eupomatia alkaloids. Alkaloids 24: 1–23. 54. ANNONACEAE SOURSOP FAMILY Bobadilla M, Zavala F, Sisniegas M, Zavaleta G, Mostacero J, Taramona L. 2005. Evaluación larvicida de suspensiones acuosas de Annona muricata Linnaeus ‹‹guanábana›› sobre Aedes aegypti Linnaeus (Diptera, Culicidae). Revista Peruana de

684

Christenhusz, Fay & Chase

Biología 12: 145–152. Chaowasku T, Johnson DM, Van der Ham RWJM, Chatrou LW. 2012. Characterisation of Hubera (Annonaceae), a new genus segregated from Polyathia and allied to Miliusa. Phytotaxa 69: 33–56. Chatrou LW, Erkens RHJ, Richardson JE, Saunders RMK, Fay MF (eds) 2012. The natural history of Annonaceae. Botanical Journal of the Linnean Society 169: 1–279. Couvreur TLP, Van der Ham RWJM, Mbele YM, Mbago FM, Johnson DM. 2009. Molecular and morphological characterisation of a new monotypic genus of Annonaceae, Mwasumbia, from Tanzania. Systematic Botany 34: 26–276. Couvreur TLP, Pirie MD, Chatrou LW, Saundrs RMK, Su YCF, Richardson JE, Erkens RHJ. 2011. Early evolutionary history of the flowering plant family Annonaceae: steady diversification and boreotropical geodispersal. Journal of Biogeography 38: 664–680. Doyle JE, Le Thomas A. 1996. Phylogenetic analysis and character evolution in Annonaceae. Adansonia 18: 279–334. Hongratanaworakit T, Buchbauer G. 2004. Evaluation of the harmonizing effect of ylang-ylang oil on humans after inhalation. Planta Medica 70: 632–636. Rainer H. 2007. Monographic studies in the genus Annona L. (Annonaceae): inclusion of the genus Rollinia A.St.-Hil. Annalen des Naturhistorischen Museum in Wien, Serie B 108: 191–205. Saunders RMK. 2010. Floral evolution in the Annonaceae: hypotheses of homeotic mutations and functional convergence. Biological Reviews 85: 571–591. Xue B, Su YCF, Thomas DC, Saunders RMK. 2012. Pruning the polyphyletic genus Polyalthia (Annonaceae) and resurrecting the genus Monoon. Taxon 61: 1021–1039. Note: A complete bibliographic overview is provided by Erkens RHJ, Mennega EA, Westra LYT. 2012. Botanical Journal of the Linnean Society 169: 41–73. 55. CALYCANTHACEAE SPICEBUSH FAMILY Blake ST. 1972. Idiospermum (Idiospermaceae), a new genus and family for Calycanthus australiensis. Contributions from the Queensland Herbarium 12: 1–37. Carlquist S. 1983. Wood anatomy of the Calycanthaceae: ecological and systematic implications. Aliso 10: 427–441. Friis EM, Eklund H, Pedersen KR, Crane PR. 1994. Virginianthus calycanthoides gen. et sp. nov. — a calycanthaceous flower from the Potomac group (Early Cretaceous) of North America. International Journal of Plant Sciences 155: 772–285. Grant V. 1950. The pollination of Calycanthus occidentalis. American Journal of Botany 37: 294–297. Li J, Ledger J, Ward T, del Tredici P. 2004. Phylogenetics of Calycanthaceae based on molecular and morphological data with a special reference to divergent paralogues of the nrDNA ITS region. Harvard Papers in Botany 9: 69–82. Mohr BAR, Eklund H. 2003. Araripia florifera, a magnoliid angiosperm from the Lower Cretaceous Crato formation (Brazil). Review of Palaeobotany and Palynology 126: 279–292. Nicely KA. 1965. A monographic study of the Calycanthaceae. Castanea 30: 38–81. Staedler YM, Weston PH, Endress PK. 2007.

Floral phyllotaxis and f loral architecture in Calycanthaceae (Laurales). International Journal of Plant Sciences 168: 285–306. Staedler YM, Weston PH, Endress PK. 2009. Comparative gynoecium structure and development in Calycanthaceae (Laurales). International Journal of Plant Sciences 170: 21–41. 56. SIPARUNACEAE FEVERTREE FAMILY Bello MA, González F, Romero de Pérez G. 2002. Morfología del androeceo, tapete y ultrastructura del pollen de Siparuna aspera (Ruiz, Pavón) A.DC. (Siparunaceae). Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 26: 155–167. Heilborn O. 1931. Studies on the taxonomy, geographical distribution and embryology of the genus Siparuna Aubl. Svensk Botanisk Tidskrift 25: 202–228. Kimoto Y, Tobe H. 2003. Embryology of Siparunaceae (Laurales): characteristics and character evolution. Journal of Plant Research 116: 281–294. Renner SS, Won H. 2001. Repeated evolution of dioecy from monoecy in Siparunaceae (Laurales). Systematic Biology 50: 700–712. Renner SS, Hausner G. 2005. Siparunaceae. Flora Neotropica 95: 1–247. 57. GOMORTEGACEAE KEULE FAMILY Brizicky GK. 1959. Variability in the floral parts of Gomortega (Gomortegaceae). Willdenowia 2: 200–207. Doweld AB. 2001. Carpology and phermatology of Gomortega (Gomortegaceae): systematic and evolutionary implications. Acta Botanica Malacitana 26: 19–37. González M. 1998. Gomortega keule. IUCN Red List of threatened species, version 2013.1, available at www.iucnredlist.org. Heo K, Kinoto Y, Riveros M, Tobe H. 2004. Embryology of Gomortegaceae (Laurales): characteristics and character evolution. Journal of Plant Research 117: 221–228. Muñoz-Concha D, Davey MR. 2011. Gomortega keule, the neglected and endangered Chilean fruit tree. European Journal of Forest Research 130: 677–693. Muñoz-Concha D, Garrido-Werner A. 2011. Ethnobotany of Gomortega keule, an endemic and endangered Chilean tree. New Zealand Journal of Botany 49: 509–513. Muñoz-Concha D, Saúd G. 2011. Flowering and fruiting phenology of the endangered Chilean tree Gomortega keule. New Zealand Journal of Botany 49: 497–502. Stern WL. 1955. Xylem anatomy and relationships of Gomortegaceae. American Journal of Botany 42: 874–885. 58. ATHEROSPERMATACEAE SOUTHERNSASSAFRAS FAMILY: Endress PK. 1980. Ontogeny, function and evolution of extreme floral construction in Monimiaceae. Plant Systematics and Evolution 134: 79–120. Poole I, Gottwald H. 2001. Monimiaceae sensu lato, an element of Gondwanan polar forests: evidence from Late Cretaceous-Early Tertiary wood floras of Antarctica. Australian Systematic Botany 14: 207–230. Renner SS, Foreman DB, Murray D. 2000. Timing transantarctic disjunctions in the Atherospermataceae (Laurales): evidence from coding

FURTHER READING and noncoding chloroplast sequences. Systematic Biology 49: 579–591. Sampson FB. 1969. Floral morphology of Laurelia novae-zelandiae A.Cunn. (subfamily Atherospermatoideae). New Zealand Journal of Botany 7: 214–240. Stanstrup J, Schmidt JS, Rasmussen HB, Mølgaard P, Guzmán A, Staerk D. 2010. Bisbenzylisoquinoline alkaloids as markers of Atherospermataceae: tetrandrine and fangchinoline from Laureliopsis philippiana. Biochemistry, Systematics and Ecology 38: 450–453. Worth JRP. 2011. Low but structured chloroplast diversity in Atherosperma moschatum (Atherospermataceae) suggests bottlenecks in response to the Pleistocene glacials. Annals of Botany 108: 1247–1256. 59. HERNANDIACEAE LANTERN-TREE FAMILY: Chen JJ, Hung HC, Sung PJ, Chen IS, Kuo WL. 2011. Aporphine alkaloids and cytotoxic lignans from the roots of Illigera luzonensis. Phytochemistry 72: 523–532. Duyfjes BEE. 1994. Illigera elegans (Hernandiaceae), a new species from Christmas Island, Indian Ocean. Blumea 38: 407–408. Heo K, Tobe H. 1995. Embryology and relationships of Gyrocarpus and Hernandia (Hernandiaceae). Journal of Plant Research 108: 327–341. Kimoto Y, Tobe H. 2008. Embryology of Illigera and Sparattanthelium (Hernandiaceae, Laurales): family characteristics and relationships. International Journal of Plant Sciences 169: 391–408. Kubitzki K. 1969. Monographie der Hernandiaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 69: 78–209. Michalak I, Zhang LB, Renner SS. 2010. Trans-Atlantic, trans-Pacific and trans-Indian Ocean dispersal in the small Gondwanan Laurales family Hernandiaceae. Journal of Biogeography 37: 1214–1226. Shutts CF. 1960. Wood anatomy of Hernandiaceae and Gyrocarpaceae. Tropical Woods 113: 85–123. 60. MONIMIACEAE BOLDO FAMILY: Endress PK. 1980. Floral structure and relationships of Hortonia (Monimiaceae). Plant Systematics and Evolution 133: 199–221. Jordaan M, Lötter M. 2012. Taxonomic and nomenclatural notes on the monotypic genus Xymalos and general information on the family Monimiaceae. Bothalia 42: 51–56. Lorence DH. 1985. A monograph of the Monimiaceae (Laurales) in the Malagasy region (S.W. Indian Ocean). Annals of the Missouri Botanical Garden 72: 1–165. Philipson WR. 1987. A classification of the Monimiaceae. Nordic Journal of Botany 7: 25–29. Poole I, Gottwald H. 2001. Monimiaceae sensu lato, an element of Gondwanan polar forests: evidence from Late Cretaceous-Early Tertiary wood floras of Antarctica. Australian Systematic Botany 14: 207–230. Renner SS, Strijk JS, Strasberg D, Thébaud C. 2010. Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long-distance dispersal, but not West Gondwana. Journal of Biogeography 37: 1227–1238. Romanov MS, Endress PK, Bobrov AVFC, Melikian AP, Bejarano AP. 2007. Fruit structure and systematics of Monimiaceae s.s. (Laurales). Botanical Journal of the Linnean Society 153: 265–285.

61. LAURACEAE BAY-LAUREL FAMILY Buzgo M, Chanderbali AS, Kim S, Zheng Z, Oppenheimer DG, Soltis PS, Soltis DE. 2007. Floral developmental morphology of Persea americana (avocado, Lauraceae): the oddities of male organ identity. International Journal of plant Sciences 168: 261–284. Chanderbali AS, Van der Werff H, Renner SS. 2001. Phylogeny and historical biogeography of Lauraceae: evidence from the chloroplast and nuclear genomes. Annals of the Missouri Botanical Garden 88: 104–134. Ferguson DK. 1974. On the taxonomy of recent and fossil species of Laurus (Lauraceae). Botanical Journal of the Linnean Society 68: 51–72. Fijridiyanto IA, Murakami N. 2009. Phylogeny of Litsea and related genera (Laureae-Lauraceae) based on analysis of rpb2 gene sequences. Journal of Plant Research 122: 283–298. Kimoto Y, Utami N, Tobe H. 2006. Embryology of Eusideroxylon (Cryptocaryeae, Lauraceae) and character evolution in the family. Botanical Journal of the Linnean Society 150: 187–201. Kvaček Z. 1971. Fossil Lauraceae in the stratigraphy of the North-Bohemian Tertiary. Sbornik Geologických vĕd Paleontologie 13: 47–86. Poole I, Richter HG, Francis JE. 2000. Evidence for Gondwana origins for Sassafras (Lauraceae)? Late Cretaceous fossil wood of Antarctica. IAWA Journal 21: 463–475. Rohwer JG. 1986. Prodromus einer Monographie der Gattung Ocotea Aubl. (Lauraceae), sensu lato. Mitteilungen aus dem Institut für allgemeine Botanik in Hamburg 20: 3–278. Rohwer JG, Richter HG, Van der Werff H. 1991. Two new genera of Neotropical Lauraceae, and critical remarks on the generic delimitation. Annals of the Missouri Botanical Garden 78: 338–400. Rohwer JG. 2000. Toward a phylogenetic classification of the Lauraceae: evidence from matK sequences. Systematic Botany 25: 60–71. Rohwer JG, Rudolph B. 2005. Jumping genera: the phylogenetic positions of Cassytha, Hypodaphnis, and Neocinnamomum (Lauraceae) based on different analysis of trnK intron sequences. Annals of the Missouri Botanical Garden 92: 153–178. Rohwer JG. 2009. The timing of nectar secretion in staminal and staminodial glands in Lauraceae. Plant Biology 11: 490–492. Snow DW. 1981. Tropical frugivorous birds and their food plants: a world survey. Biotropica 13: 1–14. Van der Merwe JJM, Van Wyk, AE, Kok PDF. 1990. Pollen types in the Lauraceae. Grana 29: 185–196. Van Rijckevorsel P. 2002. Wood anatomy and taxonomy of the Lauraceae family. World of Woods 55(8): 4–16. Weber JZ. 1981. A taxonomic revision of Cassytha (Lauraceae) in Australia. Journal of the Adelaide Botanical Garden 3: 187–262. Note: For older literature see the bibliography of Kostermans AJGH. 1964. Bibliographia lauracearum. Archipel, Bogor. 62. CHLORANTHACEAE PEARL-ORCHID FAMILY Antonelli A, Sanmartín I. 2011. Mass extinction, gradual cooling, or rapid radiation? Reconstructing the spatiotemporal evolution of the ancient angiosperm genus Hedyosmum (Chloranthaceae) using empirical and simulated approaches. Systematic Biology 60: 596–615. Carlquist S. 1987. Presence of vessels in Sarcandra (Chloranthaceae): comments on vessel origins

in angiosperms. American Journal of Botany 74: 1765–1771. Carlquist S. 1992. Wood anatomy and stem of Chloranthus; summary of wood anatomy of Chloranthaceae, with comments on relationships, vessellessness, and the origin of monocotyledons. IAWA Bulletin 13: 3–16. Doria MG, Pabón-Mora N, González F. 2012. Reassessing inflorescence and floral morphology and development in Hedyosmum (Chloranthaceae). International Journal of Plant Sciences 173: 735–750. Doyle JA, Eklund H, Herendeen PS. 2003. Floral evolution in Chloranthaceae: implications of a morphological phylogenetic analysis. International Journal of Plant Sciences 164(5 suppl.): S365–S382. Eklund H, Friis EM, Pedersen KR. 1997. Chloranthaceous floral structures from the Late Cretaceous of Sweden. Plant Systematics and Evolution 207: 13–42. Eklund H, Doyle JA, Herendeen PS. 2004. Morphological phylogenetic analysis of living and fossil Chloranthaceae. International Journal of Plant Sciences 165: 107–151. Endress PK. 1987. The Chloranthaceae: reproductive structures and phylogenetic position. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 109: 153–226. Friis EM, Crane PR, Pederson KR. 1986. Floral evidence for Cretaceous chloranthoid angiosperms. Nature 320: 163–164. Leroy JP. 1983. The origin of angiosperms: an unrecognized ancestral dicotyledon, Hedyosmum (Chloranthales), with a strobiloid flower is living today. Taxon 32: 169–175. Swamy BGL. 1953. The morphology and relationships of the Chloranthaceae. Journal of the Arnold Arboretum 34: 375–411. Todzia CA. 1988. Chloranthaceae. Flora Neotropica Monograph 48: 1–139. Verdcourt B. 1986. Chloranthaceae. Flora Malesiana I, 10: 123–144. Zhang LB, Renner SS. 2003. The deepest splits in Chloranthaceae as resolved by chloroplast sequences. International Journal of Plant Sciences 164(5, suppl.): S383–S392. Zhang Q, Antonelli A, Feild TS, Kong HZ. 2011. Revisiting taxonomy, morphological evolution, and fossil calibration strategies in Chloranthaceae. Journal of Systematics and Evolution 49: 315–329. 63. ACORACEAE SWEET-FLAG FAMILY Bogner J. 2011. Acoraceae. Flora Malesiana ser. 1, 20: 1–13. Carlquist S. 2012a. Monocot xylem revisited: new information. Botanical Review 78: 87–150. Carlquist S, Schneider EL. 1997. Origins and nature of vessels in monocotyledons. I. Acorus. International Journal of Plant Sciences 158: 51–56. Duvall MR, Learn GH, Eguiarte LE, Clegg MT. 1993. Phylogenetic analysis of rbcL sequences identifies Acorus calamus as the primal extant monocotyledon. Proceedings of the National Academy of Sciences of the USA 90: 4641–4644. Grayum MH. 1987. A summary of evidence and arguments supporting the removal of Acorus from the Araceae. Taxon 36: 723–729. Kaplan DR. 1970. Comparative foliar histogenesis of Acorus calamus and its bearing on the phyllode theory of monocotyledonous leaves. American Journal of Botany 57: 331–336.

Plants of the World

685

FURTHER READING Rana TS, Mahar KS, Pandey MM, Srivastava SK, Rawat AKS. 2013. Molecular and chemical profiling of ‘sweet flag’ (Acorus calamus L.) germplasm from India. Physiology and Molecular Biology of Plants 19: 231–237. Todorova MN, Ognyanov IV, Shatar S. 1995. Chemical composition of essential oil from Mongolian Acorus calamus L. rhizomes. Journal of Essential Oil Research 7: 191–193. 64. ARACEAE CALLA-LILY FAMILY Arber A. 1919. On the vegetative morphology of Pistia and the Lemnaceae. Proceedings of the Royal Society 91: 96–103. Bierzychudek P. 1982. The demography of jack-in-thepulpit, a forest perennial that changes sex. Ecology Monographs 52: 335–351 Bogner J. 1979. A critical list of the aroid genera. Aroideana 1: 63–73. Bogner J, Nicolson DH. 1991. A revised classification of Araceae with dichotomous keys. Willdenowia 21: 35–50. Boyce PC. 1993. The genus Arum. Royal Botanic Gardens, Kew. Cabrera L., Salazar GA, Chase MW, Mayo SJ, Bogner J, Dávila P. 2008. Phylogenetic relationships of aroids and duckweeds (Araceae) inferred from coding and noncoding plastid DNA. American Journal of Botany 95: 1153–1165 Chandra S. 1984. Edible aroids. Clarendon Press, Oxford. Croat TB. 1980. Flowering behavior of the Neotropical genus Anthurium (Araceae). American Journal of Botany 67: 888–904. Culley DD, Rejmankova E, Kvet J, Frye JB. 1981. Production, chemical quality and use of duckweeds (Lemnaceae) in aquaculture, waste management and animal feeds. Journal of the World Mariculture Society 12: 27–49. Cusimano N, Bogner J, Mayo SJ, Boyce PC, Wong SY, Hesse M, Hetterscheid WLA, Keating RC, French JC. 2011. Relationships within the Araceae: comparison of morphological patterns with molecular phylogenies. American Journal of Botany 98: 654–668. Den Hartog C, Van der Plas F. 1970. A synopsis of the Lemnaceae. Blumea 18: 355–368. Dötterl S, David A, Boland W, Silberbauer-Gottsberger I, Gottsberger G. 2012. Evidence for behavioral attractiveness of methoxylated aromatics in a dynastid scarab beetle-pollinated Araceae. Journal of Chemical Ecology 38: 1539–1543. eMonocot Team. CATE Araceae website. http:// araceae.e-monocot.org (accessed 2 January 2014). Ertl PO. 1932. Vergleichende Untersuchungen über die Entwicklung der Blattnervatur der Araceen. Flora 126: 115–248. Espíndola A, Buerki S, Bedalova M, Küpfer P, Alvarez N. 2010. New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history. Botanical Journal of the Linnean Society 163: 14–32. Eyde RH. Nicolson DH. Sherwin P. 1967. A survey of floral anatomy in Araceae. American Journal of Botany 54: 478–497. Grayum MH. 1990. Evolution and phylogeny of the Araceae. Annals of the Missouri Botanical Garden 43: 1–167. Hillman WS. 1961. The Lemnaceae or duckweeds. A review of the descriptive and experimental literature. Botanical Review 27: 221–287. Hillman WS, Culley DD. 1978. The uses of duckweeds.

686

Christenhusz, Fay & Chase

American Scientist. 66: 442–451. Hotta M. 1970. Study of the family Araceae — general remarks. Japanese Journal of Botany 20: 269–310. Hugett B, Tomlinson PB. 2010. Aspects of vessel dimensions in the aerial roots of epiphytic Araceae. International Journal of Plant Sciences 171: 362–369. Landolt E. 1986–1987. The family Lemnaceae — a monographic study Vol. 1 and 2. Veröffentlichungen des Geobotanisches Institutes an der Eidggenössischen Technischen Hochschule Stiftung Rübel Zürich 71: 1–566; 95: 1–638. Les DH, Crawford DJ, Landolt E, Gabel JD, Kimball RT. 2002. Phylogeny and systematics of Lemnaceae, the duckweed family. Systematic Botany 27: 221–240. Mayo SJ, Bogner J, Boyce PC. 1997. The genera of Araceae. Royal Botanic Gardens, Kew. Meeuse BDJ, Rashin I. 1988. Review. Sexual reproduction in the arum lily family, with emphasis on thermogenicity. Sexual Plant Reproduction 1: 3–15. Nauheimer K, Boyce PC, Renner SS. 2012. Giant taro and its relatives: a phylogeny of the large genus Alocasia (Araceae) sheds light on Miocene floristic exchange in the Malesian region. Molecular Phylogenetics and Evolution 63: 43–51. Nauheimer K, Metzler D, Renner SS. 2012. Global history of the ancient monocot family Araceae inferred with models accounting for past continental positions and previous ranges based on fossils. New Phytologist 195: 938–950. Nicolson DH. 1987. History of Araceae systematics. Aroideana 10: 23–30. Petersen G. 1989. Cytology and systematics of Araceae. Nordic Journal of Botany 9: 119–166. Plowman T. 1969. Folk uses of New World aroids. Economic Botany 23: 97–122. Stockey RA, Hoffman GL, Rothwell GW, 1997. The fossil monocot Limnobiophyllum scutatum: resolving the phylogeny of Lemnaceae. American Journal of Botany 84: 355–368. Ulrich S, Hesse M, Bröderbauer D, Bogner J, Weber M, Halbritter H. 2013. Calla palustris (Araceae): new palynological insights with special regard to its controversial systematic position and to closely related genera. Taxon 62: 701–712. 65. TOFIELDIACEAE FALSE-ASPHODEL FAMILY Azuma H, Tobe H. 2011. Molecular phylogenetic analysis of Tofieldiaceae (Alismatales): family circumscription and intergeneric relationships. Journal of Plant Research 124: 349–357. Campbell LM, Dorr, LJ. 2013. A synopsis of Harperocallis (Tofieldiaceae, Alismatales) with ten new combinations. Phytokeys 21: 37–52. McDaniel ST. 1968. Harperocallis, a new genus of the Liliaceae from Florida. Journal of the Arnold Arboretum 49: 35–40. Remizowa MV, Sokoloff DD. 2003. Inflorescence and floral morphology in Tofieldia (Tofieldiaceae) compared with Araceae, Acoraceae and Alismatales s. str. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzen-geographie 124: 255–271. Remizowa MV, Sokoloff DD, Campbell LM, Stevenson DW, Rudall PJ. 2011. Harperocallis is congeneric with Isidrogalvia (Tofieldiaceae, Alismatales): evidence from comparative floral morphology. Taxon 60: 1076–1094. Sterling C. 1979. Comparative morphology of the

carpel in the Liliaceae: Tofieldieae. Botanical Journal of the Linnean Society 79: 321–332. Tamura MN, Fuse S, Azuma H, Hasebe M. 2004. Biosystematic studies in the family Tofieldiaceae I. Phylogeny and circumscription of the family inferred from DNA sequences of matK and rbcL. Plant Biology 6: 562–567. Utech FH. 1978. Floral vascular anatomy of Pleea tenuifolia Michx. (Liliaceae-Tofieldieae) and its reassignment to Tofieldia. Annals of the Carnegie Museum 47: 423–453. Zomlefer WB. 1997. The genera of Tofieldiaceae in the southeastern United States. Harvard Papers in Botany 2: 179–194. 66. ALISMATACEAE WATER-PLANTAIN FAMILY Argue CL. 1976. Pollen studies in the Alismataceae with special reference to taxonomy. Pollen and Spores 18: 157–201. Chen L-Y, Chen J-M, Gituru RW, Temam TD, Wang Q-F. 2012. Generic phylogeny and historical biogeography of Alismataceae, inferred from multiple DNA sequences. Molecular Phylogenetics and Evolution 63: 407–416. Haynes RR. Holm-Nielsen LB. 1985. A generic treatment of Alismatidae in the Neotropics with special reference to Brazil. Acta Amazonica 15: 153–193. Lehtonen S. 2006. Phylogenetics of Echinodorus (Alismataceae) based on morphological data. Botanical Journal of the Linnean Society 150: 291–305. Lehtonen S. 2009. Systematics of the Alismataceae — a morphological evaluation. Aquatic Botany 91: 279–290. Lehtonen S. 2009. On the origin of Echinodorus grandiflorus (Alismataceae) in Florida (“E. floridanus”), and its estimated potential as an invasive species. Hydrobiologia 635: 107–112. Lehtonen S, Myllys L. 2008. Cladistic analysis of Echinodorus (Alismataceae): simultaneous analysis of molecular and morphological data. Cladistics 24: 218–239. Les DH, Garvin DK, Wimpee CF. 1993. Phylogenetic studies in the monocot subclass Alismatidae: evidence for a reappraisal of the aquatic order Najadales. Molecular Phylogenetics and Evolution 2: 304–314. Stant MY. 1964. Anatomy of the Alismataceae. Journal of the Linnean Society, Botany 59: 1–42. 67. BUTOMACEAE FLOWERING-RUSH FAMILY Brown JS, Eckert CG. 2005. Evolutionary increase in sexual and clonal reproductive capacity during biological invasion in an aquatic plant Butomus umbellatus (Butomaceae). American Journal of Botany 92: 495–502. Singh V, Sattler R. 1974. Floral development of Butomus umbellatus. Canadian Journal of Botany 52: 223–230. Stant MY. 1967. Anatomy of the Butomaceae. Journal of the Linnean Society, Botany 60: 31–60. 68. HYDROCHARITACEAE FROGBIT FAMILY Ancibor E. 1979. Systematic anatomy of vegetative organs of the Hydrocharitaceae. Botanical Journal of the Linnean Society 78: 237–266. Appert O. 1996. Zur Verbreitung und Biologie des Gattung Appertiella (Hydrocharitaceae). Botanica Helvetica 106: 57–71. Chen L-Y, Chen J-M, Gituru RW, Wang Q-F. 2012. Genetic phylogeny, historical biogeography and character evolution of the cosmopolitan aquatic

FURTHER READING plant family Hydrocharitaceae. BMC Evolutionary Biology 12: 30. Cook CDK. 1982–1985. [a series of revisions of most genera of Hydrocharitaceae]. Aquatic Botany 13: 485–504, 505–513; 14: 177–204; 15: 1–52; 16: 213–249; 17: 1–27; 18: 263–274; 19: 73–96; 20: 131–177; 21: 111–156, 157–164. Cox PA. 1985. Noodles on the tide: why is water-borne pollen so long and stringy? Natural History 94: 36–41. Cox PA, Tomlinson PB. 1988. Pollination ecology of a seagrass, Thalassia testudinum (Hydrocharitaceae), in St. Croix. American Journal of Botany 75: 958–965. He J-B, Sun X-Z, Zhong Y, Huang D. 1991. Cladistic studies on the genus Ottelia (Hydrocharitaceae). Journal of Wuhan Botanical Research 9: 121–129. Kaul RB. 1970. Evolution and adaptation of inflorescences in the Hydrocharitaceae. American Journal of Botany 57: 708–715. Les DH, Garvin DK, Wimpee CF. 1993. Phylogenetic studies in the monocot subclass Alismatidae: evidence for a reappraisal of the aquatic order Najadales. Molecular Phylogenetics and Evolution 2: 304–314. Les DH, Cleland MA, Waycott M. 1997. Phylogenetic studies in Alismatidae II: evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22: 443–463. Les DH, Jacobs SWL, Tipepry NP, Chen L, Moody ML, Wilstermann-Hildebrand M. 2008. Systematics of Vallisneria (Hydrocharitaceae). Systematic Botany 33: 49–65. Miki S. 1937. The origin of Najas and Potamogeton. Botanical Magazine (Tokyo) 51: 472–480. Pettitt JM, Jermy AC. 1975. Pollen in hydrophilous angiosperms. Micron 5: 377–405. Shaffer-Fehre M. 1991. The position of Najas within subclade Alismatidae (Monocotyledones) in the light of new evidence from seed coat structures in the Hydrocharitoideae (Hydrocharitales). Botanical Journal of the Linnean Society 107: 189–209. Tanaka N, Setoguchi H, Murata J. 1997. Phylogeny of the family Hydrocharitaceae inferred from rbcL and matK gene sequence data. Journal of Plant Research 110: 329–337. Wilder GJ. 1975. Phylogenetic trends in the Alismatidae (Monocotyledonae). Botanical Gazette 136: 159–170. 69. SCHEUCHZERIACEAE RANNOCH-RUSH FAMILY Buchenau F. 1903. Scheuchzeriaceae. Das Pflanzenreich IV. 14: 1–20. Engelmann, Leipzig. Stenar H. 1935. Embryologische Beobachtungen über Scheuchzeria palustris L. Botaniska Notiser 1935: 78–86. Tallis JH, Birks HJB. 1965. The past and present distribution of Scheuchzeria palustris L. in Europe. Journal of Ecology 53: 287–298. 70. APONOGETONACEAE WATERBLOMMETJIE FAMILY Grimsson F, Zetter R, Halbritter H, Grimm GW. 2013. Aponogeton pollen from the Cretaceous and Paleogene of North America and west Greenland: implications for the origin and palaeobiogeography of the genus. Review of Palaeobotany and Palynology 200: 161–187. Les DH, Moody ML, Jacobs SWL. 2005. Phylogeny and systematics of Aponogeton (Aponogetonaceae):

the Australian species. Systematic Botany 30: 503–519. Van Bruggen HWE. 1985. Monograph of the genus Aponogeton (Aponogetonaceae) Bibliotheca Botanica 137: 1–76. Van Bruggen HWE. 1991. Neue Erkentnisse über die Aponogetonaceae. Aqua-Planta 16: 46–54. Wright H, Van Doorn WG, Gunawardena AHLAN. 2009. In vivo study of developmental programmed cell death using the lace plant (Aponogeton madagascariensis; Aponogetonaceae) leaf model system. American Journal of Botany 96: 865–876. 72. MAUNDIACEAE MAUND’S-ARROWGRASS FAMILY Sokoloff DD, Von Mering S, Jacobs SWL, Remizowa MV. 2013. Morphology of Maundia supports its isolated phylogenetic position in the early-divergent monocot order Alismatales. Botanical Journal of the Linnean Society 173: 12–45. 71. JUNCAGINACEAE ARROWGRASS FAMILY Larsen K. 1966. Cytotaxonomical note on Lilaea. Botaniska Notiser 119: 496–497. Von Mering S, Kadereit JW. 2010. Phylogeny, systematics and recircumscription of Juncaginaceae — a cosmopolitan wetland family, pp. 55–79. In: Seberg O, et al. (eds). Diversity, phylogeny and evolution in the monocotyledons. Aarhus University Press, Århus. 73. ZOSTERACEAE EELGRASS FAMILY Cambridge ML, Carstairs SA, Kuo J. 1983. An unusual method of vegetative propagation in Australian Zosteraceae. Aquatic Botany 15: 201–203. Cooper LW, McRoy CP. 1988. Anatomical adaptation to rocky substrates and surf exposure by the seagrass genus Phyllospadix. Aquatic Botany 32: 365–381. Coyer JA, Hoarau G, Kuo J, Tronholm A, Veldink J, Olsen JL. 2013. Phylogeny and temporal divergence of the seagrass family Zosteraceae using one nuclear and three chloroplast loci. Systematics and Biodiversity 11: 271–284. den Hartog C. 1970. The seagrasses of the world. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde II, 59: 1–275. Kato Y, Aioi K, Omori Y, Takahata N, Satta Y. 2003. Phylogenetic analysis of Zostera species based on rbcL and matK sequences: implications for the origin and diversification of seagrasses in Japanese waters. Genes and Genetic Systems 78: 329–342. Kuo J, Seto K, Nasu T, Iizumi H, Aioi K. 1989. Notes on Archaeozostera in relation to Zosteraceae. Aquatic Botany 34: 317–328. Les DH, Cleland MA, Waycott M. 1997. Phylogenetic studies in Alismatidae, II: Evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22: 443–463. Reusch TBH, Boström C, Stam WT, Olsen JL. 1999. An ancient eelgrass clone in the Baltic. Marine Ecology Progress Series 183: 301–304. 74. POTAMOGETONACEAE PONDWEED FAMILY Charlton WA, Posluszny U. 1991. Meristic variation in Potamogeton flowers. Botanical Journal of the Linnean Society 106: 265–293. Daghlian CP. 1981. A review of the fossil record of monocotyledons. Botanical Review (Lancaster) 47: 517–555.

Guo YH, Cook CDK. 1990. The floral biology of Groenlandia densa (L.) Fourreau (Potamogetonaceae). Aquatic Botany 38: 283–288. Haynes RR. 1978. The Potamogetonaceae in the southeastern United States. Journal of the Arnold Arboretum 59: 170–191. Les DH. 1983. Taxonomic implications of aneuploidy and poly ploidy in Potamogeton (Potamogetonaceae). Rhodora 85: 301–323. Les DH, Sheridan DJ. 1990. Biochemical heterophylly and flavonoid evolution in North American Potamogeton (Potamogetonaceae). American Journal of Botany 77: 453–465. Les DH, Cleland MA, Waycott M. 1997. Phylogenetic studies in Alismatidae, II: evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22: 443–463. Lindqvist C, de Laet J, Haynes RR, Aagesen L, Keener BR, Albert VA. 2006. Molecular phylogenetics of an aquatic plant lineage, Potamogetonaceae. Cladistics 22: 568–588. Miki S. 1937. The origin of Najas and Potamogeton. Botanical Magazine (Tokyo) 51: 472–480. Muencher WC. 1936. The germination of seeds of Potamogeton. Annals of Botany 50: 805–821. Posluszny U. 1981. Unicarpellate floral development in Potamogeton zosteriformis. Canadian Journal of Botany 59: 495–504. Sorsa P. 1988. Pollen morphology of Potamogeton and Groenlandia (Potamogetonaceae) and its taxonomic significance. Annales Botanici Fennici 25: 179–199. Wieglieb G. 1988. Notes on pondweeds, outline for a monographical treatment of the genus Potamogeton. Feddes Repertorium 99: 249–266. 75. POSIDONIACEAE TAPEWEED FAMILY Arnaud-Haond S, Duarte CM, Diaz-Almela E, Marbà N, Sintes T, Serrão EA. 2012. Implications of extreme life span in clonal organisms: millenary clones in meadows of the threatened seagrass Posidonia oceanica. PLoS ONE 7: e30454. Cannon JFM. 1979. An experimental investigation of Posidonia balls. Aquatic Botany 6: 407–410. Den Hartog C. 1970. The seagrasses of the world. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde II, 59: 1–275. Kuo J. 1983. Notes on the biology of Australian seagrasses. Proceedings of the Linnean Society of New South Wales 106: 225–245. Ma G, Zhang X, Bunn E, Dixon K. 2012. Megasporogenesis and embryogenesis in three sympatric Posidonia seagrass species. Aquatic Biology 100: 1–7. McConchie CA, Knox RB. 1989. Pollen-stigma interaction in the seagrass Posidonia australis. Annals of Botany 63: 235–248. Remizowa M, Sokoloff DD, Calvo S, Tomasello A, Rudall PJ. 2012. Flowers and inflorescences of the seagrass Posidonia (Posidoniaceae, Alismatales). American Journal of Botany 99: 1592–1602. 76. RUPPIACEAE TASSELWEED FAMILY Friis EM. 1985. Angiosperm fruits and seeds from the Middle Miocene of Jutland (Denmark). Kongelige Danske Videnskabernes Selskabs Biologiska Skrifter 24: 1–165. Ito Y, Ohi-Toma T, Murata J, Tanaka N. 2010. Hybridisation and polyploidy of an aquatic plant, Ruppia (Ruppiaceae), inferred from plastid and nuclear DNA phylogenies. American Journal of Botany 97: 1156–1167.

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FURTHER READING Jacobs SWL, Brock MA. 1982. A revision of the genus Ruppia (Potamogetonaceae) in Australia. Aquatic Botany 14: 325–337. Les DH, Cleland MA, Waycott M. 1997. Phylogenetic studies in Alismatidae, II: evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22: 443–463. 77. CYMODOCEACEAE TURTLE-GRASS FAMILY Cox PA, Humphries CJ. 1993. Hydrophilous pollination and breeding system evolution in seagrasses: a phylogenetic approach to the evolutionary ecology of Cymodoceaceae. Botanical Journal of the Linnean Society 113: 217–226. Den Hartog C. 1970. The seagrasses of the world. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde II, 59: 1–275. Ducker SC, Foord NJ, Knox RB. 1977. Biology of Australian seagrasses: the genus Amphibolis C.Agardh (Cymodoceaceae). Australian Journal of Botany 25: 67–95. Kuo J. 2013. Chromosome numbers of Australian Cymodoceaceae. Plant Systematics and Evolution 299: 1443–1448. Kuo J, Kirkman H. 1990. Anatomy of viviparous seagrass seedlings of Amphibolis and Thalassodendron and their nutrient supply. Botanica Marina 33: 117–126. McConchie CA, Ducker SC, Knox RB. 1982. Biology of Australian seagrasses: floral development and morphology in Amphibolis (Cymodoceaceae). Australian Journal of Botany 30: 251–264. McMillan C, Bragg LH. 1987. Comparison of fruits of Syringodium (Cymodoceaceae) from Texas, the US Virgin islands and the Philippines. Aquatic Botany 28: 97–100. Petersen G, Seberg O, Short FT, Fortes MD. 2014. Complete genomic congruence but non-monophyly of Cymodocea (Cymodoceaceae). Taxon 63: 3–8. Pettitt JM, McConchie CA, Ducker SC, Knox RB. 1980. Unique adaptations for submarine pollination in seagrasses. Nature 286: 487–489. Waycott M, Walker DI, James SH. 1996. Genetic uniformity in a dioecious seagrass Amphibolis antarctica. Heredity 76: 578–585. 78. PETROSAVIACEAE OZE-SO FAMILY Cameron KM, Chase MW, Rudall PJ. 2003. Recircumscription of the monocotyledonous family Petrosaviaceae to include Japonolirion. Brittonia 55: 214–225. Logacheva MD, Schelkunov MI, Nuraliev MS, Samigullin TH, Penin AA. 2014. The plastid genome of mycoheterotrophic monocot Petrosavia stellaris exhibits both gene losses and multiple rearrangements. Genome Biology and Evolution 6: 238–246. Remizowa M. 2011. Floral morphology in Japonolirion and Petrosavia (Petrosaviales). Botanicheskii Zhurnal 96: 198–214. Stant MY. 1970. Anatomy of Petrosavia stellaris Becc., a saprophytic monocotyledon. Botanical Journal of the Linnean Society 63 (Suppl. 1): 147–161. Takahashi H, Nishio E, Hayashi H. 1993. Pollination biology of the saprophytic species Petrosavia sakuraii (Makino) van Steenis in central Japan. Journal of Plant Research 106: 213–217. Tobe H. 2008. Embryology of Japonolirion (Petrosaviaceae, Petrosaviales): a comparison

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Christenhusz, Fay & Chase

with other monocots. Journal of Plant Research 121: 407–416. Utech FH. 1984. Floral vascular anatomy of Japonolirion osense Nakai (Liliaceae) and its tribal relationship. Annals of the Carnegie Museum 53: 447–461. 79. NARTHECIACEAE BOG-ASPHODEL FAMILY Fuse S, Lee NS, Tamura MN. 2012. Biosystematic studies on the family Nartheciaceae (Dioscoreales) I. Phylogenetic relationships, character evolution and taxonomic re-examination. Plant Systematics and Evolution 298: 1575–1584. Hara H. 1967. The status of the genus Metanarthecium Maxim. Journal of Japanese Botany 42: 312–316. Merckx V, Schols V, Geuten K, Huysmans S, Smets EF. 2008. Phylogenetic relationships in Nartheciaceae (Dioscoreales) with focus on pollen and orbicule morphology. Belgian Journal of Botany 141: 64–77. Remizowa M, Sokoloff D, Rudall PJ. 2006. Evolution of the monocot gynoecium: evidence from comparative morphology and development in Tofieldia, Japonolirion, Petrosavia and Narthecium. Plant Systematics and Evolution 258: 183–209. Remizowa M, Sokoloff DD, Kondo K. 2008. Floral evolution in the monocot family Nartheciaceae (Dioscoreales): evidence from anatomy and development of Metanarthecium luteo-viride Maxim. Botanical Journal of the Linnean Society 158: 1–18. Zomlefer WB. 1997. The genera of Nartheciaceae in the southeastern United States. Harvard Papers in Botany 2: 195–211. 80. BURMANNIACEAE BLUETHREADS FAMILY Jonker FP. 1938. A monograph of the Burmanniaceae. Mededelingen van het Botanisch Museum en Herbarium van de Rijksuniversiteit te Utrecht 51: 1–279. Maas PJM, Maas-van de Kamer H, Van Benthem J, Snelders HCM, Rübsamen T. 1986. Burmanniaceae. Flora Neotropica Monographs 42: 1–189. Merckx VSFT, Schols V, Maas-van de Kamer H, Maas P, Huysmans S, Smets EF. 2006. Phylogeny and evolution of Burmanniaceae (Dioscoreales) based on nuclear and mitochondrial data. American Journal of Botany 93: 1684–1698. Merckx VSFT, Chatrou LW, Lemaire B, Sainge MN, Huysmans S, Smets EF. 2008. Diversification of mycoheterotrophic angiosperms: evidence from Burmanniaceae. BMC Evolutionary Biology 8: 178. Rao VS. 1970. Certain salient features in the floral anatomy of Burmannia, Gymnosiphon and Thismia. Journal of the Indian Botanical Society 48: 22–29. 81. DIOSCOREACEAE YAM FAMILY Ayensu ES. 1966. Taxonomic status of Trichopus: anatomical evidence. Journal of the Linnean Society, Botany 59: 425–430. Bucherer E. 1889. Beiträge zur Morphologie und Anatomie der Dioscoreaceen. Bibliotheca Botanica Band 3, Heft 16: 1–34. Burkill IH. 1960. The organography and the evolution of Dioscoreaceae, the family of the yams. Journal of the Linnean Society, Botany 56: 319–412. Caddick LR, Rudall PJ, Wilkin P, Chase MW. 2000. Yams and their allies: systematics of Dioscoreales. Pp. 475–487, in Wilson, K. L. and D. A. Morrison, (eds) Monocots: systematics and evolution. CSIRO Publishing, Collingwood.

Caddick LR, Rudall PJ, Wilkin P, Hedderson TAJ, Chase MW. 2002. Phylogenetics of Dioscoreales based on combined analysis of morphological and molecular data. Botanical Journal of the Linnean Society 138: 123–144. Caddick LR, Wilkin P, Rudall PJ, Hedderson TAJ, Chase MW. 2002. Yams reclassified: a recircumscription of Dioscoreaceae and Dioscoreales. Taxon 51: 103–114. Drenth E. 1972. A revision of the family Taccaceae. Blumea 20: 367–406. Schols P, Furness CA, Merckx V, Wilkin P, Smets E. 2005. Comparative pollen development in Dioscoreales. International Journal of Plant Sciences 166: 909–924. Viruel J, Catalán P, Segarra-Moragues JG. 2010. New microsatellite loci in the dwarf yams Dioscorea group Epipetrum (Dioscoreaceae). American Journal of Botany 97: e121–e123. Wilkin P, Schols P, Chase MW, Chayamarit K, Furness CA, Huysmans S, Rakotonasolo F, Smets E, Thapyai C. 2005. A plastid gene phylogeny of the yam genus, Dioscorea: roots, fruits and Madagascar. Systematic Botany 30: 736–749. 82. TRIURIDACEAE THREETAILS FAMILY Ambrose BA, Espinosa-Matís S, Vásquez-Santana S, Márquez-Guzmán J, Alvarez-Buylla ER. 2006. Comparative developmental series of the Mexican triurids support a euanthial interpretation for the unusual reproductive axes of Lacandonia schismatica (Triuridaceae). American Journal of Botany 93: 15–35. Gandolfo MA, Nixon KC, Crepet WL, Stevenson DW, Friis EM. 1998. Oldest known fossils of monocotyledons. Nature 394: 532–533. Gandolfo MA, Nixon KC, Crepet WL. 2002. Triuridaceae fossil flowers from the Upper Cretaceous of New Jersey. American Journal of Botany 89: 1940–1957. Maas PHM, Rübsamen T. 1986. Triuridaceae. Flora Neotropica Monograph 40: 1–55. Maas-van de Kamer H, Maas PJM. 1994. Triuridopsis, a new monotypic genus in Triuridaceae. Plant Systematics and Evolution 192: 257–262. Márquez-Guzmán J, Engleman M, Martínez-Mena A, Martínez E, Ramos C. 1989. Anatomia reproductiva de Lacandonia schismatica (Lacandoniaceae). Annals of the Missouri Botanical Garden 76: 124–127. Melo A, Alves M. 2012. The discovery of Lacandonia (Triuridaceae) in Brazil. Phytotaxa 40: 21–25. Mennes CB, Smets EF, Moses SN, Merckx VSFT. 2013. New insights into the long-debated evolutionary history of Triuridaceae (Pandanales). Molecular Phylogenetics and Evolution 69: 994–1004. Rübsamen-Weustenfeld T. 1991. Morphologische, embryologische und systematische Untersuchungen an Triuridaceae. Bibliotheca Botanica 140: 1–113. Vergara-Silva F, Espinosa-Matías S, Ambrose BA, Vázquez-Santana S, Martínez-Mena A, Márquez-Guzmán J, Martínez E, Meyerowitz EM, Álvarez-Buylla ER. 2003. Inside-out flowers characteristic of Lacandonia schismatica evolved at least before its divergence from a closely related taxon, Triuris brevistyla. International Journal of Plant Sciences 164: 345–357. 83. VELLOZIACEAE BABOON-TAIL FAMILY Ayensu ES. 1973. Biological and morphological aspects of the Velloziaceae. Biotropica 5: 135–335. Behnke H-D, Treutelein J, Wink M, Kramer K,

FURTHER READING Schneider C, Kao P-C. 2000. Systematics and evolution of Velloziaceae, with special reference to sieve-element plastids and rbcL sequence data. Botanical Journal of the Linnean Society 134: 93–129. Behnke H-D, Hummel E, Hillmer S, Sauer-Gürth H, Sonzalez J, Wink M. 2013. A revision of African Velloziaceae based on leaf anatomy characters and rbcL nucleotide sequences. Botanical Journal of the Linnean Society 171: 22–94. Kao P-C. 1980. A new genus of Amaryllidaceae from China. Acta Phytotaxonomica Sinica (Chengdu) 1: 1–3. Mello-Silva R, Santos DYAC, Salatino MJF, Motta LB, Cattai MB, Sasaki D, Lovo J, Pita PB, Rocini C, Rodrigues CDN, Zarrei M, Chase MW. 2011. Five vicariant genera from Gondwana: the Velloziaceae as shown by molecules and morphology. Annals of Botany 108: 87–102. Menezes NL de, Mello-Silva R de, Mayo SJ. 1994. A cladistic analysis of the Velloziaceae. Kew Bulletin 49: 71–92. Smith LB, Ayensu ES. 1976. A revision of American Velloziaceae. Smithsonian Contributions to Botany 30: 1–173. Williams CA, Harborne JB, Menezes NL de. 1991. The utility of leaf flavonoids as taxonomic markers in the subfamily and generic classification of the Velloziaceae. Biochemical Systematics and Ecology 19: 483–495. 84. STEMONACEAE BAIBU FAMILY Ayensu ES. 1968. Comparative vegetative anatomy of the Stemonaceae (Roxburghiaceae). Botanical Gazette 129: 160–165. Bouman F, Devente N. 1992. A comparison of the structure of ovules and seeds in Stemona (Stemonaceae) and Pentastemona (Pentastemonaceae). Blumea 36: 501–514. Duyfjes BEE. 1991. Stemonaceae and Pentastemonaceae; with miscellaneous notes on members of both families. Blumea 36: 239–252. Ham RWJM. 1991. Pollen morphology of the Stemonaceae. Blumea 36: 127–159. Jiang R-W, Hon P-M, Xu Y-T, Chan Y-M, Xu H-X, Shaw PC, But PP. 2006. Isolation and chenotaxonomic significance of tuberostemospironine-type alkaloids from Stemona tuberosa. Phytochemistry 67: 52–57. Rogers GK. 1982. The Stemonaceae in the southeastern United States. Journal of the Arnold Arboretum 63: 327–336. Rudall PJ, Cunniff J, Wilkin P, Caddick LR. 2005. Evolution of dimery, pentamery and the monocarpellary condition in the monocot family Stemonaceae (Pandanales). Taxon 54: 701–711. Rudall PJ, Bateman RM. 2006. Morphological phylogenetic analysis of Pandanales: testing contrasting hypotheses of floral evolution. Systematic Botany 31: 223–238. Tomlinson PB, Ayensu ES. 1968. Morphology and anatomy of Croomia pauciflora (Stemonaceae). Journal of the Arnold Arboretum 49: 260–275. Van Steenis CGGJ. 1982. Pentastemona, a new 5-merous genus of monocotyledons from north Sumatra (Stemonaceae). Blumea 28: 151–163. Whetstone RD. 1984. Notes on Croomia pauciflora (Stemonaceae). Rhodora 86: 131–137. 85. CYCLANTHACEAE PANAMA HAT-PALM FAMILY Bennett BC, Alarcón R, Cerón C. 1992. The ethnobotany of Carludovica palmata Ruiz, Pavón

(Cyclanthaceae) in Amazonian Ecuador. Economic Botany 46: 233–240. Eriksson R. 1994. The remarkable weevil pollination of the Neotropical Carludovicoideae (Cyclanthaceae). Plant Systematics and Evolution 189: 75–81. Eriksson R. 1994. Phylogeny of the Cyclanthaceae. Plant Systematics and Evolution 190: 31–47. Franz NM. 2004. Analysing the history of the derelomine flower weevil-Carludovica association (Coleoptera: Curculionideae; Cyclanthaceae). Biological Journal of the Linnean Society 81: 483–517. French JC, Clancy K, Tomlinson PB. 1983. Vascular patterns in stems of the Cyclanthaceae. American Journal of Botany 70: 1386–1400. Gottsberger G. 1991. Pollination of some species of the Carludovicoideae, and remarks on the origin and evolution of the Cyclanthaceae. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 113: 221–235. Harling G. 1958. Monograph of the Cyclanthaceae. Acta Horti Bergiani 18: 1–428. Smith SY, Collinson ME, Rudall PJ. 2008. Fossil Cyclanthus (Cyclanthaceae, Pandanales) from the Eocene of Germany and England. American Journal of Botany 95: 688–699. Tomlinson PB, Wilder GJ. 1984. Systematic anatomy of Cyclanthaceae (Monocotyledoneae) — an overview. Botanical Gazette 145: 535–549. 86. PANDANACEAE SCREWPALM FAMILY Buerki S, Callmander MW, Devey DS, Chapell L, Gallaher T, Munzinger J, Haevermans T, Forest F. 2012. Straightening out the screw-pines: a first step in understanding phylogenetic relationships within Pandanaceae. Taxon 61: 1010–1020. Callmander MW, Chassot P, Küpfer P, Lowry II PP. 2003. Recognition of Martellidendron, a new genus of Pandanaceae, and its biogeographic implications. Taxon 52: 747–762. Callmander MW, Lowry II PP, Forest F, Devey DS, Beentje H, Buerki S. 2012. Benstonea Callm., Buerki (Pandanaceae): characterization, circumscription, and distribution of a new genus of screw-pines, with a synopsis of accepted species. Candollea 67: 323–343. Callmander MW, Booth T, Beentje H, Buerki S. 2013. Update on the systematics of Benstonea (Pandanaceae): when a visionary taxonomist foresees phylogenetic relationships. Phytotaxa 112: 57–60. Cox PA. 1981. Bisexuality in the Pandanaceae: new findings in the genus Freycinetia. Biotropica 13: 195–198. Cox PA. 1990. Pollination and the evolution of breeding systems in Pandanaceae. Annals of the Missouri Botanical Garden 77: 816–840. Huynh K-L. 1991–1992. The flower structure in the genus Freycinetia, Pandanaceae. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 112: 295–328; 114: 417–441. Jarzen DM. 1983. The fossil pollen record of the Pandanaceae. Garden’s Bulletin of the Straits Settlement 36: 163–175. Nadaf A, Zanan R. 2012. Indian Pandanaceae — an overview. Springer, Heidelberg. North CA, Willis AJ. 1971. Contributions to the anatomy of Sararanga (Pandanaceae). Botanical Journal of the Linnean Society 64: 411–421. Poppendieck H-H. 1987. Monoecy and sex changes in Freycinetia (Pandanaceae). Annals of the Missouri Botanical Garden 74: 314–320.

Stone BC. 1974. Towards an improved infrageneric classification in Pandanus (Pandanaceae). Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 94: 459–540. 87. CAMPYNEMATACEAE GREENMOUNTAINLILY FAMILY Dahlgren R, Lu A-M. 1985. Campynemanthe (Campynemaceae): morphology, microsporogenesis, early ovule ontogeny and relationships. Nordic Journal of Botany 5: 321–330. Goldblatt P. 1986. Systematics and relationships of the bigeneric Pacific family Campynemataceae (Liliales). Bulletin du Muséum Nationale d’Histoire Naturelle B, Adansonia IV, 8: 117–132. Lowry II PP, Goldblatt P, Tobe H. 1987. Notes on the floral biology, cytology, and embryology of Campynemanthe (Liliales: Campynemataceae). Annals of the Missouri Botanical Garden 74: 573–576. 88. CORSIACEAE GHOST-FLOWER FAMILY Cribb PJ. 1985. The saprophytic genus Corsia in the Solomon Islands. Kew Magazine 2: 320–323. Cribb PJ, Wilkin P, Clements M. 1995. Corsiaceae: a new family for the Falkland Islands. Kew Bulletin 50: 171–172. Fay MF, Chase MW, Rønsted N, Devey DS, Pillon Y, Pires JC, Petersen G, Seberg O, Davis JI. 2006. Phylogenetics of Liliales: summarized evidence from combined analyses of five plastid and one mitochondrial loci. Aliso 22: 559–565. Ibisch P, Neinhuis C, Rojas NP. 1996. On the biology, biogeography, and taxonomy of Arachnitis Phil. nom. cons. (Corsiaceae) in respect to a new record from Bolivia. Willdenowia 26: 321–332. Mennes CB, Lam VKY, Rudall PJ, Lyon SP, Graham SW, Smets EF, Merckx VSFT. 2015. Ancient Gondwana break-up explains the distribution of the mycoheterotrophic family Corsiaceae (Liliales). Journal of Biogeography 42: 1123–1136. Neyland R, Hennigan M. 2003. A phylogenetic analysis of large-subunit (26S) ribosome DNA sequences suggests that the Corsiaceae are polyphyletic. New Zealand Journal of Botany 41: 1–11. Rudall PJ, Eastman A. 2002. The questionable affinities of Corsia (Corsiaceae): evidence from floral anatomy and pollen morphology. Botanical Journal of the Linnean Society 138: 315–324. Van Royen P. 1972. Corsiaceae of New Guinea and surrounding areas. Webbia 27: 223–255. Zhang D, Saunders RMK, Hu C-M. 1999. Corsiopsis chinensis gen. et sp. nov. (Corsiaceae): first record of the family in Asia. Systematic Botany 24: 311–314. 89. MELANTHIACEAE BUNCHFLOWER FAMILY Buxbaum F. 1925. Vergleichende Anatomie der Melanthioideae. Repertorium Specierum Novarum Regni Vegetabile, Beihefte 29: 1–80. Fukuda I. 1990. Breeding systems in American and Asian Trillium species by means of chromosome analysis. Plant Species Biology 5: 65–72. Fuse S, Tamura MN. 2000. A phylogenetic analysis of the plastid matK gene with emphasis on Melanthiaceae sensu lato. Plant Biology 2: 415–427. Gates RR. 1918. A systematic study of the North American Melanthiaceae from the genetic standpoint. Journal of the Linnean Society, Botany 44: 131–172.

Plants of the World

689

FURTHER READING Kato H, Terauchi R, Utech FH, Kawano S. 1995. Molecular systematics of the Trilliaceae sensu lato as inferred from rbcL sequence data. Molecular Phylogenetics and Evolution 4: 184–193. Kim JS, Hong J-K, Chase MW, Fay MF, Kim J-H. 2013. Familial relationships of the monocot order Liliales based on a molecular phylogenetic analysis using four plastid loci: matK, rbcL, atpB and atpF-H. Botanical Journal of the Linnean Society 171: 5–21. Kupchan SM, Zimmerman JH, Afonso A. 1961. The alkaloids and taxonomy of Veratrum and related genera. Lloydia 24: 1–22. Kurabayashi M. 1958. Evolution and variation in Japanese species of Trillium. Evolution 12: 286–310. Pellicer J, Fay MF, Leitch IJ. 2010. The largest eukaryotic genome of them all? Botanical Journal of the Linnean Society 164: 10–15. Pellicer J, Kelly LJ, Leitch IJ, Zomlefer WB, Fay MF. 2013. A universe of dwarfs and giants: genome size and chromosome evolution in the monocot family Melanthiaceae. New Phytologist 201: 1484–1497. Takahashi M, Kawano S. 1989. Pollen morphology of the Melanthiaceae and its systematic implications. Annals of the Missouri Botanical Garden 76: 863–876. Zomlefer WB. 1996. The Trilliaceae in the southeastern United States. Harvard Papers in Botany 9: 91–120. Zomlefer WB, Williams NH, Whitten WM, Judd WS. 2001. Generic circumscription and relationships in the tribe Melanthieae (Liliales, Melanthiaceae), with emphasis on Zigadenus: evidence from ITS and trnL-F sequence data. American Journal of Botany 88: 1657–1669. Zomlefer WB, Judd WS, Whitten WM, Williams NH. 2006. A synopsis of Melanthiaceae (Liliales) with focus on character evolution in tribe Melanthieae. Aliso 22: 204–208. 90. PETERMANNIACEAE PETERMANN’S-VINE FAMILY Conran JG. 1988. Embryology and possible relationships of Petermannia cirrhosa (Petermanniaceae). Nordic Journal of Botany 8: 13–17. Conran JG, Christophel DC, Scriven LJ. 1994. Petermanniopsis angleseaënsis: an Australian fossil net-veined monocotyledon from Eocene Victoria. International Journal of Plant Sciences 155: 816–827. Conran JG, Christophel DC. 1999. A redescription of the Aústralian Eocene fossil monocotyledon Petermanniopsis (Lilianae: aff. Petermanniaceae). Transactions of the Royal Society of South Australia 123: 61–67. Tomlinson PB, Ayensu ES. 1969. Notes on the vegetative morphology and anatomy of the Petermanniaceae (Monocotyledones). Botanical Journal of the Linnean Society 62: 17–26. 91. ALSTROEMERIACEAE INCA-LILY FAMILY Aagesen L., Sanso AM. 2003. The phylogeny of the Alstroemeriaceae, based on morphology, rps16 intron, and rbcL sequence data. Systematic Botany 28: 47–69. Aker S, Healy W. 1990. The phytogeography of the genus Alstroemeria. Herbertia 46: 76–87. Arroyo SC, Leuenberger BE. 1988. Leaf morphology and taxonomic history of Luzuriaga (Philesiaceae). Willdenowia 17: 159–172.

690

Christenhusz, Fay & Chase

Buxbaum 1951. Die Grundachse von Alstroemeria und die Einheit ihres morphologischen Typus mit dem der echten Liliaceen. Phytomorphology 1: 170–184. Buxbaum F. 1954. Morphologie des Blüte und Frucht von Alstroemeria un der Anschluß der Alstroemerioideae bei den echten Liliaceae. Oesterreichische Botanische Zeitschrift 101: 337–352. Chacón J, Sousa A, Baeza CM, Renner SS. 2012. Ribosomal DNA distribution and a genus-wide phylogeny reveal patterns of chromosomal evolution in Alstroemeria (Alstroemeriaceae). American Journal of Botany 99: 1501–1512. Chacón J, De Assis MC, Meerow AW, Renner SS. 2012. From East Gondwana to Central America: historical biogeography of the Alstroemeriaceae. Journal of Biogeography 39: 1806–1818. Conran JG. 1988. Observations on the pollination ecology of Drymophila moorei Baker (Luzuriagaceae) in southeast Queensland. Victorian Naturalist 105: 43–47. Hofreiter A. 2006. Bomarea edulis (Tussac) Herb. a nearly forgotten pre-Columbian cultivated plant and its closest relatives (Alstroemeriaceae). Feddes Repertorium 117: 85–95. Hofreiter A. 2007. Biogeography and ecology of the Alstroemeriaceae-Luzuriagaceae clade in the highmountain regions of Central and South America. Harvard Papers in Botany 12: 259–284. Sanso AM, Xifreda CC. 2001. Generic delimitation between Alstroemeria and Bomarea (Alstroemeriaceae). Annals of Botany 88: 1057–1069. Sarwar AKMG, Hoshino Y, Araki H. 2010. Pollen morphology and infrageneric classification of Alstroemeria (Alstroemeriaceae). Grana 49: 227–242. 92. COLCHICACEAE NAKED-LADIES FAMILY Baker JG. 1879. A synopsis of Colchicaceae and the aberrant tribes of Liliaceae. Journal of the Linnean Society of London, Botany 17: 405–510. Buxbaum F. 1936. Die Entwicklungslinien der Lilioideae. I. Die Wurmbaeoideae. Botanisches Archiv 38: 213–293. Del Hoyo A, Pedrola-Monfort J. 2008. Phylogeny of Androcymbium (Colchicaceae) based on morphology and DNA sequences. Plant Systematics and Evolution 273: 151–167. Del Hoyo A, García-Marín JL, Pedrola-Monfort J. 2009. Temporal and spatial diversification of the African disjunct genus Androcymbium (Colchicaceae). Molecular Phylogenetics and Evolution 53: 848–861. Hara H. 1988. A revision of the Asiatic species of the genus Disporum (Liliaceae). University Museum, the University of Tokyo, Bulletin 31: 163–209. Hayashi K, Yoshida S, Kato H, Utech FH, Wigham D, Kawano S. 1998. Molecular systematics of the genus Uvularia and selected Liliales based on matK and rbcL gene sequence data. Plant Species Biology 13: 120–146. Le Roux LG, Robbertse PJ. 1994. Tuber ontogeny, morphology and vegetative reproduction of Gloriosa superba L. South African Journal of Botany 60: 321–324. Manning JC, Forest F, Vinnersten A. 2007. The genus Colchicum L. redefined to include Androcymbium Willd. based on molecular evidence. Taxon 56: 872–882. Nguyen TPA, Kim JS, Kim J-H. 2013. Molecular phylogenetic relationships and implications for the circumscription of Colchicaceae (Liliales).

Botanical Journal of the Linnean Society 172: 255–269. Nordenstam B. 1982. A monograph of the genus Ornithoglossum (Liliaceae). Opera Botanica 64: 1–51. Nordenstam B. 1986. The genus Wurmbea (Colchicaceae) in the Cape Region. Opera Botanica 87: 1–41. Persson K, Petersen G, Del Hoyo A, Seberg O, Jørgensen T. 2011. A phylogenetic analysis of the genus Colchicum L. (Colchicaceae) based on sequences from six plastid regions. Taxon 60: 1349–1365. Vinnersten A, Larsson S. 2010. Colchicine is still a chemical marker for the expanded Colchicaceae. Biochemical Systematics and Ecology 38: 1193–1198. Vinnersten A, Manning J. 2007. A new classification of Colchicaceae. Taxon 56: 171–178. Vinnersten A, Reeves G. 2003. Phylogenetic relationships within Colchicaceae. American Journal of Botany 90: 1455–1462. 93. PHILESIACEAE CHILEAN-BELLFLOWER FAMILY Cave MS. 1966. The female gametophytes of Lapageria rosea and Philesia magellanica. Gayana Botánica 15: 25–31. Schulze W. 1982. Beiträge zur Taxonomie der Liliifloren VII. Philesiaceae. Wissenschaftliche Zeitschrift der Friedrich-Schiller-Universität Jena 31: 277–283. 94. RIPOGONACEAE SUPPLEJACK FAMILY Conran JG, Clifford HT. 1985. The taxonomic affinities of the genus Ripogonum. Nordic Journal of Botany 5: 215–219. Conran JG, Carpenter RJ, Jordan JG. 2009. Early Eocene Ripogonum (Liliales: Ripogonaceae) leaf fossils from southern Australia. Australian Systematic Botany 22: 219–228. Macmillan BW. 1972. The biological flora of New Zealand. Ripogonum scandens J.R. et G. Forst. (Smilacaceae), supplejack, kareao. New Zealand Journal of Botany 10: 641–672. 95. SMILACACEAE CATBRIER FAMILY Arber A. 1920. Tendrils of Smilax. Botanical Gazette 69: 438–442. Cameron KM, Fu C. 2006. A nuclear rDNA phylogeny of Smilax (Smilacaceae). Aliso 22: 598–605. Chen S-C, Qiu Y-X, Wang A-L, Cameron KM, Fu C-X. 2006. A phylogenetic analysis of Smilacaceae based on morphological data. Acta Phytotaxonomica Sinica 44: 113–125. Chen S-C, Zhang X-P, Ni S-F, Fu C-X, Cameron KM. 2006. The systematic value of pollen morphology in Smilacaceae. Plant Systematics and Evolution 259: 19–37. Judd WS. 1998. The Smilacaceae in the southeastern United States. Harvard Papers in Botany 3: 147–169. Kong HH, Wang AL, Lee J, Fu CX. 2007. Studies of systematic evolution and karyotypic variation in Smilax and Heterosmilax (Smilacaceae). Acta Phytotaxonomica Sinica 45: 257–273. Koyama T. 1984. A taxonomic revision of the genus Heterosmilax (Smilacaceae). Brittonia 36: 184–205. Mangaly JK. 1968. A cytotaxonomic study of the herbaceous species of Smilax: section Coprosmanthus. Rhodora 70: 55–82, 247–273. Qi Z, Cameron KM, Li P, Zhao Y, Chen S, Chen

FURTHER READING G, Fu C. 2013. Phylogenetics, character evolution, and distribution patterns of the greenbriers, Smilacaceae (Liliales), a near cosmopolitan family of monocots. Botanical Journal of the Linnean Society 173: 535–548. Schulze W. 1982. Beiträge zur Taxonomie der Liliifloren VIII. Smilacaceae. Wissenschaftliche Zeitschrift der Friedrich-Schiller-Universität Jena 31: 285–289. Sun Z, Dilcher DL. 1988. Fossil Smilax from Eocene sediments in western Tennessee. American Journal of Botany 75 (suppl. 6): 118. 96. LILIACEAE LILY FAMILY Berg RY. 1959. Seed dispersal, morphology, and taxonomic position of Scoliopus, Liliaceae. Skrifter Utgitt av det Norske Videnskaps-Akademi i Oslo. Matematisk-Naturvidenskapelig Klasse 4: 1–56. Berg RY. 1962. Morphology and taxonomic position of Medeola, Liliaceae. Skrifter Utgitt av det Norske Videnskaps-Akademi i Oslo. MatematiskNaturvidenskapelig Klasse Ny Seri 3: 1–55. Chen SC, Hsia KC. 1977. A taxonomic study of the Chinese drug bei-mu (Fritillaria). Acta Phytotaxonomica Sinica 15: 31–46. Christenhusz MJM, Govaerts R, David JC, Hall T, Borland K, Roberts PS, Tuomisto A, Buerki S, Chase MW, Fay MF. 2013. Tiptoe through the tulips — cultural history, molecular phylogenetics and classification of Tulipa (Liliaceae). Botanical Journal of the Linnean Society 172: 280–328. Fay MF, Chase MW, Rønsted N, Devey DS, Pillon Y, Pires LC, Petersen G, Seberg O, Davis JI. 2006. Phylogenetics of Liliales: summarized evidence from combined analyses of five plastid and one mitochodrial loci. Aliso 22: 559–565. Gao Y-D, Harris AJ, Zhou S-D, He X-J. 2013. Evolutionary events in Lilium (including Nomocharis, Liliaceae) are temporally correlated with orogenies of the Q-T plateau and the Hengduan Mountains. Molecular Phylogenetics and Evolution 68: 443–460. Hayashi K, Kawano S. 2000. Molecular systematics of Lilium and allied genera (Liliaceae): phylogenetic relationships among Lilium and related genera based on the rbcL and matK sequence data. Plant Species Biology 15: 73–93. Kim JS, Hong J-K, Chase MW, Fay MF, Kim J-H. 2013. Familial relationships of the monocot order Liliales based on a molecular phylogenetic analysis using four plastid loci: matK, rbcL, atpB and atpF-H. Botanical Journal of the Linnean Society 171: 5–21. Lee CS, Kim S-C, Yeau SH, Lee NS. 2011. Major lineages of the genus Lilium (Liliaceae) based on nrDNA ITS sequences, with special emphasis on the Korean species. Journal of Plant Biology 54: 159–171. Patterson TB, Givnish TJ. 2002. Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from rbcL and ndhF sequence data. Evolution 56: 233–252. Peruzzi L, Leitch IJ, Caparelli KF. 2009. Chromosome diversity and evolution in Liliaceae. Annals of Botany 103: 459–475. Peterson A, Levichev IG, Peterson J. 2008. Systematics of Gagea and Lloydia (Liliaceae) and infrageneric classification of Gagea based on molecular and morphological data. Molecular Phylogenetics and Evolution 46: 446–465. Peterson A, Harpke D, Peruzzi L, Levichev IG, Tison J-M, Peterson J. 2009. Hydridisation drives

speciation in Gagea (Liliaceae). Plant Systematics and Evolution 278: 138–148. Rønsted N, Law S, Thornton H, Fay MF, Chase MW. 2005. Molecular phylogenetic evidence for the monophyly of Fritillaria and Lilium (Liliaceae; Liliales) and the infrageneric classification of Fritillaria. Molecular Phylogenetics and Evolution 35: 509–527. Shinwari ZK, Terauchi R, Utech FH, Kawano S. 1994. Recognition of the New World Disporum section Prosartes as Prosartes (Liliaceae) based on the sequence data of the rbcL gene. Taxon 43: 353–366. Zarrei M, Wilkin P, Fay MF, Ingrouille MJ, Zarre S, Chase MW. 2009. Molecular systematics of Gagea and Lloydia (Liliaceae; Liliales): implications of analyses of nuclear ribosomal and plastid DNA sequences for infrageneric classification. Annals of Botany 104: 125–142. 97. ORCHIDACEAE ORCHID FAMILY Albert VA. 1994. Cladistic relationships of the slipper orchids (Cypripedioideae: Orchidaceae) from congruent morphological and molecular data. Lindleyana 9: 115–132. Arditti J (ed.). 1994. Orchid biology: reviews and perspectives, VI. Wiley, New York. Bohman B, Phillips RD, Menz M, Berntsson BW, Flematti GR, Barrow RA, Dixon KW, Peakall R. 2014. Discovery of pyrazines as pollinator sex pheromones and orchid semiochemicals: implications for the evolution of sexual deception. New Phytologist 203: 939–952. Cameron KM. 2004. Utility of plastid psaB gene sequence of investigating intrafamilial relationships within Orchidaceae. Molecular Phylogenetics and Evolution 31: 1157–1180. Cameron KM, Arditti J, Kull T (eds). 2007. Orchid biology: reviews and perspectives, IX. Memoirs of the New York Botanical Garden 95. Cameron KM. 2009. On the value of nuclear and mitochondrial gene sequences for reconstructing the phylogeny of vanilloid orchids. Annals of Botany 104: 377–385. Cameron KM. 2011. Vanilla orchids. Natural history and cultivation. Timber Press, Portland, London. Chase MW, Hills HG. 1992. Orchid phylogeny, flower sexuality, and fragrance-seeking bees. Bioscience 42: 43–49. Chase MW, Williams NH, de Faria AD, Neubig KM, Amaral MCE, Whitten WM. 2009. Floral convergence in Oncidiinae (Cymbidieae; Orchidaceae): an expanded concept of Gomesa and a new genus Nohawilliamsia. Annals of Botany 104: 387–402. Chase MW, Cameron KM, Freudenstein JV, Pridgeon AM. 2015. An updated classification of Orchidaceae. Botanical Journal of the Linnean Society 177: 151–174. Cozzolino S, Windmer A. 2005. Orchid diversity: an evolutionary consequence of deception? Trends in Ecology and Evolution 20: 487–495. Darwin C. 1862. The various contrivances by which orchids are fertilized by insects. John Murray, London. Dixon KW, Kell SP, Barrett RL, Cribb PJ. eds. 2003. Orchid conservation. Natural History Publications (Borneo), Kota Kinabalu. Freudenstein JV, Rasmussen FN. 1996. Pollinium development and number in the Orchidaceae. American Journal of Botany 83: 813–824. Freudenstein JV, Rasmussen FN. 1999. What does morphology tell us about orchid relationships?

— A cladistic analysis. American Journal of Botany 86: 225–248. Freudenstein JV, van den Berg C, Goldman DH, Kores PJ, Molvray M, Chase MW. 2004. An expanded plastid DNA phylogeny of Orchidaceae and analysis of jacknife branch support strategy. American Journal of Botany 91: 149–157. Górniak M, Paun O, Chase MW. 2010. Phylogenetic relationships within Orchidaceae based on a low-copy nuclear coding gene, Xdh: congruence with organellar and nuclear ribosomal DNA results. Molecular Phylogenetics and Evolution 56: 784–795. Gravendeel B, Smithson A, Slik FJW, Schuiteman A. 2004. Epiphytism and pollinator specialisation: drivers for orchid diversity? Philosophical Transactions of the Royal Society of London B 359: 1523–1536. Kocyan A. 2010. Apostasioideae — the least known orchid subfamily. Malesian Orchid Journal 5: 125–138. Micheneau C, Fournel J, Warren BH, Hugel S, GauvinBialecki A, Pailler T, Strasberg D, Chase MW. 2010. Orthoptera, a new order of pollinator. Annals of Botany 105: 355–364. Micheneau C, Johnson SD, Fay MF. 2009. Orchid pollination: from Darwin to the present day. Botanical Journal of the Linnean Society 161: 1–9. Peakall R. 2007. Speciation in the Orchidaceae: confronting the challenge. Molecular Ecology 16: 2834–2837. Pemberton RW. 2013. Pollination in slipper orchids (Cypripedioideae): a review. Lankesteriana 13: 65–73. Phillips RD, Dixon KW, Peakall R. 2012. Low population genetic differentiation in the Orchidaceae: implications for the diversification of the family. Molecular Ecology 21: 5200–5220. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 1999. Genera orchidacearum. Volume 1. General introduction, Apostasioideae, Cypripedioideae. Oxford University Press, Oxford. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 2001. Genera orchidacearum. Volume 2. Orchidoideae (Part 1). Oxford University Press, Oxford. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 2003. Genera orchidacearum. Volume 3. Orchidoideae (Part 2), Vanilloideae. Oxford University Press, Oxford. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 2005. Genera orchidacearum. Volume 4. Epidendroideae (Part 1). Oxford University Press, Oxford. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 2009. Genera orchidacearum, vol. 5: Epidendroideae (part 2). Oxford University Press, Oxford. Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN, (eds). 2014. Genera orchidacearum, vol. 6: Epidendroideae (part 3). Oxford University Press, Oxford. Ramírez SR. 2009. Quick guide: orchid bees. Current Biology 19: 1061–1063. Ramírez SR, Gravendeel B, Singer RB, Marshall CR, Pierce NE. 2007. Dating the origin of Orchidaceae from a fossil orchid with its pollinator. Nature 448: 1042–1045. Rudall PJ, Bateman RM. 2002. Roles of synorganisation, zygomorphy and heterotopy in floral evolution: the gynostemium and labellum of orchids and other liliod monocots. Biological Review 77: 403–441.

Plants of the World

691

FURTHER READING Tremblay RL, Ackerman JD, Zimmerman JK, Calvo RN. 2005. Variation in sexual reproduction in orchids and its evolutionary consequence: a spasmodic journey to diversification. Biological Journal of the Linnean Society 84: 1–54. Van den Berg C, Goldman DH, Freudenstein JV, Pridgeon AM, Cameron KM, Chase MW. 2005. An overview of the phylogenetic relationships within Epidendroideae inferred from multiple DNA regions and recircumscription of Epidendreae and Arethuseae (Orchidaceae). American Journal of Botany 92: 613–624. Yam TW, Arditti J, Cameron KM. 2009. “The orchids have been a splendid sport” — an alternative look at Charles Darwin’s contribution to orchid biology. American Journal of Botany 96: 2128–2154. 98. BORYACEAE PINCUSHION-LILY FAMILY Chase MW, Rudall P, Conran JG. 1997. New circumscription and a new family of asparagoid lilies: genera formerly included in the Anthericaceae and associated taxa. Kew Bulletin 51: 667–680. Conran JG, Temby A. 2000. Embryology and affinities of the Boryaceae (Asparagales), p. 401–406 in Wilson KL, Morrison DA (eds). Monocots: systematics and evolution. CSIRO, Collingwood. Gaff DF, Loveys BR. 1984. Abscisic acid content and effects during dehydration of detached leaves of desiccation-tolerant plants. Journal of Experimental Botany 35: 1350–1358. 99. BLANDFORDIACEAE CHRISTMAS-BELLS FAMILY Di Fulvio TE, Cave MS. 1964. Embryology of Blandfordia nobilis Smith (Liliaceae) with special reference to its taxonomic position. Phytomorphology 14: 487–499. Johnson KA, Morrison DA, Goldsack G. 1994. Postfire flowering patterns in Blandfordia nobilis (Liliaceae). Australian Journal of Botany 42: 49–60. Kocyan A, Endress PK. 2001. Floral structure and development and systematic aspects of some “lower” Asparagales. Plant Systematics and Evolution 229: 187–216. Porter CL, Morrison DA, Johnson KA. 1992. Morphological variation within the genus Blandfordia (Liliaceae) in relation to its environment. Australian Systematic Botany 5: 373–386. Pyke GH, Day LP, Wale KA. 1988. Pollination ecology of Christmas bells (Blandfordia nobilis Sm.): effects of adding artificial nectar on pollen removal and seed set. Australian Journal of Ecology 13: 279–284. 100. ASTELIACEAE PINEAPPLE-GRASS FAMILY Birch JL, Keeley SC, Morden CW. 2012. Molecular phylogeny and dating of Asteliaceae (Asparagales): Astelia s.l. evolution provides insight into the Oligocene history of New Zealand. Molecular Phylogenetics and Evolution 65: 102–115. Mildenhall DC. 1980. New Zealand Late Cretaceous and Cenozoic plant biogeography: a contribution. Palaeogeography, Palaeoclimatology and Palaeoecology 31: 197–233. Moore LB. 1966. Australian asteliads (Liliaceae) with special reference to New Zealand species of Astelia subgenus Tricella. New Zealand Journal of Botany 4: 201–240. Rudall PJ, Chase MW, Cutler DF, Rusby J, De Bruijn AY. 1998. Anatomical and molecular systematics

692

Christenhusz, Fay & Chase

of Asteliaceae and Hypoxidaceae. Botanical Journal of the Linnean Society 127: 1–42. Skottsberg C. 1960. Astelia on Mauritius. Svensk Botanisk Tidskrift 54: 477–482. Wheeler CA. 1966. Cytotaxonomy of the large asteliads (Liliaceae) of the North Island of New Zealand. New Zealand Journal of Botany 4: 95–113. 101. LANARIACEAE LAMBTAILS FAMILY De Vos MP. 1963. Studies on the embryology and relationships of South African genera of the Haemodoraceae: Lanaria Ait. Journal of South African Botany 29: 79–90. Dora G, Edwards JM. 1991. Taxonomic status of Lanaria lanata and isolation of a novel biflavone. Journal of Natural Products 54: 796–801. Rudall PJ, Chase MW, Cutler DF, Rusby J, De Bruijn AY. 1998. Anatomical and molecular systematics of Asteliaceae and Hypoxidaceae. Botanical Journal of the Linnean Society 127: 1–42. 102. HYPOXIDACEAE STARGRASS FAMILY Baker JG. 1878. A synopsis of Hypoxidaceae. Journal of the Linnean Society, Botany 17: 93–126. Brackett A. 1923. Revision of the American species of Hypoxis. Rhodora 69: 120–147. Friedmann F. 1984. Note sur les Hypoxidaceae des Seychelles et description du genre nouveau Hypoxidia. Adansonia 4: 453–460. Judd WS. 2000. The Hypoxidaceae in the southeastern United States. Harvard Papers in Botany 5: 79–98. Kocyan A, Endress PK. 2001. Floral structure and development and systematic aspects of some “lower” Asparagales. Plant Systematics and Evolution 229: 187–216. Kocyan A, Snijman DA, Forest F, Devey DS, Freudenstein JV, Wiland-Szymańska J, Chase MW, Rudall PJ. 2011. Molecular phylogenetics of Hypoxidaceae — evidence from plastid DNA data and inferences on morphology and biogeography. Molecular Phylogenetics and Evolution 60: 122–136. Nordal I, Laane MM, Holt E, Staubo I. 1985. Taxonomic studies of the genus Hypoxis in East Africa. Nordic Journal of Botany 5: 15–30. Rudall PJ, Chase MW, Cutler DF, Rusby J, De Bruijn AY. 1998. Anatomical and molecular systematics of Asteliaceae and Hypoxidaceae. Botanical Journal of the Linnean Society 127: 1–42. Snijman DA, Kocyan A. 2013. The genus Pauridia (Hypoxidaceae) amplified to include Hypoxis sect. Ianthe, Saniella and Spiloxene, with revised nomenclature and typification. Phytotaxa 116: 19–33. Thompson MF. 1976, 1978, 1979. Studies in the Hypoxidaceae. I, II, III. Bothalia 12: 111–117, 429–435, 621–625. Wiland-Szymanska J. 2009. The genus Hypoxis L. (Hypoxidaceae) in East Tropical Africa: variability, distribution and conservation status. Biodiversity Research and Conservation 14: 1–129. 103. DORYANTHACEAE GYMEA-LILY FAMILY Blunden G, Yi Y, Jewers K. 1973. The comparative leaf anatomy of Agave, Beschorneria, Doryanthes and Furcraea species (Agavaceae: Agaveae). Botanical Journal of the Linnean Society 66: 157–179. Hoffmann C-C, Zetter R. 2010. Upper Cretaceous sulcate pollen from the Timerdyakh formation, Vilui Basin (Siberia). Grana 10: 170–193. Kocyan A, Endress PK. 2001. Floral structure and

development and systematic aspects of some “lower” Asparagales. Plant Systematics and Evolution 229: 187–216. Newman IV. 1928, 1929. The life history of Doryanthes excelsa. Part 1 and part 2. Proceedings of the Linnaean Society of New South Wales 53: 499–558; 54: 411–435. Wunderlich R. 1950. Die Agavaceae Hutchinsons im Lichte ihrer Embryologie, ihres Gynözeum-, Staublatt- und Blattbaues. Oesterreichische Botanische Zeitschrift 97: 437–502. 104. IXIOLIRIACEAE TARTAR-LILY FAMILY Arroyo SC. 1982. Anatomía vegetativa de Ixiolirion Fisch. ex Herb. (Liliales) y su significado taxonómico. Parodiana 1: 271–286. Dönmez EO, Işik S. 2008. Pollen morphology of Turkish Amaryllidaceae, Ixioliriaceae and Iridaceae. Grana 47: 15–38. Traub HP. 1943. The Ixiolirion tribe. Herbertia 9: 53–59. 105. TECOPHILAEACEAE CHILEAN-CROCUS FAMILY Arroyo S. 1986. Leaf anatomy in the Tecophilaeaceae. Botanical Journal of the Linnean Society 93: 323–328. Brummitt RK, Rudall PJ, Banks H, Johnson MAT, Docherty KA, Jones K, Chase MW, Rudall PJ. 1998. Taxonomy of the Cyanastroideae (Tecophilaeaceae): a multidisciplinary approach. Kew Bulletin 53: 769–803. Buerki S, Manning JC, Forest F. 2013. Spatio-temporal history of the disjunct family Tecophilaeaceae: a tale involving the colonisation of three Mediterranean-type ecosystems. Annals of Botany 111: 361–373. Carter S. 1962. Revision of Walleria and Cyanastrum (Tecophilaeaceae). Kew Bulletin 16: 185–195. De Vos MP. 1950. Die ontwikkeling van die saadknop en saad by Cyanella capensis: ‘n gefal van polyembryonie. South African Journal of Science 46: 220–226. Dulberger R, Ornduff R. 1980. Floral morphology and reproductive biology of four species of Cyanella (Tecophilaeaceae). New Phytologist 86: 45–56. Kocyan A, Endress PK. 2001. Floral structure and development and systematic aspects of some “lower” Asparagales. Plant Systematics and Evolution 229: 187–216. Manning JC, Goldblatt P. 2012. A revision of Tecophilaeaceae subfam. Tecophilaeoideae in Africa. Bothalia 42: 21–41. Simpson MG. 1985. Pollen ultrastructure of the Tecophilaeaceae. Grana 24: 77–92. Zavada MS, Scott G. 1993. Pollen morphology of Cyanella species (Tecophilaeaceae). Grana 32: 189–192. 106. IRIDACEAE IRIS FAMILY Arber A. 1921. The leaf structure of the Iridaceae. Annals of Botany 35: 301–336. Goldblatt P. 1990. Phylogeny and classification of Iridaceae. Annals of the Missouri Botanical Garden 77: 607–627. Goldblatt P. 1993. The woody Iridaceae: Nivenia, Klattia, Witsenia. Systematics, biology and evolution. Timber Press, Portland. Goldblatt P, Manning JC. 1998. Gladiolus in southern Africa. Fernwood Press, Vlaeberg. Goldblatt P, Manning JC. 2008. The iris family: natural history and evolution. Timber Press, Portland.

FURTHER READING Goldblatt P, Rodríguez A, Powell MP, Davies TJ, Manning JC, Van der Bank M, Savolainen V. 2008. Iridaceae ‘out of Australasia’? Phylogeny, biogeography, and divergence time based on plastid DNA sequences. Systematic Botany 33: 495–508. Harpke D, Meng S, Rutten T, Kerndorff H, Blattner FR. 2013. Phylogeny of Crocus (Iridaceae) based on one chloroplast and two nuclear loci: ancient hybridisation and chromosome number evolution. Molecular Phylogenetics and Evolution 66: 617–627. Karst L, Wilson CA. 2012. Phylogeny of the New World genus Sisyrinchium (Iridaceae) based on analyses of plastid and nuclear DNA sequence data. Systematic Botany 37: 87–95. Manning JC, Goldblatt P. 1990. Endothecium in Iridaceae and its systematic implications. American Journal of Botany 77: 527–532. Reeves, G. Chase MW, Goldblatt P, Rudall P, Fay, MF, Cox AV, Lejeune B, Souza-Chies T. 2001. Molecular systematics of Iridaceae: evidence from four plastid DNA regions. American Journal of Botany. 88: 2074–2087. Rodríguez A, Sytsma KJ. 2006. Phylogenetics of the “tiger-flower” group (Tigridieae, Iridaceae): molecular and morphological evidence. Aliso 22: 412–424. Tillich H-J. 2003. Seedling morphology in Iridaceae: indications of relationships within the family and to related families. Flora 198: 220–242. 107. XERONEMATACEAE POOR-KNIGHTSLILY FAMILY Chase MW, Rudall PJ, Fay MF. 2000. Xeronemataceae, a new family of asparagoid lilies from New Caledonia and New Zealand. Kew Bulletin 55: 865–870. Moore LB. 1957. The species of Xeronema (Liliaceae). Pacific Science 11: 355–362. 108. ASPHODELACEAE GRASSTREE FAMILY Adams SP, Leitch IJ, Bennett MD, Chase MW, Leitch AR. 2000. Ribosomal DNA evolution and phylogeny in Aloe (Asphodelaceae). American Journal of Botany 87: 1578–1583. Beaumont J, Cutler DF, Reynolds T, Vaughan JG. 1985. The secretory tissue of aloes and their allies. Israel Journal of Botany 34: 265–282. Carter S, Lavranos JJ, Newton LE, Walker CC. 2011. Aloes: the definitive guide. University of Chicago Press, Chicago. Chase MW, Reveal JW, Fay MF. 2009. A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161: 132–136. Couper RA. 1960. New Zealand Mesozoic and Cenozoic plant microfossils. Palynological Bulletin of the New Zealand Geological Survey 32: 1–87. Devey DS, Leitch I, Rudall PJ, Pires JC, Pillon Y, Chase MW. 2006. Systematics of Xanthorrhoeaceae sensu lato, with an emphasis on Bulbine. Aliso 22: 345–351. Ely F, Luque Arias R. 2006. Estudio morfoanatómico comparado de Eccremis coarctata (Ruiz, Pav.) Baker (Phormiaceae) en diferentes altitudes de la Cordillera de la Mérida. Plantula 4: 23–37. Fahn A. 1961. The anatomical structure of Xanthorrhoeaceae Dumort and its taxonomic position. Recent Advances in Botany 1: 155–160. Grace OM, Klopper RR, Smith GF, Crouch NR, Figueiredo E, Rønsted N, Van Wyk AE.

2013. A revised generic classification for Aloe (Xanthorrhoeaceae subfam. Asphodeloideae). Phytotaxa 76: 7–14. Holland PG. 1978. An evolutionary biogeography of the genus Aloe. Journal of Biogeography 5: 213–226. Lamont BB, Wittkuhn R, Korczynskyj D. 2004. Ecology and ecophysiology of grasstrees. Australian Journal of Botany 52: 561–582. Manning J, Boatwright JS, Daru BH, Maurin O, Van der Bank M. 2014. A molecular phylogeny and generic classification of Asphodelaceae subfamily Alooideae: a final resolution of the prickly issue of polyphyly in the alooids? Systematic Botany 39: 55–74. Reynolds T. 2004. Aloes, the genus Aloe. CRC Press, Boca Raton. Riley HP, Majumbar SK. 1979. The Aloineae: a biosystematic survey. University Press of Kentucky, Lexington. Rudall P, Chase MW. 1996. Systematics of Xanthorrhoeaceae sensu lato: new evidence for polyphyly. Telopea 6: 185–203. Schlitter J. 1940. Morphologie der Liliaceengattung Dianella Lam. Mitteilungen der Botanisches Museum der Universität Zürich 163: 5–283. Tuzlaci E. 1987. Revision of the genus Asphodeline (Liliaceae) — a new infrageneric classification. Candollea 42: 559–576. Van Jaarsveld EJ, Smith GF, Van Wyk, B-E. 1994. A cladistic analysis of Gasteria (Aloaceae). South African Journal of Botany 90: 467–470. Wendelbo P. 1964. On the genus Eremurus (Liliaceae) in southwest Asia. Acta Universitatis Bergensis, Series Mathematica rerumque Naturalium 5: 1–45. Wurdack KJ, Dorr LJ. 2009. The South American genera of Hemerocallidaceae (Eccremis and Pasithea): two introductions to the New World. Taxon 58: 1122–1132. 109. AMARYLLIDACEAE ONION FAMILY Chase MW, Reveal JW, Fay MF. 2009. A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161: 132–136. Fay MF, Chase MW. 1996. Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, Amaryllidaceae and Agapanthoideae. Taxon 45: 441–451. Fay MF, Rudall PJ, Chase MW. 2006. Molecular studies of subfamily Gilliesioideae (Alliaceae). Aliso 22: 367–371. Friesen N, Fritsch RM, Blattner FR. 2006. Phylogeny and new infrageneric classification in Allium (Alliaceae) based on nuclear ribosomal DNA ITS sequences. Aliso 22: 372–395. García N, Meerow AW, Soltis DE, Soltis PS. 2014. Testing deep reticulate evolution in Amaryllidaceae tribe Hippeastreae (Asparagales) with ITS and chloroplast sequence data. Systematic Botany 39: 75–89. Gurushidze M, Fritsch RM, Blattner FR. 2010. Specieslevel phylogeny of Allium subg. Melanocrommyum: incomplete lineage sorting, hybridisation and trnF gene duplication. Taxon 59: 829–840. Kwembeya EG, Bjora CS, Stedje B, Nordal I. 2007. Phylogenetic relationships of the genus Crinum (Amaryllidaceae) with emphasis on tropical African species: evidence from trnL-F and nuclear ITS DNA sequence data. Taxon 56: 801–810. Leighton FM. 1965. The genus Agapanthus L’Héritier. Journal of South African Botany, Supplement 4: 50.

Lledó MD, Davis AP, Crespo MB, Chase MW, Fay MF. 2004. Phylogenetic analysis of Leucojum and Galanthus (Amaryllidaceae) based on plastid matK and nuclear ribosomal spacer (ITS) DNA sequences and morphology. Plant Systematics and Evolution 246: 223–243. Meerow AW, Clayton JR. 2004. Generic relationships among the baccate-fruited Amaryllidaceae (tribe Haemantheae) inferred from plastid and nuclear non-coding DNA sequences. Plant Systematics and Evolution 244: 141–155. Meerow AW, Fay MF, Guy CL, Li Q-B, Zaman FQ, Chase MW. 1999. Systematics of Amaryllidaceae based on cladistic analysis of plastid rbcL and trnL-F sequence data. American Journal of Botany 86: 1325–1345. Meerow AW, Francisco-Ortega J, Kuhn DN, Schnell RJ. 2006. Phylogenetic relationships and biogeography within the Eurasian clade of Amaryllidaceae based on plastid ndhF and nrDNA ITS sequences; lineage sorting in a reticulate area. Systematic Botany 31: 42–60. Meerow AW, Guy CL, Li Q-B, Yang S-L. 2000. Phylogeny of the American Amaryllidaceae based on nrDNA ITS sequences. Systematic Botany 25: 708–726. Meerow AW, Lehmiller DJ, Clayton JR. 2003. Phylogeny and biogeography of Crinum L. (Amaryllidaceae) inferred from nuclear and limited plastid non-coding DNA sequences. Botanical Journal of the Linnean Society 141: 349–363. Meerow AW, Snijman DA. 2006. The never-ending story: multigene approaches to the phylogeny of Amaryllidaceae. Aliso 22: 355–366. Pellicer J, Hidalgo O, Walker J, Chase MW, Christenhusz MJM, Shackleford G, Leitch IJ, Fay MF. 2017. Genome size dynamics in tribe Gilliesieae (Amaryllidaceae subfamily Allioideae) in the context of polyploidy and unusual incidence of Robertsonian translocations. Botanical Journal of the Linnean Society 184: 16–31. Rønsted N, Symonds MRW, Birkholm T, Christensen SB, Meerow AW, Molander M, Mølgaard P, Petersen G, Rasmussen N, Van Staden J, Stafford GI, Jäger AK. 2012. Can phylogeny predict chemical diversity and potential medicinal activity of plants? A case study of Amaryllidaceae. BMC Evolutionary Biology 12: 182. Rudall PJ, Bateman RM, Fay MF, Eastman A. 2002. Floral anatomy and systematics of Alliaceae with particular reference to Gilliesia, a presumed insect mimic with strongly zygomorphic flowers. American Journal of Botany 89: 1867–1883. Santos-Gally R, Vargas P, Arroyo J. 2012. Insights into Neogene Mediterranean biogeography based on phylogenetic relationships of mountain and lowland lineages of Narcissus (Amaryllidaceae). Journal of Biogeography 39: 782–789. Scotland RW. 2013. Some observations on the homology of the daffodil corona. Pp. 297–303 in Wilkin P, Mayo SJ (eds). Early events in monocot evolution. Cambridge University Press, Cambridge. Vosa CG. 1975. The cytotaxonomy of the genus Tulbaghia. Annali Botanica 34: 47–121. 110. ASPARAGACEAE HYACINTH FAMILY Alvarez A, Köhler W. 1987. Morphología del polen de las Agavaceae y algunos géneros afines. Grana 26: 25–46.
Arber A. 1924. Danaë, Ruscus and Semele, a morphological study. Annals of Botany 38: 229–260. Blunden H, Yi Y, Jewers K. 1978. Steroidal sapogenins

Plants of the World

693

FURTHER READING from leaves of Agaveae species. Phytochemistry 17: 1923–1925. Bogler DJ. 1995. Systematics of Dasylirion: taxonomy and molecular phylogeny. Boletín de la Sociedad Botánica de México 56: 69–76. Bogler DJ, Pires JC, Francisco-Ortega J. 2006. Phylogeny of Agavaceae based on ndhF, rbcL, and ITS sequences: implications of molecular data for classification. Aliso 22: 313–328. Bogler DJ, Simpson BB. 1995. Chloroplast DNA study of the Agavaceae. Systematic Botany 20: 191–205. Bogler DJ, Simpson BB. 1996. Phylogeny of Agavaceae based on ITS rDNA sequence variation. American Journal of Botany 83: 1225–1235. Brittan NH. 1981. Revision of the genus Thysanotus R. Br. (Liliaceae). Brunonia 4: 67–181. Brown NE. 1915. Sansevieria: a monograph of all the known species. Bulletin of Miscellaneous Information of the Royal Gardens, Kew 1915: 185–261. Cave MS. 1948. Sporogenesis and embryo-sac development in Hesperocallis and Leucocoryne in relation to their systematic position. American Journal of Botany 35: 343–349. Chase MW, Reveal JW, Fay MF. 2009. A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161: 132–136. Chung MG, Jones SBJ. 1987. Pollen morphology of Hosta Tratt. (Funkiaceae) and related genera. Bulletin of the Torrey Botanical Club 116: 31–44. Clary KH, Simpson BB. 1995. Systematics and character evolution in the genus Yucca (Agavaceae): evidence from morphology and molecular analysis. Boletín de la Sociedad Botánica de México 56: 77–88. Conran JG. 1997. Paracordyline kerguelensis, an Oligocene monocotyledon macrofossil from the Kerguélen Islands. Alcheringa 21: 129–140. Conran JG, Rudall PJ, Chase MW. 1997. Two new monocotyledon families: Anemarrhenaceae and Behniaceae (Lilianae: Asparagales). Kew Bulletin 52: 995–999. Cooney-Sovetts C, Sattler R. 1986. Phylloclade development in the Asparagaceae: an example of homoeosis. Botanical Journal of the Linnean Society 94: 327–371. Erwin DM, Stockey RA. 1991. Soleredera rhizomorpha gen. et sp. nov., a permineralized monocotyledon from the Middle Eocene Princeton Chert of British Columbia, Canada. Botanical Gazette 152: 231–247. Fay MF, Chase MW. 1996. Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, Amaryllidaceae and Agapanthoideae. Taxon 45: 441–451. Fay MF, Rudall PJ, Sullivan S, Stobart KL, De Bruijn AY, Reeves G, Quamuz-Zaman F, Hong W-P, Joseph J, Hahn WJ, Conran JG, Chase MW. 2000. Phylogenetic studies of Asparagales based on four plastid DNA regions. Pp. 360–371, in Wilson KL, Morrison DA (eds) Monocots: systematics and evolution. CSIRO, Collingwood. Gentry HS. 1982. Agaves of continental North America. University of Arizona Press, Tucson. Hernández L. 1995. Análisis cladístico de la familia Agavaceae. Boletín de la Sociedad Botánica de México 56: 57–68. Hernández L. 1995. Taxonomic study of the Mexican genus Hemiphylacus (Hyacinthaceae). Systematic Botany 20: 546–554.

694

Christenhusz, Fay & Chase

Hume HH. 1961. Ophiopogon-Liriope complex. Baileya 9: 134–158. LaFrankie JV. 1986. Transfer of species of Smilacina to Maianthemum (Liliaceae). Taxon 35: 584–589. Lu AM. 1985. Embryology and possible relationships of Eriospermum. Nordic Journal of Botany 5: 229–240. Marais W, Reilly J. 1978. Chlorophytum and its related genera (Liliaceae). Kew Bulletin 32: 653–663. Nordal I, Eriksen RE, Fosing M. 1990. Studies in the generic delimitation of Anthericaceae. Mitteilungen aus dem Staatsinstitut für Allgemeine Botanik in Hamburg 23b: 535–559. Obermeyer AA. 1978. Ornithogalum: a revision of the southern African species. Bothalia 12: 323–376. Perry PL. 1994. A revision of the genus Eriospermum (Eriospermaceae). Contributions of the Bolus Herbarium 17: 1–320. Pfosser M, Speta F. 1999. Phylogenetics of Hyacinthaceae based on plastid DNA sequences. Annals of the Missouri Botanical Garden 86: 852–875. Pires JC, Fay MF, Davis WS, Hufford L, Rova J, Chase MW, Sytsma KJ. 2001. Molecular and phylogenetic analysis of Themidaceae (Asparagales). Kew Bulletin 56: 601–626. Pires JC, Maureira IJ, Rebman JP, Salazar GA, Cabrera LI, Fay MF, Chase MW. 2004. Molecular data confirm the phylogenetic placement of the enigmatic Hesperocallis (Hesperocallidaceae) with Agave. Madroño 51: 307–311. Pires JC, Sytsma KJ. 2002. A phylogenetic evaluation of a biosystematic framework: Brodiaea and related petaloid monocots (Themidaceae). American Journal of Botany 89: 1342–1359. Pires JC, Maureira IJ, Givnish TJ, Sytsma KJ, Seberg O, Petersen G, Davis JI, Stevenson DW, Rudall PJ, Fay MF, Chase MW. 2006. Phylogeny, genome size, and chromosome evolution of Asparagales. Aliso 22: 287–304. Rudall PJ, Chase MW. 1996. Systematics of the Xanthorrhoeaceae sensu lato: evidence for polyphyly. Telopea 6: 185–203. Rudall PJ, Engelman EM, Hanson L, Chase MW. 1998. Systematics of Hemiphylacus, Anemarrhena, and Asparagus. Plant Systematics and Evolution 211: 181–199. Seberg O, Petersen G, Davis JI, Pires JC, Stevenson DW, Chase MW, Fay MF, Devey DS, Jørgensen T, Sytsma KJ, Pillon Y. 2012. Phylogeny of the Asparagales based on three plastid and two mitochondrial genes. American Journal of Botany 99: 875–889. Schmid WG. 1991. The genus Hosta. Timber Press, Portland. Speta F. 1998. Systematische Analyse der Gattung Scilla L. s.l. (Hyacinthaceae). Phyton 38: 1–141. Steele PR, Hertwick KL, Mayfield D, McKain MR, Leebens-Mack J, Pires JC. 2012. Quality and quantity of data recovered from massively parallel sequencing: examples in Asparagales and Poaceae. American Journal of Botany 99: 330–348. Tidwell WD, Parker LR. 1990. Protoyucca shadishii gen. et spec. nov., an arborescent monocotyledon with secondary growth from the Middle Miocene of northwestern Nevada, USA. Review of Paleobotany and Palynology 62: 79–95. Tomlinson PB. 1965. Notes on the anatomy of Aphyllanthes (Liliaceae) and comparison with Eriocaulaceae. Journal of the Linnean Society, Botany 59: 163–173.

111. DASYPOGONACEAE SAVIOUR-GRASS FAMILY Anway JC. 1969. The evolution and taxonomy of Calectasia cyanea R.Br. (Xanthorrhoeaceae) in terms of its present-day variation and cytogenetics. Australian Journal of Botany 17: 147–159. Barrett RL, Dixon KW. 2001. A revision of the genus Calectasia (Calectasiaceae) with eight new species described from south-west Western Australia. Nuytsia 13: 411–448. Keighery GJ. 1983. Ballistochory (explosive seed dispersal) in Baxteria R.Br. (Xanthorrhoeaceae). Western Australian Naturalist 15: 163–166. Rudall PJ, Chase MW. 1996. Systematics of the Xanthorrhoeaceae sensu lato: evidence for polyphyly. Telopea 6: 185–203. Rudall PJ, Conran JG. 2012. Systematic placement of Dasypogonaceae among commelinid monocots: evidence from flowers and fruits. Botanical Review 78: 398–415. 112. ARECACEAE PALM FAMILY: Asmussen CB, Chase MW. 2001. Coding and noncoding plastid DNA in palm systematics. American Journal of Botany 88: 1103–1117. Balick MJ, Beck HT et al. 1990. Useful palms of the world. Columbia University Press, New York. Dransfield J, Ferguson IK, Uhl NW. 1990. The coryphoid palms: patterns of variation and evolution. Annals of the Missouri Botanical Garden 77: 802–815. Dransfield J, Uhl NW, Asmussen CB, Baker WJ, Harley M, Lewis C. 2008. Genera palmarum: the evolution and classification of palms. Royal Botanic Gardens, Kew. Fisher JB, Maidman KJ. 1998. Branching and architecture in palms: value for systematics. Memoirs of the New York Botanical Garden 83: 35–46. Henderson A. 1986. A review of pollination studies in the Palmae. Botanical Review 52: 221–259. Kaplan DR, Dengler NG, Dengler RE. 1982. The mechanism of plication inception in palm leaves. Canadian Journal of Botany 60: 2939–2975, 2999–3016. Moore HE Jr. 1973. The major groups of palms and their distribution. Gentes Herbarium 11: 27–141, Moore HE Jr, Uhl WR. 1982. Major trends of evolution in palms. Botanical Review 48: 1–69. PALMweb. Palms of the world online. http://www. palmweb.org (accessed 2 January 2012). Sowunmi MA. 1972. Pollen morphology of the Palmae and its bearing on taxonomy. Review of Palaeobotany and Palynology 13: 1–80. Tomlinson PB. 1990. The structural biology of palms. Clarendon Press, Oxford. Uhl NW. 1972. Inflorescence and flower structure in Nypa fruticans (Palmae). American Journal of Botany 59: 729–743. Uhl NW, Dransfield J. 1987. Genera palmarum, a classification of palms based on the work of Harold E. Moore, Jr. L.H. Bailey Hortorium, Cornell. Zona S, Henderson A. 1989. A review of animal-mediated seed dispersal of palms. Selbyana 11: 6–11. 113. HANGUANACEAE SUSUM FAMILY Lee DW, Yap KP, Liew FY. 1975. Serological evidence on the distinctness of the monocotyledonous families Flagellariaceae, Hanguanaceae and Joinvilleaceae. Botanical Journal of the Linnean Society 70: 77–81. Maury P. 1888. Sur les affinités du genre Susum.

FURTHER READING Bulletin de la Société Botanique de France 35: 410–417. Rudall PJ, Stevenson DW, Linder HP. 1999. Structure and systematics of Hanguana, a monocotyledon of uncertain affinity. Australian Journal of Botany 12: 311–330. Tillich H-J. 1996. Seeds and seedlings in Hanguanaceae and Flagellariaceae (monocotyledons). Sendtnera 3: 187–197. Tillich H-J, Sill E. 1999. Morphological and anatomical systematics of Hanguana Blume (Hanguanaceae) and Flagellaria L. (Flagellariaceae) with a description of a new species, Hanguana bogneri spec. nov. Sendtnera 6: 215–238. 114. COMMELINACEAE SPIDERWORT FAMILY Burns JH, Faden RB, Steppan SJ. 2011. Phylogenetic studies in the Commelinaceae subfamily Commelinoideae inferred from nuclear ribosomal and chloroplast DNA sequences. Systematic Botany 36: 268–276. Evans TM, Faden RB, Simpson MG, Sytsma KJ. 2000. Phylogenetic relationships in the Commelinaceae: I. A cladistic analysis of morphological data. Systematic Botany 25: 668–691. Evans TM, Sytsma KJ, Faden RB, Givnish TJ. 2003. Phylogenetic relationships in the Commelinaceae: II. A cladistic analysis of rbcL sequences and morphology. Systematic Botany 28: 270–292. Faden RB, Hunt DR. 1991. The classification of the Commelinaceae. Taxon 40: 19–31. Owens SJ. 1981. Self-incompatibility in the Commelinaceae. Annals of Botany 47: 567–581. Tomlinson PB. 1966. Anatomical data in the classification of Commelinaceae. Botanical Journal of the Linnean Society 59: 371–395. 115. PHILYDRACEAE FROGSMOUTH FAMILY Hamann U. 1966. Embryologische, morphologischanatomische und systematische Untersuchungen and Philydraceen. Willdenowia, Beihefte 4: 1–178. Saarela JM, Prentis PJ, Rai HS, Graham SW. 2008. Phylogenetic relationships in the monocot order Commelinales, with a focus on Philydraceae. Botany 86: 719–731. Skottsberg C. 1932. Bemerkungen über die Philydraceen. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 65: 253–274. 116. PONTEDERIACEAE WATER-HYACINTH FAMILY Daumann E. 1965. Das Blütennektarium bei den Pontederiaceen und die systematische Stellung dieser Familie. Preslia 37: 407–412. Eckenwalder JE, Barrett SCH. 1986. Phylogenetic systematics of Pontederiaceae. Systematic Botany 11: 373–391. Graham SW, Kohn JR, Morton BR, Eckenwalder JE, Barrett SCH. 1998. Phylogenetic congruence and discordance among one morphological and three molecular data sets from Pontederiaceae. Systematic Biology 47: 545–567. Kohn JR, Graham SW, Morton BR, Doyle JJ, Barrett SCH. 1996. Reconstruction of the evolution of reproductive characters in Pontederiaceae using evidence from chloroplast DNA restriction-site variation. Evolution 50: 1454–1469. Lunau K. 2006. Stamens and mimic stamens as components of floral colour patterns. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 127: 13–41.

Ness RW, Graham SW, Barrett SCH. 2011. Reconciling gene and genome duplication events: using multiple gene families to infer the phylogeny of the aquatic plant family Pontederiaceae. Molecular Biology and Evolution 28: 3009–3018. Price SD, Barrett SCH. 1984. The function and adaptive significance of tristyly in Pontederia cordata (Pontederiaceae). Biological Journal of the Linnean Society 21: 315–329. Strange A, Rudall PJ, Prychid CJ. 2004. Comparative f loral anatomy of Pontederiaceae. Botanical Journal of the Linnean Society 144: 395–408. 117. HAEMODORACEAE KANGAROO-PAW FAMILY Cheadle VI. 1968. Vessels in Haemodorales. Phytomorphology 18: 412–420. Hopper SD, Burbridge AH. 1978. Assortative pollination by red wattlebirds in a hybrid population of Anigozanthos Labill. (Haemodoraceae). Australian Journal of Botany 26: 335–350. Hopper SD, Smith RJ, Fay MF, Manning JC, Chase MF. 2009. Molecular phylogenetics of Haemodoraceae in the Greater Cape and southwest Australian floristic regions. Molecular Phylogenetics and Evolution 51: 19–30. Hopper SD, Fay MF, Rossetto M, Chase MW. 1999. A molecular phylogenetic analysis of the bloodroot and kangaroo paw family, Haemodoraceae: taxonomic, biogeographic and conservation implications. Botanical Journal of the Linnean Society 131: 285–299. MacFarlane TD, Hopper SD, Purdie RW, George AS, Patrick SJ. 1987. Haemodoraceae. Flora of Australia 45: 55–148. Simpson MG. 1983. Pollen ultrastructure of the Haemodoraceae and its taxonomic significance. Grana 22: 79–103. Simpson MG. 1990. Phylogeny and classification of the Haemodoraceae. Annals of the Missouri Botanical Garden 77: 722–784. 118. STRELITZIACEAE TRAVELLER’S-PALM FAMILY Fisher JB. 1976. Development of dichotomous branching and axillary buds in Strelitzia (Monocotyledonae). Canadian Journal of Botany 54: 578–592. Frost SK, Frost PGH. 1981. Sunbird pollination of Strelitzia nicolai. Oecologia 49: 379–384. Kress WJ, Schatz GE, Andrianifahanana M, Morland HS. 1994. Pollination of Ravenala madagascariensis (Strelitziaceae) by lemurs in Madagascar: evidence for an archaic evolutionary system? American Journal of Botany 81: 542–551. Tomlinson PB. 1960. The anatomy of Phenakospermum (Musaceae). Journal of the Arnold Arboretum 41: 287–297. 119. LOWIACEAE ORCHID-LILY FAMILY Holttum RE. 1970. The genus Orchidantha (Lowiaceae). Garden’s Bulletin Singapore 25: 239–246. Johansen LB. 2005. Phylogeny of Orchidantha (Lowiaceae) and the Zingiberales based on six DNA regions. Systematic Botany 30: 106–117. Kirchoff BK, Kunze H. 1995. Inf lorescence and f loral development in Orchidantha maxillarioides (Lowiaceae). International Journal of Plant Sciences 156: 159–171. Pedersen LB. 2001. Four new species of Orchidantha (Lowiaceae) from Sabah. Nordic Journal of Botany 21: 121–128.

Pedersen LB, Johansen B. 2004. Anatomy of the unusual stigma in Orchidantha (Lowiaceae). American Journal of Botany 91: 299–305. Sakai S, Inoue T. 1999. A new pollination system: dung beetle pollination discovered in Orchidantha inouei (Lowiaceae, Zingiberales) in Sarawak, Malaysia. American Journal of Botany 86: 56–61. 120. HELICONIACEAE PARROT-FLOWER FAMILY Andersson L. 1985. Revision of Heliconia subgen. Stenochlamys. Opera Botanica 82: 1–123. Andersson L. 1992. Revision of Heliconia subgen. Taenostrobus and subgen. Heliconia (MusaceaeHeliconioideae). Opera Botanica 111: 1–98. Berry F, Kress WJ. 1991. Heliconia. An identification guide. Smithsonian Institution Press, Washington. Kirchoff BK, Lagomarsino LP, Newman WH, Bartlett ME, Specht CD. 2009. Early floral development of Heliconia latispatha (Heliconiaceae), a key taxon for understanding the evolution of flower development in the Zingiberales. American Journal of Botany 96: 580–593. Kress WJ. 1983. Crossability barriers in Neotropical Heliconia. Annals of Botany 52: 131–147. Kress WJ. 1990. The taxonomy of Old World Heliconia (Heliconiaceae). Allertonia 6: 1–58. Tomlinson PB. 1959. An anatomical approach to the classification of the Musaceae. Journal of the Linnean Society, Botany 55: 799–809. 121. MUSACEAE BANANA FAMILY Fahn A. 1953. The origin of the banana inflorescence. Kew Bulletin 1953: 299–306. Häkkinen M, Väre H. 2008. Typification and checklist of Musa names (Musaceae) with nomenclatural notes. Adansonia 30: 63–112. Heslop-Harrison JS, Schwarzacher T. 2007. Domestication, genomics and the future for banana. Annals of Botany 100: 1073–1084. Lane IE. 1955. Genera and generic relationships in Musaceae. Mitteilungen der Botanischen Staatssammlung München 13: 114–131. Liu A-Z, Kress WJ, Lum D-Z. 2010. Phylogenetic analyses of the banana family (Musaceae) based on nuclear ribosomal (ITS) and chloroplast (trnL-F) evidence. Taxon 59: 20–28. Manchester SR, Kress WJ. 1993. Fossil bananas (Musaceae): Ensete oregonense sp. nov. from the Eocene of western North America and its phytogeographic significance. American Journal of Botany 80: 1264–1272. Nur N. 1976. Studies on pollination in Musaceae. Annals of Botany 40: 167–177. Pillay M, Tenkouaro A. (eds) 2011. Banana breeding: progress and challenges. CRC Press, Boca Raton. Simmonds NW. 1962. The evolution of the bananas. Longmans, London. Tomlinson PB. 1959. An anatomical approach to the classification of the Musaceae. Journal of the Linnean Society, Botany 55: 799–809. Väre H, Häkkinen M. 2011. Typification and check-list of Ensete Horan. names (Musaceae) with nomenclatural notes. Adansonia 33: 191–200. 122. CANNACEAE CANNA-LILY FAMILY Almeida AMR, Brown A, Specht CD. 2013. Tracking the development of the petaloid fertile stamen in Canna indica: insights into the origin of androecial petaloidy in the Zingiberales. AoB Plants 5: plt009. Gade D. 1966. Achira, the edible Canna, its cultivation

Plants of the World

695

FURTHER READING and use in the Peruvian Andes. Economic Botany 20: 407–415. Grootjen CJ, Bouman F. 1988. Seed structure in Cannaceae: taxonomic and ecological implications. Annals of Botany 61: 363–371. Kirchoff BK. 1983. Floral organogenesis in five genera of the Marantaceae and in Canna. American Journal of Botany 70: 508–523. Pai RM. 1965. Morphology of the flower in Cannaceae. Journal of Biological Science 8: 4–8. Maas-van de Kamer H, Maas PJM. 2008. The Cannaceae of the world. Blumea 53: 247–318. Tanaka N. 2001. Taxonomic revision of the family Cannaceae in the New World and Asia. Makinoa new series 1: 1–74. Tanaka N, Uchiyama H, Matoba H, Koyama T. 2009. Karyological analysis of the genus Canna. Plant Systematics and Evolution 280: 45–51. Tomlinson PB. 1961. The anatomy of Canna. Journal of the Linnean Society, Botany 66: 467–473. Trivedi BS, Verma CL. 1970. Silicified pseudostem of Canna L. from early Eocene of Deccan Intertrappean beds, M.P., India. Current Science 39: 442–443. 123. MARANTACEAE PRAYER-PLANT FAMILY Andersson L. 1981. The Neotropical genera of Marantaceae. Circumscription and relationships. Nordic Journal of Botany 1: 218–245. Andersson L. 1986. Revision of Maranta subg. Maranta (Marantaceae). Nordic Journal of Botany 6: 729–756. Andersson L, Chase MW. 2001. Phylogenetic classification of Marantaceae. Botanical Journal of the Linnean Society 135: 275–287. Borchsenius F, Suárez Suárez LS, Prince LM. 2012. Molecular phylogeny and redefined generic limits of Calathea (Marantaceae). Systematic Botany 37: 620–635. Holttum RE. 1951. The Marantaceae of Malaya. Gardens’ Bulletin Singapore 13: 254–296. Kennedy H. 1978. Systematics and pollination of the “closed-f lowered” species of Calathea (Marantaceae). Universit y of California Publications in Botany 71: 1–90. Kirchoff BK. 1983. Floral organogenesis in five genera of the Marantaceae and in Canna. American Journal of Botany 70: 508–523. Prince LM, Kress WJ. 2006. Phylogenetic relationships and classification in Marantaceae: insights from plastid DNA sequence data. Taxon 55: 281–296. Suksathan P, Gustafsson MH, Borchsenius F. 2009. Phylogeny and generic delimitation of Asian Marantaceae. Botanical Journal of the Linnean Society 159: 381–395. Tomlinson PB. 1961. Morphological and anatomical characteristics of the Marantaceae. Journal of the Linnean Society of London 58: 55–78. 124. COSTACEAE SPIRAL-GINGER FAMILY Grootjen CJ, Bouman F. 1981. Development of the ovule and seed in Costus cuspidatus (N. E. Br.) Maas (Zingiberaceae), with special reference to the information of the operculum. Botanical Journal of the Linnean Society 83: 27–39. Maas PJM. 1972. Costoideae (Zingiberaceae). Flora Neotropica, Monograph 8. Haffner, New York. Punt W. 1968. Pollen morphology of the American species of the subfamily Costoideae (Zingiberaceae). Review of Palaeobotany and Palynology 7: 31–43. Specht CD. 2006. Systematics and evolution of the

696

Christenhusz, Fay & Chase

tropical monocot family Costaceae (Zingiberales): a multiple dataset approach. Systematic Botany 31: 89–106. Specht CD, Kress WJ, Stevenson DW, DeSalle R. 2001. A molecular phylogeny of Costaceae (Zingiberales). Molecular Phylogenetics and Evolution 21: 333–345. Specht CD, Stevenson DW. 2006. A new phylogeny-based generic classification of Costaceae (Zingiberales). Taxon 55: 153–163. 125. ZINGIBERACEAE GINGER FAMILY Burtt BL. 1972. General introduction to papers on Zingiberaceae. Notes of the Royal Botanic Gardens Edinburgh 31: 155–165. Clark WD, Gaut BS, Duvall MR, Clegg MT. 1993. Phylogenetic relationships of the BromeliifloraeCommelinif lorae-Zingiberif lorae complex of monocots based on rbcL sequences. Annals of the Missouri Botanical Garden 80: 987–998. Hickey LH, Peterson RK. 1978. Zingiberopsis, a fossil genus of the ginger family from Late Cretaceous to Early Eocene sediments of western interior North America. Canadian Journal of Botany 56: 1136–1152. Holttum RE. 1950. The Zingiberaceae of the Malay Peninsula. Gardens’ Bulletin Singapore 13: 1–249. Kress WJ. 1990. The phylogeny and classification of Zingiberales. Annals of the Missouri Botanical Garden 77: 698–721. Kress WJ, Prince LM, Williams KJ. 2002. The phylogeny and a new classification of gingers: evidence from molecular data. American Journal of Botany 89: 1682–1696. Kress WJ, Liu A-Z, Newman M, Li Q-J. 2005. The molecular phylogeny of Alpinia (Zingiberaceae): a complex and polyphyletic genus of gingers. American Journal of Botany 92: 167–178. Kress WJ, Newman M, Poulsen AD, Specht CD. 2007. An analysis of generic circumscriptions in tribe Alpinieae (Alpinioideae: Zingiberaceae). Garden’s Bulletin Singapore 59: 113–128. Maas PJM. 1977. Renealmia, Costoideae (Additions), Zingiberaceae. Flora Neotropica monograph 18. New York Botanical Garden, Bronx. Ngamriabsakul C, Newman MF, Cronk QCB. 2004. The phylogeny of the tribe Zingibereae (Zingiberaceae) based on ITS (nrDNA) and trnL-F (cpDNA) sequences. Edinburgh Journal of Botany 60: 483–507. Pai RM. 1965. The floral anatomy of Elettaria cardamomum Maton: a reinvestigation. New Phytologist 64: 187–204. Ravindran PN, Babu KN (eds). 2005. Ginger, the genus Zingiber. CRC Press, Boca Raton. Sakai S, Kawakita A, Ooi K, Inoue T. 2013. Variation in the strength of association among pollination systems and floral traits: evolutionary changes in the floral traits of Bornean gingers (Zingiberaceae). American Journal of Botany 100: 546–555. Särkinen T, Newman MF, Maas PJM, Maas H, Poulsen AD, Harris DJ, Richardson JE, Clark A, Hollingsworth M, Pennington RT. 2007. Recent oceanic long-distance dispersal and divergence in the amphi-Atlantic rain forest genus Renealmia L.f. (Zingiberaceae). Molecular Phylogenetics and Evolution 44: 968–980. Tomlinson PB. 1956. Studies in the systematic anatomy of Zingiberaceae. Journal of the Linnean Society, Botany 55: 547–592. Williams KJ, Kress WJ, Manos PS. 2004. The phylogeny, evolution and classification of the genus

Globba (Zingiberaceae): appendages do matter. American Journal of Botany 91: 100–114. Wood TH, Whitten WM, Williams NH. 2000. Phylogeny of Hedychium and related genera (Zingiberaceae) based on ITS sequence data. Edinburgh Journal of Botany 57: 261–270. Wu T-L, Wu Q-G, Chen Z-Y. 1996. Proceedings of the second symposium on the family Zingiberaceae. University Press, Zhongshan. Xia Y-M, Kress WJ, Prince LM. 2004. Phylogenetic analysis of Amomum (Alpinioideae: Zingiberaceae) using ITS and matK sequence data. Systematic Botany 29: 334–344. Záveská E, Fér T, Šída P. Krak K, Marhold K, LeongŠkorničková J. 2012. Phylogeny of Curcuma (Zingiberaceae) based on plastid and nuclear sequences: proposal of the new subgenus Ecomata. Taxon 61: 747–763. 126. TYPHACEAE BULRUSH FAMILY Cook CDK, Nicholls MS. 1986/87. A monographic study of the genus Sparganium (Sparganiaceae). Botanica Helvetica 96: 213–267; 97: 1–44. Kaul RB. 1972. Adaptive leaf architecture in emergent and floating Sparganium. American Journal of Botany 59: 270–278. Kim K, Choi H-K. 2011. Molecular systematics and character evolution of Typha (Typhaceae) inferred from nuclear and plastid DNA sequence data. Taxon 60: 1417–1428. Krattinger K. 1978. Biosystematische Untersuchungen innerhalb der Gattung Typha L. Mitteilungen aus dem Botanischen Museum der Universität Zürich 298: 1–270. Müller-Doblies D. 1970. Über die Verwandtschaft von Typha und Sparganium im Infloreszenz- und Blütenbau. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 89: 451–562. Sulman JD, Drew BT, Drummond C, Hayasaka E, Sytsma KJ. 2013. Systematics, biogeography, and character evolution of Sparganium (Typhaceae): diversification of a widespread, aquatic lineage. American Journal of Botany 100: 2023–2039. Yeo RR. 1964. Life history of the common cattail. Weeds 12: 284–288. 127. BROMELIACEAE PINEAPPLE FAMILY Arenas P, Arroyo SC. 1988. Las espécies comestibles del género Bromelia (Bromeliaceae) del Gran Chaco. Candollea 43: 645–660. Barfuss MH, Samuel R, Till W, Stuessy TF. 2005. Phylogenetic relationships in subfamily Tillandsioideae (Bromeliaceae) based on DNA sequence data from seven plastid regions. American Journal of Botany 92: 337–351. Benzing DH. 1994. How much is known about Bromeliaceae in 1994? Selbyana 15: 1–7. Benzing DH, Givnish TJ, Bermides D. 1985. Absorptive trichomes in Brocchinia reducta (Bromeliaceae) and their evolutionary and systematic significance. Systematic Botany 10: 81–91. Bromeliad Society International. website: http://www. bsi.org/new/ Brown GK, Gilmartin AJ. 1989. Stigma types in Bromeliaceae — a systematic survey. Systematic Botany 14: 110–132. Butcher D, Gouda E. 2014. The new bromeliad taxon list. Bromelia Contact Group. Accessed online on 12 January 2014. http://botu07.bio.uu.nl/bcg/ taxonList.

FURTHER READING 128. RAPATEACEAE TOW-TOW FAMILY Carlquist S. 1961. Pollen morphology of Rapateaceae. Aliso 5: 39–66. Carlquist S. 1966. Anatomy of Rapateaceae: roots and stems. Phytomorpology 16: 17–38. Givnish T, Evans T, Sytsma K. 1994. Molecular evolution and adaptive radiation in South American elements of the plant family Rapateaceae. American Journal of Botany 81 (6, suppl.): 158. Givnish TJ, Millam KC, Evans TM, Hall JC, Pires JC, Berry PE, Sytsma KT. 2004. Ancient vicariance or recent long-distance dispersal? Inferences about phylogeny and South American-African disjunctions in Rapateaceae and Bromeliaceae based on ndhF sequence data. International Journal of Plant Sciences 165 (4 suppl.): S35–S54. Oriani A, Scatena VL. 2013. The taxonomic value of floral characters in Rapateaceae (Poales — monocotyledons). Plant Systematics and Evolution 299: 291–303. Venturelli M, Bouman F. 1988. Development of ovule and seed in Rapateaceae. Botanical Journal of the Linnean Society 97: 267–294. 129. XYRIDACEAE YELLOW-EYED-GRASS FAMILY Campbell LM. 2012. Pollen morphology of Xyridaceae (Poales) and its systematic potential. Botanical Review 78: 428–439. Carlquist S. 1960. Anatomy of Guayana Xyridaceae: Abolboda, Orectanthe, and Achlyphila. Memoirs of the New York Botanical Garden 10: 65–117. Kral R. 1988. The genus Xyris (Xyridaceae) in Venezuela and contiguous northern South America. Annals of the Missouri Botanical Garden 75: 522–722. Kral R. 1992. A treatment of American Xyridaceae exclusive of Xyris. Annals of the Missouri Botanical Garden 79: 819–885. Oriani A, Scatena VL. 2011. Reproductive biology of Abolboda pulchella and A. poarchon (Xyridaceae: Poales). Annals of Botany 107: 611–619. Oriani A, Scatena VL. 2012. Floral anatomy of xyrids (Poales): contributions to their reproductive biology, taxonomy, and phylogeny. International Journal of Plant Sciences 173: 767–779. Remizowa MV, Kuznetsov AN, Kuznetsova SP, Rudall PJ, Nuraliev MS, Sokoloff DD. 2012. Flower development and vasculature in Xyris grandis (Xyridaceae, Poales): a case study for examining petal diversity in monocot flowers with a double perianth. Botanical Journal of the Linnean Society 170: 93–111. Sajo MG, Wanderley MGL, de Menezes NL. 1997. Observações anatômicas sobre a vascularização floral em Xyris L. (Xyridaceae). Boletim Botânica da Universidade de São Paulo 16: 15–19. Sajo MG, Rudall PJ. 1999. Systematic vegetative anatomy and ensiform leaf development in Xyris (Xyridaceae). Botanical Journal of the Linnean Society 130: 171–182. Stützel T. 1990. “Appendices” am Gynoecium der Xyridaceen. Morphogenie, Funktion und systematische Bedeutung. Beiträge zur Biologie der Pflanzen 65: 275–299. 130. ERIOCAULACEAE PIPEWORT FAMILY Alves PGM, Scatena VL, Trovó M. 2013. Anatomy of scapes, bracts and leaves of Paepalanthus sect. Diphyomene (Eriocaulaceae, Poales) and its taxonomic implications. Brittonia 65: 262–272. Coan AI, Stützel T, Scatena VL. 2012. Comparative

embryology and taxonomic considerations in Eriocaulaceae (Poales). Feddes Repertorium 121: 268–284. Gomes de Andrade MJ, Giuletti AM, Rapini A, de Queiroz LP, Concição de Souza A, Macahado de Almeida PR, van den Berg C. 2010. A comprehensive phylogenetic analysis of Eriocaulaceae: evidence from nuclear (ITS) and plastid (psbAtrnH and trnL-F) DNA sequences. Taxon 59: 379–388. Martius CFP, von. 1835. Die Eriocaulaceen als selbständige Pflanzenfamilie, afgestellt und erläutert. Nova Acta Academiae Caesariae Leopoldino-Carolinae Germanicae Naturae Curiosum 17, 1: 1–72. Rosa MM, Scatena VL. 2007. Floral anatomy of Paepalanthoideae (Eriocaulaceae, Poales) and their nectariferous structures. Annals of Botany 99: 131–139. Scatena VL, Giulietti AM, Borba EL, van den Berg C. 2005. Anatomy of Brazilian Eriocaulaceae: correlation with taxonomy and habitat using multivariate analysis. Plant Systematics and Evolution 253: 1–22. Trovó M., Andrade MG, Sano O, Ribeiro P, van den Berg C. 2013. Molecular phylogenetics and biogeography of Neotropical Paepalanthoideae with emphasis on Brazilian Paepalanthus (Eriocaulaceae). Botanical Journal of the Linnean Society 171: 225–243. Trovó M, Stützel T. 2011. Diaspores in Eriocaulaceae: morphology, mechanisms, and implications. Feddes Repertorium 122: 464–465. 131. MAYACACEAE BOG-MOSS FAMILY Lourteig A. 1952. Mayacaceae. Notulae Systematicae 14: 234–248. Oriani A, Scatena VL. 2012. Floral anatomy of xyrids (Poales): contributions to their reproductive biology, taxonomy, and phylogeny. International Journal of Plant Sciences 173: 767–779. Stevenson D. 1983. Systematic implications of the floral morphology of the Mayacaceae. American Journal of Botany 70: 32. Uphof JCT. 1924. The physiological anatomy of Mayaca fluviatilis. Annals of Botany 38: 389–393. Venturelli M, Bouman F. 1986. Embryology and seed development in Mayaca fluviatilis. Acta Botanica Neerlandica 35: 497–516. 132. THURNIACEAE PALMIET FAMILY Cutler DF. 1965. Vegetative anatomy of Thurniaceae. Kew Bulletin 19: 431–441. Munro SL, Linder HP. 1997. The embryology and systematic relationships of Prionium serratum (Juncaceae: Juncales). American Journal of Botany 84: 850–860. Zimmermann MH, Tomlinson PB. 1968. Vascular construction and development in the aerial stem of Prionium (Juncaceae). American Journal of Botany 55: 1100–1109. 133. JUNCACEAE RUSH FAMILY Drábková LZ. 2010. Phylogenetic relationships within Juncaceae: evidence from five regions of plastid, mitochondrial and nuclear ribosomal DNA, with notes on morphology. Pp. 389–416, in Seberg O, et al. (eds). Diversity, phylogeny, and evolution in the monocotyledons. Århus, Aarhus University Press. Drábková LZ, Kirchner J, Seberg O, Petersen G, Vlacek C. 2003. Phylogeny of the Juncaceae based on rbcL sequences, with species emphasis on Luzula DC and Juncus L. Plant Systematics and

Evolution 240: 133–147. Kirchner J (et al.). 2002. Juncaceae (in three parts). Species plantarum, Flora of the World 6. Oriani A, Stützel T, Scatena VL. 2012. Contributions to the floral anatomy of Juncaceae (Poales — monocotyledons). Flora 207: 334–340. Snogerup S. 1993. A revision of Juncus subgen. Juncus (Juncaceae). Willdenowia 23: 23–73. Yamaguchi T, Tsukaya H. 2010. Evolutionary and developmental studies of unifacial leaves in monocots: Juncus as a model system. Journal of Plant Research 123: 35–41. 134. CYPERACEAE SEDGE FAMILY Bruhl J. 1991. Comparative development of some taxonomically critical floral/inflorescence features in Cyperaceae. Australian Journal of Botany 39: 119–127. Bruhl J. 1995. Sedge genera of the world: relationships and a new classification of the Cyperaceae. Australian Journal of Botany 8: 125–305. Inglis C. 1994. Checklist of economically important Cyperaceae. Cyperaceae Newsletter 13: 13–15. Kukkonen I. 1986. Special features of the inflorescence structure in the family Cyperaceae. Annales Botanici Fennici 23: 107–120. Muasya AM, Simpson DA, Chase MW. 2001. Phylogenetic relationships in Cyperus s.l. (Cyperaceae) inferred from plastid DNA sequence data. Botanical Journal of the Linnean Society 138: 145–153. Muasya AM, Simpson DA, Verboom GA, Goetghebeur P, Naczi RFC, Chase MW, Smets E. 2009. Phylogeny of Cyperaceae based on DNA sequence data: current progress and future prospects. Botanical Review 75: 2–21. Simpson DA, Furness CA, Hodkinson T, Muasya AM, Chase MW. 2003. Phylogenetic relationships in Cyperaceae subfamily Mapanioideae inferred from pollen and plastid DNA sequence data. American Journal of Botany 90: 1071–1087. Simpson DA, Muasya AM, Alves M, Bruhl JJ, Dhooge S, Chase MW, Furness CA, Ghamkhar K, Goetghebeur P, Hodkinson TR, Marchant AD, Nieuborg R, Reznicek AA, Roalson EH, Smets E, Starr JR, Thomas WW, Wilson KL, Zhang X. 2007. Phylogeny of Cyperaceae based on DNA sequence data–a new rbcL analysis. Aliso 23: 72–83. Vrijdaghs A, Reynders M, Larridon I, Muasya AM, Smets E, Goetghebeur P. 2010. Spikelet structure and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic evidence. Annals of Botany 105: 555–571. 135. RESTIONACEAE FYNBOS FAMILY Bell TL, Pate JS. 1993. Morphotypic differentiation in the SW Australian restiad Lyginia barbata R. Br. (Restionaceae). Australian Journal of Botany 41: 91–104. Briggs BG, Johnson LAS. 1998. New genera and species of Australian Restionaceae (Poales) Telopea 7: 345–373. Briggs BG, Linder HP. 2009. A new subfamilial and tribal classification of Restionaceae (Poales). Telopea 12: 333–345. Briggs BG, Marchant AD, Perkins AJ. 2014. Phylogeny of the restiid clade (Poales) and implications for the classification of Anarthriaceae, Centrolepidaceae and Australian Restionaceae. Taxon 63: 24–46. Carlquist S. 1976. Alexgeorgea, a bizarre new genus of Restionaceae from Western Australia. Australian Journal of Botany 24: 281–295.

Plants of the World

697

FURTHER READING Chanda S. 1966. On the pollen morphology of the Centrolepidaceae, Restionaceae and Flagellariaceae. Grana 7: 16–36. Hamann U. 1975. Neu Untersuchungen zur Embryologie und Systematik der Centrolepidaceae. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 96: 154–191. Harborne JB. 1979. Correlations between f lavonoid chemistry, anatomy and geography in the Restionaceae. Phytochemistry 18: 1323–1327. Hardy CR, Moline P, Linder HP. 2008. A phylogeny for the African Restionaceae, and new perspectives on morphology’s role in generating complete species phylogenies for large clades. International Journal of Plant Sciences 169: 337–390. Linder HP, Hardy CR. 2010. A generic classification of the Restioneae (Restionaceae), southern Africa. Bothalia 40: 1–35. Linder HP, Rudall PJ. 1993. The megagametophyte in Anarthria (Anarthriaceae, Poales) and its implication for the phylogeny of the Poales. American Journal of Botany 80: 1455–1464. Meney KA, Pate JS (eds). 1998. Australian rushes — biology, identification and conservation of Restionaceae and allied families. University of Western Australia Press, Perth. Rourke JP. 1974. On restios and roofs. Veld, Flora 4: 57–59. Sokoloff D, Remizowa MV, Linder HP, Rudall PJ. 2009. Morphology and development of the gynoecium in Centrolepidaceae: the most remarkable range of variation in Poales. American Journal of Botany 96: 1925–1940. Williams CA, Harborne JB, Greenham J, Briggs BG, Johnson LAS. 1997. Flavonoid evidence and the classification of Anarthriaceae within the Poales. Phytochemistry 45: 1189–1196. 136. FLAGELLARIACEAE WHIP-VINE FAMILY Chanda S. 1966. On the pollen morphology of the Centrolepidaceae, Restionaceae and Flagellariaceae. Grana 7: 16–36. Lee DW, Yap Kim Pin, Liew Foo Yew. 1975. Serological evidence on the distinctness of the monocotyledonous families Flagellariaceae, Hanguanaceae and Joinvilleaceae. Botanical Journal of the Linnean Society 70: 77–81. Mary Z, Pattan Shetty JK, Yoganarasimhan SN. 1985. Pharmacognostical studies on Flagellaria indica L. (Flagellariaceae). Journal of Economic and Taxonomic Botany 6: 1–8. Rudall PJ, Linder HP. 1988. Megagametophyte and nucellus in Restionaceae and Flagellariaceae. American Journal of Botany 75: 1777–1786. Sack FD. 1994. Structure of the stomatal complex of the monocot Flagellaria indica. American Journal of Botany 81: 339–344. Sajo MG, Rudall PJ. 2012. Morphological evolution in the graminid clade: comparative floral anatomy of the grass relatives Flagellariaceae and Joinvilleaceae. Botanical Journal of the Linnean Society 170: 393–404. Smithson E. 1956. The comparative anatomy of the Flagellariaceae. Kew Bulletin 11: 491–501. 137. JOINVILLEACEAE OHE FAMILY Brongniart MM, Gris A. 1861. Note sur le genre Joinvillea de Gaudichaud et sur la famille des Flagellariées. Bulletin de la Société Botanique de France 8: 264–269. Christophersen E. 1931. Notes on Joinvillea. Bernice P. Bishop Museum Bulletin 9: 2–7.

698

Christenhusz, Fay & Chase

Lee DW, Yap Kim Pin, Liew Foo Yew. 1975. Serological evidence on the distinctness of the monocotyledonous families Flagellariaceae, Hanguanaceae and Joinvilleaceae. Botanical Journal of the Linnean Society 70: 77–81. Newell TK. 1969. A study of the genus Joinvillea (Flagellariaceae). Journal of the Arnold Arboretum 50: 527–555. Smithson E. 1956. The comparative anatomy of the Flagellariaceae. Kew Bulletin 11: 491–501. Stone BC. 1981. Nomenclature of Joinvillea (Joinvilleaceae). Garden’s Bulletin of the Straits Settlement III, 34: 223–225. Tomlinson PB, Smith AC. 1970. Joinvilleaceae, a new family of monocotyledons. Taxon 19: 887–889. 138. ECDEIOCOLEACEAE KWONGAN-RUSH FAMILY Briggs BG. 2011. Ecdeiocolea rigens, a new species of Ecdeiocoleaceae (Poales) from Western Australia. Telopea 13: 69–75. Briggs BG, Johnson LAS. 1998. Georgeantha hexandra, a new genus and species of Ecdeiocoleaceae (Poales) from Western Australia. Telopea 7: 307–312. Cutler DF, Airy Shaw HK. 1965. Anarthriaceae and Ecdeiocoleaceae: two new monocotyledonous families, separated from the Restionaceae. Kew Bulletin 19: 160–168. Marchant AD, Briggs BG. 2007. Ecdeiocoleaceae and Joinvilleaceae, sisters of Poaceae (Poales): evidence from rbcL and matK data. Telopea 11: 437–450. Rudall PJ. 1990. Development of the ovule and megagametophyte in Ecdeiocolea monostachya. Australian Systematic Botany 3: 365–374. Rudall PJ, Stuppy W, Cunniff J, Kellogg EA, Briggs BG. 2005. Evolution of reproductive structures in grasses (Poaceae) inferred by sister-group comparison with their putative closest living relatives, Ecdeiocoleaceae. American Journal of Botany 92: 1432–1443. 139. POACEAE GRASS FAMILY Arber A. 1934. The Gramineae: a study of cereal, bamboo, and grass. Cambridge, University Press. Bamboo Phylogeny Group. 2012. An updated tribal and subtribal classification of the bamboos (Poaceae: Bambusoideae). Journal of the American Bamboo Society 24: 1–10. Barkworth ME, Arriaga MO, Smith JF, Jacobs SWL, Valdés-Reyna J, Bushman BS. 2008. Molecules and morphology in South American Stipeae (Poaceae). Systematic Botany 33: 719–731. Chase A. 1964. First book of grasses. The structure of grasses explained for beginners. Washington, Smithsonian Institution. Clayton WD, Renvoize SA (1986). Genera graminum. Grasses of the world. Royal Botanic Gardens, Kew. Clayton WD, Vorontsova MS, Harman KT, Williamson H. (2006 onwards). GrassBase — The online world grass flora. http://www.kew.org/data/grassbase/ index.html (accessed 2 January 2014). Dillon SL, Shapter FM, Henry RJ, Cordeiro G, Izquierdo L, Lee LS. 2007. Domestication to crop improvement: genetic resources for Sorghum and Saccharum (Andropogoneae). Annals of Botany 100: 975–989. Doyle JJ, Davis JI, Soreng RJ, Garvin D, Anderson MJ. 1992. Chloroplast DNA inversions and the origin of the grass family (Gramineae). Proceedings of the National Academy of Sciences of the USA 89: 7722–7726.

Duvall MR, Leseberg CH, Grennan CP, Morris IM. 2010. Molecular evolution and phylogenetics of complete chloroplast genomes in grasses. Pp. 437–450 in Seberg O, et al. (eds). Diversity, phylogeny, and evolution in the monocotyledons. Aarhus, University Press. Fuller DQ. 2007. Contrasting patterns in crop domestication and domestication rates: recent archaeobotanical insights from the Old World. Annals of Botany 100: 903–924. Fuller DQ. 2009. A comparative view of the evolution of grasses under domestication. New Phytologist 183: 273–290. Grass Phylogeny Working Group. 2001. Phylogeny and subfamilial classification of the grasses (Poaceae). Annals of the Missouri Botanical Garden 88: 373–457. Hastorf CA. 2009. Rio Balsas most likely region for maize domestication. Proceedings of the National Academy of Sciences of the USA 106: 4957–4958. Jacobs SWL, Everett J. (eds) 2000. Grasses: systematics and evolution. Melbourne, CSIRO. Jacobs SWL, Bayer R, Everett J, Arriaga MO, Barkworth ME, Sabin-Badereau A, Torres MA, Vazquez FM, Bagnall N. 2007. Systematics of the tribe Stipeae (Gramineae) using molecular data. Aliso 23: 349–361. Kellogg EA. 2000. The grasses: a case study in macroevolution. Annual Review of Ecology and Systematics 31: 217–238. Kellogg EA. 2001. Evolutionary history of the grasses. Plant Physiology 125: 1198–1205. Kellogg EA. 2013. C4 photosynthesys. Current Biology 23: R594–R599. McClure FA. 1968. The bamboos: a fresh perspective. Cambridge, Harvard University Press. Retallack GJ. 2001. Cenozoic expansion of grasslands and climate cooling. Journal of Geology 109: 407–426. Sang T. 2009. Genes and mutations underlying domestication transitions in grasses. Plant Physiology 149: 63–70. Steele PR, Hertwick KL, Mayfield D, McKain MR, Leebens-Mack J, Pires JC. 2012. Quality and quantity of data recovered from massively parallel sequencing: examples in Asparagales and Poaceae. American Journal of Botany 99: 330–348. Sweeney M, McCouch S. 2007. The complex history of the domestication of rice. Annals of Botany 100: 951–957. Vorontsova MS, Simon BK. 2012. Updating classifications to reflect monophyly: 10–20 percent of species names change in Poaceae. Taxon 61: 735–746. 140. CERATOPHYLLACEAE HORNWORT FAMILY Dilcher DL. 1989. The occurrence of fruits with affinities to Ceratophyllaceae in lower and midCretaceous sediments. American Journal of Botany 76: 162. Iwamoto A, Shimizo A, Ohba H. 2003. Floral development and phyllotactic variation in Ceratophyllum demersum (Ceratophyllaceae). American Journal of Botany 90: 1124–1130. Les DH. 1985. The taxonomic significance of plumule morphology in Ceratophyllum (Ceratophyllaceae). Systematic Botany 10: 338–346. Les DH. 1988. The origin and affinities of the Ceratophyllaceae. Taxon 37: 326–345. Les DH. 1989. The evolution of achene morphology in Ceratophyllum (Ceratophyllaceae), IV. Summary

FURTHER READING of proposed relationships and evolutionary trends. Systematic Botany 14: 254–262. Les DH, Garvin DK, Wimpee CF. 1991. Molecular evolutionary history of ancient aquatic angiosperms. Proceedings of the National Academy of Sciences of the U.S.A. 88: 10119–10123. 141. EUPTELEACEAE  ASIAN-ELM FAMILY Endress PK. 1969. Gesichtspunkte zur systematischen Stellung der Eupteleaceen (Magnoliales), Berichte der Schweizerischen Botanischen Gesellschaft 79: 229–278. Nast CG, Bailey IW. 1946. Morphology of Euptelea and comparison with Trochodendron. Journal of the Arnold Arboretum 27: 186–192. Ren Y, Li H-F, Zhao L, Endress PK. 2007. Floral morphogenesis in Euptelea (Eupteleaceae, Ranunculales). Annals of Botany 100: 185–193. Smith AC. 1946. A taxonomic review of Euptelea. Journal of the Arnold Arboretum 27: 175–185. Wang W, Lu A-M, Ren Y, Endress ME, Chen Z-D. 2009. Phylogeny and classification of Ranunculales: evidence from four molecular loci and morphological data. Perspectives in Plant Ecology, Evolution and Systematics 11: 81–110. 142. PAPAVERACEAE  POPPY FAMILY Brückner C. 1985. Frucht- und Samenanatomie von Pteridophyllum racemosum unde die Position der monotypischen Gattung in den Papaverales. Feddes Repertorium 96: 199–213. Carlquist S, Zona S. 1988. Wood anatomy of Papaveraceae with comments on vessel restriction patterns. IAWA Bulletin new series 9: 253–267. Carolan JC, Hook ILI, Chase MW, Kadereit JW, Hodkinson TR. 2006. Phylogenetics of Papaver and related genera based on DNA sequences from ITS nuclear ribosomal DNA and plastid trnL intron and trnL-F intergenic spacers. Annals of Botany 98: 141–155. Gunn CR. 1980. Seeds and fruits of Papaveracae and Fumariaceae. Seed Science and Technology 8: 3–58. Hoot SB, Kadereit JW, Blattner FR, Jork KB, Schwarzbach AE, Crane PR. 1997. Data congruence and phylogeny of the Papaveraceae s.l. based on four data sets: atpB and rbcL sequences, trnK restriction sites, and morphological characters. Systematic Botany 22: 575–590. Kadereit JW. 1988. Sectional affinities and geographical distribution in the genus Papaver L. (Papaveraceae). Beiträge zur Biologie der Pflanzen 63: 139–156. Kadereit JW, Blattner FR, Jork KB, Schwarzbach A. 1994. Phylogenetic analysis of the Papaveraceae s.l. (incl. Fumariaceae, Hypecoaceae, and Pteridophyllum) based on morphological characters. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 116: 361–390. Kadereit JW, Erbar C. 2011. Evolution of gynoecium morphology in Old World Papaveroideae: a combined phylogenetic/ontogenetic approach. American Journal of Botany 98: 1243–1251. Kadereit JW, Preston CD, Valtueña FJ. 2011. Is Welsh poppy, Meconopsis cambrica (L.) Vig. (Papaveraceae), truly a Meconopsis? New Journal of Botany 1: 80–87. Kadereit JW, Schwarzbach A, Jork KB. 1997. The phylogeny of Papaver s.l. (Papaveraceae): polyphyly or monophyly? Plant Systematics and Evolution 204: 75–98.

Kadereit JW, Sytsma KJ. 1992. Disassembling Papaver: a restriction site analysis of chloroplast DNA. Nordic Journal of Botany 12: 205–217. Lidén M. 1986. Synopsis of Fumarioideae (Papaveraceae) with a monography of the tribe Fumarieae. Opera Botanica 88: 1–133. Lidén M, Fukuhara T, Rylander J, Oxelman B. 1997. Phylogeny and classification of Fumariaceae, with emphasis on Dicentra s.l. based on the plastid gene rps16 intron. Plant Systematics and Evolution 206: 411–420. Pérez-Gutiérrez MA, Romero-García AT, Salinas MJ, Blanca G, Fernández NC, Suárez-Santiago VN. 2012. Phylogeny of the tribe Fumarieae (Papaveraceae s.l.) based on chloroplast and nuclear DNA sequences: evolutionary and biogeographic implications. American Journal of Botany 99: 517–528. Sugiura T. 1940. Chromosome studies on Papaveraceae with special reference to the phylogeny. Cytologia 10: 558–576. 143. CIRCAEASTERACEAE WITCH’S-STAR FAMILY Foster AS. 1963. The morphology and relationships of Circaeaster. Journal of the Arnold Arboretum 44: 299–321. Junell S. 1931. Die Entwicklungsgeschichte von Circaeaster agrestis. Svensk Botaniska Tidskrift 25: 238–270. Nowicke J, Skvarla JJ. 1982. Pollen morphology and the relationships of Circaeaster, of Kingdonia, and of Sargentodoxa to the Ranunculales. American Journal of Botany 69: 990–998. Ren Y, Li Z-J, Chang H-L, Lei Y-L, Lu A-M. 2004. Floral development of Kingdonia (Ranunculaceae s.l., Ranunculales). Plant Systematics and Evolution 247: 145–153. Ren Y, Hu Z-H. 1995. The morphology of the vegetative organs of Circaeaster agrestis (Ranunculaceae) and its taxonomic significance. Cathaya 7: 177–188. Ren Y, Wang M-L, Hu Z-H. 1998. Kingdonia, embryology and its systematic significance. Acta Phytotaxonomica Sinica 36: 423–427. Tian X, Zhang L, Ren Y. 2006. Development of f lowers and inf lorescences of Circaeaster (Circaeasteraceae, Ranu nculales). Plant Systematics and Evolution 256: 89–96. Wang X-M, Zhang P, Du Q-D, He H-X, Zhao L, Ren Y, Endress PK. 2012. Heterodichogamy in Kingdonia (Circaeasteraceae, Ranunculales). Annals of Botany 109: 1125–1132. 144. LARDIZABALACEAE ZABALA-FRUIT FAMILY Carlquist S. 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Botanical Journal of the Linnean Society 88: 257–277. Christenhusz MJM. 2012. An overview of Lardizabalaceae. Curtis’s Botanical Magazine 29: 235–276 (and species treatments in the same issue: volume 29 part 3). Hoot SB, Culham A, Crane PR.1995. The utility of atpB gene sequences in resolving phylogenetic relationships: comparison with rbcL and 18S ribosomal sequences in the Lardizabalaceae. Annals of the Missouri Botanical Garden 82: 194–207. Hoot SB, Culham A, Crane PR. 1995. Phylogenetic relationships of the Lardizabalaceae and Sargentodoxaceae: chloroplast and nuclear DNA

sequence data. Plant Systematics and Evolution, Supplement 9: 195–199. Kofuji R, Ueda K, Yamaguchi K, Shimizu T. 1994. Molecular phylogeny in the Lardizabalaceae. Journal of Plant Research 107: 339–348. Qin H-N. 1997. A taxonomic revision of the Lardizabalaceae. Cathaya 8–9: 1–214. Wang F, Li DZ, Yang JB. 2002. Molecular phylogeny of the Lardizabalaceae based on trnL-F sequences and combined chloroplast data. Acta Botanica Sinica 44: 971–977. Zhang X-H, Ren Y, Tian X-H. 2009. Floral morphogenesis in Sinofranchetia (Lardizabalaceae) and its systematic significance. Botanical Journal of the Linnean Society 160: 89–92. Zhang X-H, Ren Y. 2011. Comparative floral development in Lardizabalaceae (Ranunculales). Botanical Journal of the Linnean Society 166: 171–184. 145. MENISPERMACEAE MOONSEED FAMILY: Ortiz RdC, Kellogg EA, Van Der Werff H. 2007. Molecular phylogeny of the moonseed family (Menispermaceae): implications for morphological diversification. American Journal of Botany 94: 1425–1438. Herrera F, Manchester SR, Hoot SB, Wefferling KM, Carvalho MR, Jaramillo C. 2011. Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia. American Journal of Botany 98: 2004–2017. Jacques FMB, Bertolino P. 2008. Molecular and morphological phylogeny of Menispermaceae (Ranunculales). Plant Systematics and Evolution 274: 83–97. Jacques FMB, Wang W, Ortiz RdC, Li H-L, Zhou Z-K, Chen Z-D. 2011. Integrating fossils in a molecular-based phylogeny and testing them as calibration points for divergence time estimates in Menispermaceae. Journal of Systematics and Evolution 49: 25–49. Mennega AMW. 1982. Stem structure of New World Menispermaceae. Journal of the Arnold Arboretum 63: 145–171. Miers J. 1864–1871. A complete monograph of the Menispermaceae. Contributions to Botany 3(1). Williams, Norgate, London. Thanikaimondi G. 1986. Evolution of Menispermaceae. Canadian Journal of Botany 64: 3130–3133. Wang W, Ortiz RDC, Jacques FMB, Xiang X-G, Li H-L, Lin L, Li R-Q, Liu Y, Soltis PS, Soltis DE, Chen Z-D. 2012. Menispermaceae and the diversification of tropical rainforests near the Cretaceous-Paleogene boundary. New Phytologist 195: 470–478. Wang W, Wang H-C, Chen Z-D. 2007. Phylogeny and morphological evolution of tribe Menispermeae (Menispermaceae) inferred from chloroplast and nuclear sequences. Perspectives in Plant Ecology, Evolution and Systematics 8: 141–154. Wefferling KM, Hoot SB, Neves SS. 2013. Phylogeny and fruit evolution in Menispermaceae. American Journal of Botany 100: 833–905. 146. BERBERIDACEAE BARBERRY FAMILY Kim Y-D, Kim S-H, Kim C-H, Jansen RK. 2004. Phylogeny of Berberidaceae based on sequences of the chloroplast gene ndhF. Biochemical Systematics and Ecology 32: 291–301. Loconte H, Estes JR. 1989. Phylogenetic systematics of Berberidaceae and Ranunculales (Magnoliidae). Systematic Botany 14: 565–579.

Plants of the World

699

FURTHER READING Nickol MG. 1995. Phylogeny and inf lorescences of Berberidaceae — a morphological survey. Plant Systematics and Evolution, Supplement 9: 327–340. Nowicke J. Skvarla JJ. 1981. Pollen morphology and phylogenetic relationships of the Berberidaceae. Smithsonian Contributions to Botany 50: 1–83. Sastri RLN. 1969. Floral morphology, embryology and relationships of the Berberidaceae. Australian Journal of Botany 17:69–79. Walker JC. 1969. Plant pathology. McGraw-Hill, New York. Wang W, Chen Z-D, Li R-Q, Li J-H. 2007. Phylogenetic and biogeographic diversification of Berberidaceae in the Northern Hemisphere. Systematic Botany 32: 731–742. Ying T-S. 1979. On Dysosma Woodson and Sinopodophyllum Ying, gen. nov. of the Berber-idaceae. Acta Phytotaxonomica Sinica 17: 17–23. 147. RANUNCULACEAE BUTTERCUP FAMILY Cai Y-F, Li S-W, Liu Y, Quan S, Chen W, Xie YF, Jiang H-Z, Wei E-Z, Yin N-W, Wang L, Zhang R, Huang C-L, He X-H, Jiang M-F. 2009. Molecular phylogeny of Ranunculaceae based on internal transcribed spacer sequences. African Journal of Biotechnology 8: 5215–5224. Cai Y-F, Li S-W, Chen M, Jiang M-F, Liu Y, Xie Y-F, Sun Q, Jiang H-Z, Yin N-W, Wang L, Zhang R, Huang C-L, Lei K. 2010. Molecular phylogeny of Ranunculaceae based on rbcL sequences. Biologia 65: 997–1003. Carlquist S. 1995. Wood and bark anatomy of Ranunculaceae (including Hydrastis) and Glaucidiaceae. Aliso 14: 65–103. Compton JA, Culham A, Jury SL. 1998. Reclassification of Actaea to include Cimicifuga and Souliea (Ranunculaceae): phylogeny inferred from morphology, nrDNA ITS, and cpDNA trnL-F sequence variation. Taxon 47: 593–634. Emadzade K, Lehnebach C, Lockhart P, Hörandl E. 2010. A molecular phylogeny, morphology and classification of genera of Ranunculeae (Ranunculaceae). Taxon 59: 809–828. Emadzade K, Gehrke B, Linder HP, Hörandl E. 2011. The biogeographical history of the cosmopolitan genus Ranunculus (Ranunculaceae) in the temperate to meridional zones. Molecular Phylogenetics and Evolution 58: 4–21. Hoffmann MH, Von Hagen KB, Hörandl E, Röser M, Tkach NV. 2010. Sources of the Arctic flora: origins of Arctic species in Ranunculus and related genera. International Journal of Plant Sciences 171: 90–106. Hoot SB. 1991. Phylogeny of the Ranunculaceae based on epidermal characters and micromorphology. Systematic Botany 16: 741–755. Hoot SB. 1995. Phylogeny of the Ranunculaceae based on atpB, rbcL and 18S ribosomal DNA sequence data. Plant Systematics and Evolution, Supplement 9: 241–251. Hoot SB, Kramer J, Arroyo MTK. 2008. Phylogenetic position of the South American dioecious genus Hamadryas and related Ranunculeae (Ranunculaceae). International Journal of Plant Sciences 169: 433–443. Hoot SB, Meyer KM, Manning JC. 2012. Phylogeny and reclassification of Anemone (Ranunculaceae), with emphasis on austral species. Systematic Botany 37: 139–152. Hörandl E, Emadzade K. 2011. The evolution and biogeography of alpine species in Ranunculus

700

Christenhusz, Fay & Chase

(Ranunculaceae): a global comparison. Taxon 60: 415–426. Hörandl E, Emadzade K. 2012. Evolutionary classification: a case study on the diverse plant genus Ranunculus L. (Ranunculaceae). Perspectives in Plant Ecology, Evolution and Systematics 14: 310–324. Hörandl E, Paun O, Johansson JT, Lehnebach C, Armstrong T, Chen L, Lockhart P. 2005. Phylogenetic relationships and evolutionary traits in Ranunculus s.l. (Ranunculaceae) inferred from ITS sequence analysis. Molecular Phylogenetics and Evolution 36: 305–327. Jabbour F, Renner SS. 2011. Consolida and Aconitella are an annual clade of Delphinium (Ranunculaceae) that diversified in the Mediterranean basin and the Irano-Turanian region. Taxon 60: 1029–1040. Jabbour F, Renner SS. 2012. Phylogeny of Delphinieae (Ranunculaceae) shows that Aconitum is nested within Delphinium and that Late Miocene transitions to long life cycles in the Himalayas and southwest China coincide with bursts in diversification. Molecular Phylogenetics and Evolution 62: 928–942. Jabbour F, Renner SS. 2012. Spurs in a spur: perianth evolution in the Delphinieae (Ranunculaceae). International Journal of Plant Sciences 173: 1036–1054. Jensen U, Hoot SB, Johansson JT, Kosuge K. 1995. Systematics and phylogeny of the Ranunculaceae — a revised family concept on the basis of morphological data. Plant Systematics and Evolution, Supplement 9: 273–280. Kosuge K, Tamura M. 1989. Ontogenetic studies on petals of the Ranunculaceae. Japanese Journal of Botany 64: 65–67. Langlet O. 1932. Über Chromosomenverhältnisse und Systematik der Ranunculaceae. Svensk Botanisk Tidskrift 26: 381–400. Lehnebach CA, Cano A, Monsalve C, McLenachan P, Hörandl E, Lockhart O. 2007. Phylogenetic relationships of the monotypic Peruvian genus Laccopetalum (Ranunculaceae). Plant Systematics and Evolution 264: 109–116. Lehtonen S, Christenhusz MJM, Falck D. Sensitive phylogenetics of Clematis and its position in the Ranunculaceae. Botanical Journal of the Linnean Society 182: 825–867. Mikeda O, Kita K, Handa T, Yukawa T. 2006. Phylogenetic relationships of Clematis (Ranunculaceae) based on chloroplast and nuclear DNA sequences. Botanical Journal of the Linnean Society 152: 153–168. Paun O, Lehnebach C, Johansson JT, Lockhart P, Hörandl E. 2005. Phylogenetic relationships and biogeography of Ranunculus and allied genera (Ranunculaceae) in the Mediterranean region and the European Alpine system. Taxon 54: 911–930. Pfosser M, Sun B-Y, Stuessy TF, Jang C-G, Guo Y-P, Taejin K, Hwan KC, Kato H, Sugawara T. 2011. Phylogeny of Hepatica (Ranunculaceae) and origin of Hepatica maxima Nakai endemic to Ullung Island, Korea. Stapfia 95: 16–27. Santsiuk T. 1979. A palynological study of the tribe Ranunculeae. Opera Botanica 48: 1–76. Schuettpelz E, Hoot SB. 2004. Phylogeny and biogeography of Caltha (Ranunculaceae) based on chloroplast and nuclear DNA sequences. American Journal of Botany 91: 247–253. Soza VL, Brunet J, Liston A, Salles Smith P, Di Stilio VS. 2012. Phylogenetic insights into the correlates of dioecy in meadow rues (Thalictrum,

Ranunculaceae). Molecular Biology and Evolution 63: 180–192. Sun G, Dilcher DL, Wang H, Chen Z. 2011. A eudicot from the early Cretaceous of China. Nature 471: 625–628. Tamura M. 1972. Morphology and phyletic relationships of the Glaucidiaceae. Botanical Magazine (Tokyo) 85: 29–41. Tamura M. 1987. A classification of genus Clematis. Acta Phytotaxonomica et Geobotanica 38: 33–44. Tobe H, Keating RC. 1985. The morphology and anatomy of Hydrastis (Ranunculaceae): systematic evaluation of the genus. Botanical Magazine (Tokyo) 98: 291–316. Wang W, Cheng Z-D. 2007. Generic level phylogeny of Thalictroideae (Ranunculaceae) — implications for the taxonomic status of Paropyrum and petal evolution. Taxon 56: 811–821. Wang W, Hu H, Xiang X-G, Yu S-X, Chen Z-D. 2010. Phylogenetic placements of Calathodes and Megaleranthis (Ranunculaceae): evidence from molecular and morphological data. Taxon 59: 1712–1720. Wang W, Lu A-M, Ren Y, Endress ME, Chen Z-D. 2009. Phylogeny and classification of Ranunculales: evidence from four molecular loci and morphological data. Perspectives in Plant Ecology, Evolution and Systematics 11: 81–110. 148. SABIACEAE PAO-HUA FAMILY Carlquist S, Morrell PL, Manchester SR. 1993. Wood anatomy of Sabiaceae (s.l.): ecological and systematic implications. Aliso 13: 521–549. Raju MVS. 1952. Embryology of Sabiaceae. Current Science 21: 107–108. Ronse de Craene L-P, Wanntorp L. 2008. Morphology and anatomy of the flower of Meliosma (Sabiaceae): implications for pollination biology. Plant Systematics and Evolution 271: 79–91. Van Beusekom CF. 1971. Revision of Meliosma (Sabiaceae), sect. Lorenzanea excepted, living and fossil, geography and phylogeny. Blumea 19: 355–529. Van de Water TPM. 1980. A taxonomic revision of the genus Sabia (Sabiaceae). Blumea 26: 1–64. 149. NELUMBONACEAE SACRED-LOTUS FAMILY Borsch T, Barthlott W. 1994. Classification and distribution of the genus Nelumbo Adans. (Nelumbonaceae). Beiträge zur Biologie der Pflanzen 68: 421–450. Dieringer G, Cabrera R L, Mottaleb M. 2014. Ecological relationships between thermogenesis and pollination in Nelumbo lutea (Nelumbonaceae). American Journal of Botany 101: 357–364. Esau K. 1975. The phloem of Nelumbo nucifera Gaertn. Annals of Botany 39: 901–913. Estrada-Ruiz E, Upchurch Jr GR, Wolfe JA, CevallosFerriz SRS. 2011. Comparative morphology of fossil and extant leaves of Nelumbonaceae, including a new genus from the Late Cretaceous of western North America. Systematic Botany 36: 337–351. Gupta SC, Ahluwalia R. 1977. The carpel of Nelumbo nucifera. Phytomorphology 27: 274–281. Hall TF, Penfound WT. 1944. The biology of the American lotus, Nelumbo lutea (Willd.) Pers. American Midland Naturalist 31: 744–758. Hayes V, Schneider EL, Carlquist S. 2000. Floral development of Nelumbo nucifera (Nelumbonaceae). International Journal of Plant Sciences 161: S183–S191.

FURTHER READING Ohga I. 1926. On the structure of some ancient, but still viable fruits of Indian lotus, with special reference to their prolonged dormancy. Japanese Journal of Botany 3: 1–20. Seymour RS, Schultze-Motel P. 1996. Thermoregulating lotus flowers. Nature 383: 305. Shen-Miller J, Mudgett MB, Schopf JW, Clarke S, Berger R. 1995. Exceptional seed longevity and robust growth: ancient sacred lotus from China. American Journal of Botany 82: 1367–1380. Vogel S. 2004. Contributions to the functional anatomy and biology of Nelumbo nucifera (Nelumbonaceae) I. Pathways of air circulation. Plant Systematics and Evolution 249: 9–25. Yoo M-J, Soltis PS, Soltis DE. 2010. Expression of floral MADS-box genes in two divergent water lilies: Nymphaeales and Nelumbo. International Journal of Plant Sciences 171: 121–146. 150. PLATANACEAE PLANE-TREE FAMILY Baas P. 1969. Comparative anatomy of Platanus kerrii Gagnep. Botanical Journal of the Linnean Society 62: 413–421. Boothroyd LE. 1930. The morphology and anatomy of the inflorescence and flower of the Platanaceae. American Journal of Botany 17: 678–693. Doyle JA, Endress PK. 2010. Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: Magnoliidae and eudicots. Journal of Systematics and Evolution 48: 1–35. Feng Y, Oh S-H, Manos PS. 2005. Phylogeny and historical biogeography of the genus Platanus as inferred from nuclear and chloroplast DNA. Systematic Botany 30: 786–799. Jones BMG. 1968. The origin of London plane. Proceedings of the Botanical Society of the British Isles 7: 507–508. Manchester SR. 1986. Vegetative and reproductive morphology of an extinct plane tree (Platanaceae) from the Eocene of western North America. Botanical Gazette 147: 200–226. Santamour Jr FS. 1986. Checklist of cultivated Platanus (planetree). Journal of Arboriculture 12: 78–83. Schwarzwalder Jr RN, Dilcher DL. 1991. Systematic placement of the Platanaceae in the Hamamelidae. Annals of the Missouri Botical Garden 78: 962–969. Wang H, Dilcher DL, Schwarzwalder RN, Kvaček J. 2011. Vegetative and reproductive morphology of an extinct early Cretaceous member of Platanaceae from the Braun’s Ranch locality, Kansas, U.S.A. International Journal of Plant Sciences 172: 139–157. 151. PROTEACEAE SUGARBUSH FAMILY: Barker NP, Weston PH, Rourke JP, Reeves G. 2002. The relationship of the southern African Proteaceae as elucidated by internal transcribed spacer (ITS) DNA sequence data. Kew Bulletin 57: 867–883. Barker NP, Weston PH, Rutschmann F, Sauquet H. 2007. Molecular dating of the ‘Gondwanan’ plant family Proteaceae is only partially congruent with the timing of the break-up of Gondwana. Journal of Biogeography 34: 2012–2027. Bond WJ, Midgley JJ. 1995. Kill thy neighbor: an individualistic argument for the evolution of flammability. Oikos 73: 79–85. Cardillo M, Pratt R. 2013. Evolution of a hotspot genus: geographic variation in speciation and extinction rates in Banksia (Proteaceae). BMC

Evolutionary Biology 13: 155. He T, Lamont BB, Downes KS. 2011. Banksia born to burn. New Phytologist 191: 184–196. Hoot SB, Douglas AW. 1998. Phylogeny of the Proteaceae based on atpB and atpB-rbcL intergenic spacer regions. Australian Systematic Botany 11: 301–320. Itzstein-Davey F. 2004. A spatial and temporal Eocene palaeoenvironmental study, focusing on the Proteaceae family, from Kambalda, Western Australia. Review of Palaeobotany and Palynology 131: 159–180. Lynch AJJ, Barnes RW, Cambecedes J, Vaillancourt RE. 1998. Genetic evidence that Lomatia tasmanica (Proteaceae) is an ancient clone. Australian Journal of Botany 46: 25–33. Mast A, Milton EF, Jones EH, Barker RM, Barker WR, Weston PH. 2012. Time-calibrated phylogeny of the woody Australian genus Hakea (Proteaceae) supports multiple origins of insect-pollination among bird-pollinated ancestors. American Journal of Botany 99: 472–487. Orians GH, Milewski AV. 2007. Ecology of Australia: the effect of nutrient-poor soils and intense fires. Biological Review 82: 393–423. Sauquet H, Weston PH, Barker N, Anderson CL, Cantrill DJ, Savolainen V. 2009. Using fossils and molecular data to reveal the origins of the Cape proteas (subfamily Proteoideae). Molecular Phylogenetics and Evolution 51: 31–43. Shane MW, Lambers H. 2005. Cluster roots: a curiosity in context. Plant and Soil 274: 101–125. Valente M, Reeves G, Schnitzer J, Mason IP, Fay MF, Rebelo TG, Chase MW, Barraclough TG. 2010. Diversification of the African genus Protea (Proteaceae) in the Cape biodiversity hotspot and beyond: equal rates in different biomes. Evolution 64: 745–760. Weston PH, Barker NP. 2006. A new generic classification of the Proteaceae with an annotated checklist of genera. Telopea 11: 314–344. 152. TROCHODENDRACEAE WHEEL-TREE FAMILY Bailey IW, Nast CG. 1945. Morphology and relationships of Trochodendron and Tetracentron. I. Stem, root, and leaf. Journal of the Arnold Arboretum 26: 143–153. Crane PR, Manchester SR, Dilcher DL. 1991. Reproductive and vegetative structure of Nordenskioldia (Trochodendraceae), a vesselless dicotyledon from the early Tertiairy of the Northern Hemisphere. American Journal of Botany 78: 1311–1334. Endress PK. 1986. Floral structure, systematics and phylogeny in Trochodendrales. Annals of the Missouri Botanical Garden 62: 538–589. Grimsson F, Denk T, Zetter R. 2008. Pollen, fruits and leaves of Tetracentron (Trochodendraceae) from the Cainozoic of Iceland and western North America and their palaeobiogeographic implications. Grana 47: 1–14. Hacke UG, Sperry JS, Feild TS, Sano Y, Sikkema EH, Pitterman J. 2007. Water transport in vesselless angiosperms: conducting efficiency and cavitation safety. International Journal of Plant Sciences 168: 1113–1126. Mohanna Rao PR. 1983. Seed and fruit anatomy of Trochodendron aralioides. Phytomorphology 31: 18–23. Nast CG, Bailey IW. 1945. Morphology and relationships of Trochodendron and Tetracentron.

II. Inflorescence, flower and fruit. Journal of the Arnold Arboretum 26: 267–276. Pigg KB, Wehr WC, Ickertt-Bond SM. 2001. Trochodendron and Nordenskioldia (Trochodendraceae) from the middle Eocene of Washington state, USA. International Journal of Plant Sciences 162: 1187–1198. Ren Y, Chen L, Tian X-H, Zhang X-H, Lu A-M. 2007. Discovery of vessels in Tetracentron (Trochodendraceae) and its systematic significance. Plant Systematics and Evolution 267: 155–161. Rix M, Crane P. 2007. Tetracentron sinense (Tetracentraceae). Curtis’s Botanical Magazine 24: 238–247. 153. BUXACEAE BOX FAMILY Doust AN, Stevens PF. 2005. A reinterpretation of the staminate flowers of Haptanthus. Systematic Botany 30: 779–785. Goldberg A, Nelson CH. 1989. Haptanthus, a new dicotyledonous genus from Honduras. Systematic Botany 14: 16–19. Goldberg A, Aldern HA. 2005. Taxonomy of Haptanthus Goldberg, C. Nelson. Systematic Botany 30: 773–778. Pedersen KR, Von Balthazar M, Crane PR, Friis EM. 2007. Early Cretaceous floral structures and in situ tricolpate-striate pollen: new early dicots from Portugal. Grana 46: 176–196. Shipunov A, Shipunova E. 2011. Haptanthus story: rediscovery of enigmatic flowering plant from Honduras. American Journal of Botany 98: 761–763. Van Tieghem P. 1897. Sur les Buxacées. Annales des Science Naturelles, Botanique Sér. 8, 5: 289–338. Von Balthazar M, Endress PK. 2002. Development of inflorescences and flowers in Buxaceae and the problem of perianth interpretation. International Journal of Plant Sciences 163: 847–876. Von Balthazar M, Endress PK. 2002. Reproductive structures and systematics of Buxaceae. Botanical Journal of the Linnean Society 140: 193–228. Von Balthazar M, Schatz GE, Endress PK. 2003. Female f lowers and inf lorescences of Didymelaceae. Plant Systematics and Evolution 237: 199–208. Von Balthazar M, Schönenberger J, Qiu YL. 2002. Phylogenetic relationships in Buxaceae based on nuclear internal transcribed spacers and plastid ndhF sequences. International Journal of Plant Sciences 161: 785–792. 154. MYROTHAMNACEAE RESURRECTIONSHRUB FAMILY Carlquist S. 1976. Wood anatomy of Myrothamnus flabellifolia (Myrothamnaceae) and the problem of multiperforate perforation plates. Journal of the Arnold Arboretum 57: 119–126. Carlquist S. 1990. Leaf anatomy of Geissolomataceae and Myrothamnaceae as possible indicator of relationships to Bruniaceae. Bulletin of the Torrey Botanical Club 117: 420–428. Gaff DF. 1977. Desiccation tolerant vascular plants of southern Africa. Oecologia 31: 95–109. Jäger-Zürn I. 1966. Infloreszenz- und blütenmorphologische, sowie embryologische Untersuchungen an Myriothamnus Welw. Beiträge zur Biologie der Pflanzen 42: 241–271. Moore JP, Lindsey GG, Farrant JM, Brandt WF. 2007. An overview of the desiccation-tolerant resurrection plant Myrothamnus flabellifolia. Annals of Botany 99: 211–217.

Plants of the World

701

FURTHER READING 155. GUNNERACEAE GIANT-RHUBARB FAMILY González F, Bello MA. 2009. Intra-individual variation of f lowers in Gunnera subgenus Panka (Gunneraceae) and proposed apomorphies for Gunnerales. Botanical Journal of the Linnean Society 160: 262–283. Johanson C, Bergman B. 1994. Reconstitution of the Gunnera manicata Linde symbiosis: cyanobacterial specificity. New Phytologist 126: 643–652. Khamar HJ, Breathwaite EK, Prasse CE, Fraley ER, Secor CR, Chibane FL, Elhai J, Chiu WL. 2010. Multiple roles of soluble sugars in the establishment of Gunnera-Nostoc endosymbiosis. Plant Physiology 154: 1381–1389. Santi C, Bogusz D, Franche C. 2013. Biological nitrogen fixation in non-legume plants. Annals of Botany 111: 743–767. Soltis DE, Senters AR, Zanis MJ, Kimi S, Thompson JD, Soltis PS, Ronse de Craene LP, Endress PK, Farris JS. 2003. Gunnerales are sister to other core eurosids: implications for the evolution of pentamery. American Journal of Botany 90: 461–470. Wanntorp L, Ronse de Craene LP. 2005. The Gunnera flower: key to eudicot diversification or response to pollination mode? International Journal of Plant Sciences 170: 331–342. Wanntorp L, Wanntorp H-E. 2003. The biogeography of Gunnera L.: vicariance and dispersal. Journal of Biogeography 30: 979–987. Wanntorp L, Wanntorp H-E, Oxelman B, Källersjö M. 2001. Phylogeny of Gunnera. Plant Systematics and Evolution 226: 85–107. Wanntorp L, Wanntorp H-E, Rutishauser R. 2003. On the homology of the scales in Gunnera (Gunneraceae). Botanical Journal of the Linnean Society 142: 301–308. Wanntorp L, Dettmann ME, Jarzen DM. 2004. Tracking the Mesozoic distribution of Gunnera: comparison with the fossil pollen species Tricolpites reticulatus Cookson. Review of Palaeobotany and Palynology 132: 163–174. 156. DILLENIACEAE GUINEAFLOWER FAMILY Aymard C GA. 1997. Dilleniaceae novae Neotropicae IX: Neodillenia, a new genus from the Amazon basin. Harvard Papers in Botany 10: 121–131. Craven LA, Dunlop CR. 1992. A taxonomic revision of Pachynema (Dilleniaceae). Australian Systematic Botany 5: 477–500. Endress PK. 1997. Relationships between f loral organisations, architecture, and pollination mode in Dillenia (Dilleniaceae). Plant Systematics and Evolution 206: 99–118. Gurni AA, Kubitzki K. 1981. Flavonoid chemistry and systematics of the Dilleniaceae. Biochemical Systematics and Ecology 9: 109–114. Hoogland RD. 1952. A revision of the genus Dillenia. Blumea 7: 1–145. Horn JW. 2009. Phylogenetics of Dilleniaceae using sequence data from four plastid loci (rbcL, infA, rps4, rpl16 intron). Journal of Plant Sciences 170: 794–813. Kubitzki K. 1971. Doliocarpus, Davilla und verwandte Gattungen (Dilleniaceae). Mitteilungen der Botanisches Staatssammlung München 9: 1–105. Sastri RLN. 1958. Floral morphology and embryology of some Dilleniaceae. Botaniska Notiser 111: 495–511. Tucker SC, Bernhardt P. 2000. Floral ontogeny, pattern formation, and evolution in Hibbertia and

702

Christenhusz, Fay & Chase

Adrastea (Dilleniaceae). American Journal of Botany 87: 1915–1936. 157. PERIDISCACEAE RINGFLOWER FAMILY Brenan JPM. 1952. Plants of the Cambridge Expedition, 1947–1948: II. A new order of flowering plants from the British Cameroons. Kew Bulletin 1952: 227–236. Brenan JPM. 1954. Soyauxia, a second genus of Medusandraceae. Kew Bulletin 1954: 507–511. Davis CC, Chase MW. 2004. Elatinaceae are sister to Malpighiaceae; Peridiscaceae belong to Saxifragales. American Journal of Botany 91: 262–273. Metcalfe CR. 1952. Medusandra richardsiana Brenan. Anatomy of the leaf, stem and wood. Kew Bulletin 1952: 237–244 Metcalfe CR. 1962. Notes on the systematic anatomy of Whittonia and Peridiscus. Kew Bulletin 15: 472–475. Nyananyo BL. 1987. Systematic survey of the leaf epidermis in the Medusandraceae (Rosidae). Feddes Repertorium 98: 595–598. Soltis DE, Clayton JW, Davis CC, Gitzendanner MA, Cheek M, Savolainen V, Amorim AM, Soltis PS. 2007. Monophyly and relationships of the enigmatic family Peridiscaceae. Taxon 56: 65–73. 158. PAEONIACEAE PEONY FAMILY Hiepko P. 1966. Zur Morphologie, Anatomie und Funktion des Diskus der Paeoniaceae. Berichte der Deutsche Botanische Gesellschaft 79: 233–245. Hong DY. 2012. Peonies of the World: polymorphism and diversity. Royal Botanic Gardens, Kew. Sang T, Crawford DJ, Stuessy TF. 1997. Chloroplast DNA phylogeny, reticulate evolution, and biogeography of Paeonia (Paeoniaceae). American Journal of Botany 84: 1120–1136. 159. ALTINGIACEAE SWEETGUM FAMILY Bogle AL. 1986. The floral morphology and vascular structure of the Hamamelidaceae: subfamily Liquidambaroideae. Annals of the Missouri Botanical Garden 73: 325–347. Ickert-Bond SM, Wen J. 2006. Phylogeny and biogeography of Altingiaceae: evidence from combined analysis of five non-coding chloroplast regions. Molecular Phylogenetics and Evolution 39: 512–528. Ickert-Bond SM, Wen J. 2013. A taxonomic synopsis of Altingiaceae with nine new combinations. PhytoKeys 31: 21–61. Pigg KB, Ickert-Bond SM, Wen J. 2004. Anatomically preserved Liquidambar (Altingiaceae) from the Middle Miocene of Yakima Canyon, Washington state, USA, and its biogeographic implications. American Journal of Botany 91: 499–509. Wu W, Zhou R, Huang Y, Boufford D, Shi S. 2010. Molecular evidence for natural intergeneric hybridisation between Liquidambar and Altingia. Journal of Plant Research 123: 231–239. 160. HAMAMELIDACEAE WITCH-HAZEL FAMILY Endress PK. 1977. Evolutionary trends in the Hamamelidales-Fagales group. Plant Systematics and Evolution, Supplement 1: 321–347. Endress PK. 1978. Blütenontogenese, Blütenabgrenzung und systematische Stellung der perianthlosen Hamamelidoideae. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 100: 249–317. Goldblatt P, Endress PK. 1977. Cytology and

evolution in Hamamelidaceae. Journal of the Arnold Arboretum 58: 67–71. Li J, Bogle AL. 2001. A new suprageneric classification system of Hamamelidoideae based on morphology and sequences of nuclear and chloroplast DNA. Harvard Papers in Botany 5: 499–515. Magallón SA. 2007. From fossils to molecules: phylogeny and the core eudicot floral groundplan in Hamamelidoideae (Hamamelidaceae, Saxifragales). Systematic Botany 32: 317–347. Manchester SR, Chen ZD, Lu AM, Uemura K. 2009. Eastern Asian endemic seed plant genera and their paleogeographic history throughout the Northern Hemisphere. Journal of Systematics and Evolution 47: 1–41. Xie L, Yi TS, Li R, Li DZ, Wen J. 2010. Evolution and biogeographic diversification of the witch-hazel genus (Hamamelis L., Hamamelidaceae) in the Northern Hemisphere. Molecular Phylogenetics and Evolution 56: 675–689. Zhao LC, Li DY. 2008. Anatomically preserved seeds of Corylopsis (Hamamelidaceae) from the Miocene of Yunnan, China, and their phytogeographic implications. International Journal of Plant Sciences 169: 483–191. 161. CERCIDIPHYLLACEAE CARAMEL-TREE FAMILY Crane PR, Du Val A. 2013. Cercidiphyllum magnificum. Systematic placement and fossil history of Cercidiphyllum Siebold, Zuccarini, Cercidiphyllaceae. Curtis’s Botanical Magazine 30: 177–192. Crane PR, Stockey RA. 1986. Morphology and development of pistillate inflorescences in extant and fossil Cercidiphyllaceae. Annals of the Missouri Botanical Garden 73: 382–393. Feng YK, Wang XQ, Pan KY, Hong DY. 1998. A reevaluation of the systematic positions of the Cercidiphyllaceae and Daphniphyllaceae based on rbcL gene sequence analysis, with reference to the relationships in the “lower” Hamamelidae. Acta Phytotaxonomica Sinica 36: 411–422. Qi XS, Chen C, Comes HP, Sakagushi S, Liu YH, Tanaka N, Sakio H, Qiu YX. 2012. Molecular data and ecological niche modelling reveal a highly dynamic evolutionary history of the East Asian Tertiary relict Cercidiphyllum (Cercidiphyllaceae). New Phytologist 196: 617–630. Swamy BGL, Bailey IW. 1949. The morphology and relationships of Cercidiphyllum. Journal of the Arnold Arboretum 60: 367–376. 162. DAPHNIPHYLLACEAE LAUREL-LEAF FAMILY Bhatnagar AK, Garg M. 1977. Affinities of Daphniphyllum — palynological approach. Phytomorphology 27: 92–97. Carlquist S. 1982. Wood anatomy of Daphniphyllaceae: ecological and phylogenetic considerations, review of pittosporalean families. Brittonia 34: 252–266. Feng YK, Wang XQ, Pan KY, Hong DY. 1998. A reevaluation of the systematic positions of the Cercidiphyllaceae and Daphniphyllaceae based on rbcL gene sequence analysis, with reference to the relationships in the “lower” Hamamelidae. Acta Phytotaxonomica Sinica 36: 411–422. Huang TC. 1965–1966. Monograph of Daphniphyllum. Taiwania 11: 57–98, 12: 137–234. Zhang ZY, Lu AM. 1989. On the systematic position of Daphniphyllaceae. Acta Phytotaxonomica Sinica 27: 17–26.

FURTHER READING 163. ITEACEAE SWEETSPIRE FAMILY Bohm BA, Chalmers G, Bhat UG. 1988. Flavonoids and the relationships of Itea to the Saxifragales. Phytochemistry 27: 2651–2653. Bohm BA, Yang IY, Page JE, Soltis DS. 1999. Flavonoids, DNA and relationships of Itea and Pterostemon. Biochemical Systematics and Ecology 27: 79–83. Hermsen EJ, Gandolfo MA, Nixon KC, Crepet WL. 2003. Divisestylus gen. nov. (aff. Iteaceae), a fossil saxifrage from the late Cretaceous of New Jersey, USA. American Journal of Botany 90: 1373–1383. Petrov S, Drazheva-Stamatova T. 1973. Itea L. fossil pollen in Tertiary sediments of Europe and North America. Comptes rendus de l’Académie bulgare des Sciences 26: 811–814. Wilkinson HP. 1994. Leaf and twig anatomy of the Pterostemonaceae (Engl.) Small: ecological and systematic features. Botanical Journal of the Linnean Society 115: 115–131. 164. GROSSULARIACEAE GOOSEBERRY FAMILY Bate-Smith EC. 1976. Chemistry and taxonomy of Ribes. Biochemical Systematics and Ecology 4: 13–23. Keep E. 1962. Interspecific hybridisation in Ribes. Genetica 33: 1–23. Senters AE, Soltis DE. 2003. Phylogenetic relationships in Ribes (Grossulariaceae) inferred from ITS sequence data. Taxon 52: 51–66. Stern WL, Sweitzer EM, Phipps RE. 1970. Comparative anatomy and systematics of woody Saxifragaceae. Ribes. Botanical Journal of the Linnean Society 63, suppl. 1: 215–237. Weigend M, Motley T, Mohr O. 2002. Phylogeny and classification in the genus Ribes (Grossulariaceae) based on 5S-NTS sequences and morphological and anatomical data. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 124: 163–182. 165. SAXIFRAGACEAE SAXIFRAGE FAMILY Fay MF, Rankou H, Wilford R. 2009. Saxifraga granulata. Saxifragaceae. Curtis’s Botanical Magazine 26: 74–85. Ferguson IK, Webb DA. 1970. Pollen morphology in the genus Saxifraga and its taxonomic significance. Botanical Journal of the Linnean Society 63: 295–311. Gandolfo MA, Nixon KC, Crepet WL. 1998. Tylerianthus crossmanensis gen. et sp. nov. (aff. Hydrangeaceae) from the Upper Cretaceous of New Jersey. American Journal of Botany 85: 376–386. Gornall RJ. 1987. Foliar crystals in Saxifraga and segregate genera (Saxifragaceae). Nordic Journal of Botany 7: 233–238. Gornall RJ, Bohm BA. 1985. A monograph of Boykinia, Peltoboykinia, Bolandra, and Suksdorfia (Saxifragaceae). Botanical Journal of the Linnean Society 90: 1–71. McGregor M. 2008. Saxifrages. Timber Press, Portland. Morgan DR, Soltis DE. 1993. Phylogenetic relationships among Saxifragaceae sensu lato based on rbcL sequence data. Annals of the Missouri Botanical Garden 80: 631–660. Okuyama Y, Pellmyr O, Kato M. 2008. Parallel floral adaptations to pollination by fungus gnats within the genus Mitella (Saxifragaceae). Molecular Phylogenetics and Evolution 46: 560–575. Soltis DE, Morgan DR, Grable A, Soltis PS, Kuzoff

RK. 1993. Molecular systematics of Saxifragaceae sensu stricto. American Journal of Botany 80: 1056–1081. Soltis DE, Kuzoff RK, Gornall R, Ferguson K. 1996. matK and rbcL gene sequence data indicate that Saxifraga (Saxifragaceae) is polyphyletic. American Journal of Botany 83: 371–382. Soltis DE, Kuzoff RK. 1995. Discordance between molecular and chloroplast phylogenies in the Heuchera group (Saxifragaceae). Evolution 49: 727–742. Soltis DE, Kuzoff RK, Mort ME, Zanis M, Fishbein M, Hufford L, Koontz J, Arroyo MK. 2001. Elucidating deep-level phylogenetic relationships in Saxifragaceae using sequences for six chloroplastic and nuclear DNA regions. Annals of the Missouri Botanical Garden 88: 669–693. Soltis DE, Tago-Nakazawa M, Ziang QY, Kawano S, Murat J, Wakabayashi M. 2001. Phylogenetic relationships and evolution in Chrysosplenium (Saxifragaceae) based on matK sequence data. American Journal of Botany 88: 883–893. Xiang CL, Gitzendanner MA, Soltis DE, Peng H, Lei LG. 2012. Phylogenetic placement of the enigmatic and critically endangered genus Saniculiphyllum (Saxifragaceae) inferred from combined analysis of plastid and nuclear DNA sequences. Molecular Phylogenetics and Evolution 64: 357–367. Yeo PF. 1966. A revision of the genus Bergenia Moench (Saxifragaceae). Kew Bulletin 26: 113–148. 166. CRASSULACEAE STONECROP FAMILY Eggli U (ed.). 2003. Illustrated handbook of succulent plants, IV. Crassulaceae. Springer, Berlin. Gehrig H, Gauβmann O, Marx H, Schwarzott D, Kluge M. 2001. Molecular phylogeny of the genus Kalanchoë (Crassulaceae) inferred from nucleotide sequences of the ITS-1 and ITS-2 regions. Plant Science 160: 827–835. ‘t Hart H, Eggli U (eds). 1995. Evolution and systematics of the Crassulaceae. Backhuys, Leiden. ‘t Hart H, Eggli U. 1998. Cytotaxonomic studies in Rosularia (Crassulaceae). Botanica Helvetica 98: 223–234. ‘t Hart H, Koek-Noorman J. 1989. The origin of the woody Sedoideae (Crassulaceae). Taxon 38: 535–544. Jorgensen TH, Olesen JM. 2001. Adaptive radiation of island plants: evidence from Aeonium (Crassulaceae) of the Canary Islands. Perspectives in Plant Ecology, Evolution and Systematics 4: 29–42. Mayuzumi S, Ohba H. 2004. The phylogenetic position of eastern Asian Sedoideae (Crassulaceae) inferred from chloroplast and nuclear DNA sequences. Systematic Botany 29: 587–598. Mes THM, ‘t Hart H. 1996. The evolution of growthforms in the Macaronesian genus Aeonium (Crassulaceae) inferred from chloroplast RFLPs and morphology. Molecular Ecology 5: 351–363. Mort ME, Soltis DE, Soltis PS, Francisco-Ortega J, Santos-Guerra A. 2001. Phylogenetic relationships and evolution of Crassulaceae inferred from matK sequence data. American Journal of Botany 88: 76–91. Mort ME, O’Leary TR, Carillo-Reyes P, Nowell T, Archibald JK, Randle CP. 2010. Phylogeny and evolution of Crassulaceae: past, present and future. Schumannia 6: 69–86. Ogburn RM, Edwards EJ. 2010. The ecological wateruse stategies of succulent plants. Advances in Botanical Research 55: 179–255.

Thiede J, Eggli U. 2006. Crassulaceae. In: Kubitzki K (ed.) The families and genera of vascular plants 9: 83–118. Springer, Berlin. Van Ham RCHJ, ‘t Hart H. 1998. Phylogenetc relationships in the Crassulaceae inferred from chloroplast restriction-site variation. American Journal of Botany 85: 123–134. Winter K, Smith JAC. 1996. Crassulacean acid metabolism. Springer, Berlin. Yost JM, Bontrager M, McCabe SW, Burton D, Simpson MG, Kay KM, Ritter M. 2014. Phylogenetic relationships and evolution in Dudleya (Crassulaceae). Systematic Botany 38: 1096–1104. 167. APHANOPETALACEAE GUM-VINE FAMILY Bensel CR, Palser BF. 1975. Floral anatomy in the Saxifragaceae sensu lato II. Saxifragoideae and Iteoideae. American Journal of Botany 62: 661–675. Dickison WC, Hils HM, Lucansky TW, Stern WL. 1994. Comparative anatomy and systematics of woody Saxifragaceae. Aphanopetalum Endl. Botanical Journal of the Linnean Society 114: 167–182. 168. TETRACARPAEACEAE DELICATELAUREL FAMILY Hils HM, Dickison WC, Lucansky TW, Stern WL. 1988. Comparative anatomy and systematics of woody Saxifragaceae: Tetracarpaea. American Journal of Botany 75: 1687–1700. 169. PENTHORACEAE DITCH-STONECROP FAMILY Baldwin JT, Speese BM. 1951. Penthorum: its chromosomes. Rhodora 53: 89–91. Haskins ML, Hayden WJ. 1987. Anatomy and affinities of Penthorum. American Journal of Botany 74: 164–177. Ikeda H, Itoh K. 2001. Germination and water dispersal of seeds from a threatened plant species Penthorum chinense. Ecological Research 16: 99–106. Morgan DR, Soltis DE. 1993. Phylogenetic relationships among Saxifragaceae sensu lato based on rbcL sequence data. Annals of the Missouri Botanical Garden 80: 631–660. Soltis DE, Bohm BA. 1982. Flavonoids of Penthorum sedoides. Biochemical Systematics and Ecology 10: 221–224. 170. HALORAGACEAE WATER-MILFOIL FAMILY Hernández-Castillo GR, Cevallos-Ferriz SRG. 1999. Reproductive and vegetative organs with affinities to Haloragaceae from the Upper Cretaceous Huepac chert locality of Sonora, Mexico. American Journal of Botany 86: 1717–1734. Moody ML, Les DH. 2007. Phylogenetic systematics and character evolution in the angiosperm family Haloragaceae. American Journal of Botany 94: 2005–2025. Moody ML, Les DH. 2010. Systematics of the aquatic angiosperm genus Myriophyllum (Haloragaceae). Systematic Botany 35: 121–139. Orchard AE. 1975. Taxonomic revisions in the family Haloragaceae. I. The genera Haloragis, Haloragodendron, Glischrocaryon, Meziella and Gonocarpus. Bulletin of the Auckland Institute and Museum 10: 1–293.

Plants of the World

703

FURTHER READING Orchard AE. 1985. Myriophyllum (Haloragaceae) in Australasia. II. The Australian species. Brunonia 8: 173–291. Orchard AE, Keighery GJ. 1993. The status, ecology and relationships of Meziella (Haloragaceae). Nuytsia 9: 111–117. Praglowski J. 1970. The pollen morphology of the Haloragaceae with reference to taxonomy. Grana 10: 159–239. 171. CYNOMORIACEAE TARTHUTH FAMILY Heide-Jørgensen HS. 2008. Parasitic flowering plants. Brill, Leiden. Lebling RW. 2003. The treasure of tarthuth. Saudi Aramco World 54(2): 12–17. Nickrent DL, Der JP, Anderson FE. 2005. Discovery of the photosynthetic relatives of the “Maltese mushroom” Cynomorium. BMC Evolutionary Biology 5: 38. Zhang ZH, Li CQ, Li J. 2009. Phylogenetic placement of Cynomorium in Rosales inferred from sequences of the inverted repeat region of the chloroplast genome. Journal of Systematics and Evolution 47: 297–304. 172. VITACEAE GRAPEVINE FAMILY Chen I, Manchester SR. 2011. Seed morphology of Vitaceae. International Journal of Plant Sciences 172: 1–35. Chen P, Chen L, Wen J. 2011. The first phylogenetic analysis of Tetrastigma (Miq.) Planch., the host of Rafflesiaceae. Taxon 60: 499–512. Davis CC, Wurdack KJ. 2004. Host-to-parasite gene transfer in flowering plants: phylogenetic evidence from Malpighiales. Science 305: 676–678. Ingrouille MJ, Chase MW, Fay MF, Bowman D, Van der Bank M, Bruijn ADE. 2002. Systematics of Vitaceae from the viewpoint of plastid rbcL sequence data. Botanical Journal of the Linnean Society 138: 421–432. Kerrigde G, Gackle A. 2005. Vines for wines. CSIRO, Collingwood. Lombardi JA. 2000. Vitaceae. Flora Neotropica Monograph 80. New York Botanic Garden, New York. Lu L, Wang W, Chen Z, Wen J. 2013. Phylogeny of the non-monophyletic Cayratia Juss. (Vitaceae) and implications for character evolution and biogeography. Molecular Phylogenetics and Evolution 68: 502–515. Ren H, Lu LM, Soejima A, Luke Q, Zhang DX, Chen ZD, Wen J. 2011. Phylogenetic analysis of the grape family (Vitaceae) based on noncoding plastid trnCpetN, trnH-psbA, and trnL-F sequences. Taxon 60: 629–637. Ridsdale CE. 1974. A revision of the family Leeaceae. Blumea 22: 57–100. Terral JF, Tabard E, Bouby L, Ivorra S, Pastor T, Figueiral I, Picq S, Chevance J-B, Jung C, Fabre L, Tardy C, Compan M, Bacilieri R, Lacombe R, This P. 2010. Evolution and history of grapevine (Vitis vinifera) under domestication: new morphometric perspectives to understand seed domestication syndrome and reveal origins of ancient European cultivars. Annals of Botany 105: 433–455. Trias-Blasi A, Parnell JAN, Hodkinson TR. 2012. Multi-gene phylogenetic analysis of the grape family (Vitaceae). Systematic Botany 37: 941–950. Tröndle D, Schröder S, Kassemeyer H-H, Kiefer C, Koch MA, Nick P. 2010. Molecular phylogeny of the genus Vitis (Vitaceae) based on plastid markers. American Journal of Botany 97: 1168–1178.

704

Christenhusz, Fay & Chase

Wheeler EA, LaPasha CA. 1994. Woods of the Vitaceae — fossil and modern. Review of Palaeobotany and Palynology 80: 175–207. 173. KRAMERIACEAE RATANY FAMILY Milby TH. 1971. Floral anatomy of Krameria lanceolata. American Journal of Botany 58: 569–576. Musselman LJ. 1975. Parasitism and haustorial structure in Krameria lanceolata (Krameriaceae). A preliminary study. Phytomorphology 25: 416–422. Sheahan MC, Chase MW. 1996. A phylogenetic analysis of Zygophyllaceae R.Br. based on morphological, anatomical and rbcL DNA sequence data. Botanical Journal of the Linnean Society 122: 279–300. Simpson BB. 1989. Krameriaceae. Flora Neotropica Monograph 49. New York Botanical Garden. Simpson BB. 1991. The past and present uses of rhatany (Krameria, Krameriaceae). Economic Botany 45: 397–409. Simpson BB, Neff JL, Seigler D. 1977. Krameria, free fatty acids and oil-collecting bees. Nature 267: 150–151. Simpson BB, Weeks A, Helfgott DM, Larkin LL. 2004. Species relationships in Krameria (Krameriaceae) based on ITS sequences and morphology: implications for character utility and biogeography. Systematic Botany 29: 97–108. 174. ZYGOPHYLLACEAE TWINLEAF FAMILY Beier BA. 2005. A revision of the desert shrub Fagonia (Zygophyllaceae). Systematics and Biodiversity 3: 221–263. Beier BA, Chase MW, Thulin M. 2003. Phylogenetic relationships and taxonomy of subfamily Zygophylloideae (Zygophyllaceae) based on molecular and morphological data. Plant Systematics and Evolution 240: 11–39. Bellstedt DU, Van Zyl L, Marais EM, Bytebier B, De Villiers CA, Makwarela AM, Dreyer LL. 2008. Phylogenetic relationships, character evolution and biogeography of southern African members of Zygophyllum (Zygophyllaceae) based on three plastid regions. Molecular Phylogenetics and Evolution 47: 932–949. Lia VV, Confalonieri VA, Comas CI, Hunziker JH. 2001. Molecular phylogeny of Larrea and its allies (Zygophyllaceae): reticulate evolution and the probable time of creosote bush arrival in North America. Molecular Phylogenetics and Evolution 21: 309–320. Palacios RA, Hunziker JH. 1984. Revisión taxonómica del género Bulnesia (Zygophyllaceae). Darwiniana 25: 299–320. Porter D. 1969. The genus Kallstroemia (Zygophyllaceae). Contributions from the Gray Herbarium 198: 41–153. Sands MJS. 2001. The desert dates and its relatives: a revision of the genus Balanites. Kew Bulletin 56: 1–128. Sheahan MC, Chase MW. 1996. A phylogenetic analysis of Zygophyllaceae R.Br. based on morphological, anatomical and rbcL DNA sequence data. Botanical Journal of the Linnean Society 122: 279–300. Sheahan MC, Chase MW. 2000. Phylogenetic relationships within Zygophyllaceae based on DNA sequences of three plastid regions, with special emphasis on Zygophylloideae. Systematic Botany 25: 371–384. Sheahan MC, Cutler DC. 1993. Contribution of vegetative anatomy to the systematics of the

Zygophyllaceae R. Br. Botanical Journal of the Linnean Society 113: 227–262. Yang TW, Yan YA, Xiong Z. 2000. Paternal inheritance of chloroplast DNA in interspecific hybrids in the genus Larrea (Zygophyllaceae). American Journal of Botany 87: 1452–1458. 175. QUILLAJACEAE SOAPBARK-TREE FAMILY Bate-Smith EC. 1965. Investigation of the chemistry and taxonomy of subtribe Quillajeae of the Rosaceae using comparisons of fresh and herbarium material. Phytochemistry 4: 535–539. Bello MA, Hawkins JA, Rudall PJ. 2007. Floral morphology and development in Quillajaceae and Surianaceae (Fabales), the species-poor relatives of Leguminosae and Polygalaceae. Annals of Botany 100: 1491–1505. Hooker JD. 1897. Quillaja saponaria. Curtis’s Botanical Magazine III, 53: t. 7568. 176. FABACEAE PEA FAMILY Brown GK, Murphy DJ, Kidman J, Ladiges PY. 2012. Phylogenetic connections of phyllodinous species of Acacia outside Australia are explained by geological history and human-mediated dispersal. Australian Systematic Botany 25: 390–403. Bruneau A, Forest F, Herendeen PS, Klitgaard BB, Lewis GP. 2001. Phylogenetic relationships in Caesalpinioideae (Leguminosae) as inferred from chloroplast trnL intron sequences. Systematic Botany 26: 487–514. Bruneau A, Mercure M, Lewis GP, Herendeen PS. 2008. Phylogenetic patterns and diversification in the caesalpinioid legumes. Botany 86: 697–718. Cecil CO. 2005. Gum arabic. Saudi Aramco World 56(2): 36–39. Friend SA, Quandt D, Tallury SP, Stalker HT, Hilu KW. 2010. Species, genomes, and section relationships in the genus Arachis (Fabaceae): a molecular phylogeny. Plant Systematics and Evolution 290: 185–199. Lavin M, Schrire BP, Lewis G, Pennington RT, Delgado-Salinas A, Thulin M, Hughes CE, Beyra Matos A, Wojciechowski MF. 2004. Metacommunity process rather than continental tectonic history better explains geographically structured phylogenies in legumes. Philosophical Transactions of the Royal Society of London B, 359: 1509–1522. Lavin M, Herendeen PS, Wojciechowski MF. 2005. Evolutionary rates analysis of Leguminosae implicates a rapid diversification of lineages during the Tertiary emergence. Systematic Biology 54: 575–594. Legume Phylogeny Working Group. 2013. Legume phylogeny and classification in the 21st century: progress, prospects and lessons for other speciesrich clades. Taxon 62: 217–248. Legume Phylogeny Working Group. 2013. Towards a new classification system for legumes: progress report from the 6 th International Legume Conference. South African Journal of Botany 89: 3–9. Lewis GP, Schrire B, Mackinder B, Lock M. 2005. Legumes of the World. Royal Botanic Gardens, Kew. Li QG, Li Z, Li C, Dunwell JM, Zhang YM. 2013. Comparative genomics suggests that an ancestral polyploidy event leads to enhanced root nodule symbiosis in the Papilionoideae. Molecular Biology and Evolution 30: 2602–2611. Maslin B, Miller JT, Seigler DS. 2003. Overview

FURTHER READING of the generic status of Acacia (Leguminosae: Mimosoideae). Australian Systematic Botany 16: 1–18. Orchard AE, Maslin BR. 2003. Proposal to conserve the name Acacia (Leguminosae: Mimosoideae) with a conserved type. Taxon 52: 362–363. Pennington TD (ed.). 1997. The genus Inga: botany. Royal Botanic Gardens, Kew. Pennington TD, Fernandes EC (eds). 1998. The genus Inga: utilization. Royal Botanic Gardens, Kew. Schrire BD, Lavin M, Lewis GP. 2005. Global distribution patterns of the Leguminosae: insights from recent phylogenies. Biologiske Skrifter 55: 375–422. Shoemaker RC, Schlueter J, Doyle JJ. 2006. Paleopolyploidy and gene duplication in soybean and other legumes. Current Opinions in Plant Biology 9: 104–109. Small E. 2011. Alfalfa and relatives. Evolution and classification of Medicago. NRC Research Press, Ottawa. Smith ME, Henkel TW, Aime MC, Fremier AK, Vilgalys R. 2011. Ectomycorrhizal fungi diversity and community structure on three co-occurring leguminous canopy tree species in a Neotropical rainforest. New Phytologist 192: 699–712. Weeden NF. 2007. Genetic changes accompanying the domestication of Pisum sativum: is there a common basis to the ‘domestication syndrome’ in legumes? Annals of Botany 100: 1017–1025. Note: There are numerous publications addressing a diversity of aspects relating to this family. A good general bibliography and additional information about many species is available on LOWO (Legumes of the World Online): http://www.kew. org/science-conservation/research-data/resources/ legumes-of-the-world (accessed April 2014). 177. SURIANACEAE BAY-CEDAR FAMILY Behnke HD, Kiritsis U, Patrick SJ, Kenneally KF. 1996. Form-Pfs plastids, stem anatomy and systematic affinities of Stylobasium Desf. (Stylobasiaceae). A contribution to the knowledge of sieve-element plastids in the Rutales and Sapindales. Botanica Acta 109: 346–359. Bello MA, Hawkins JA, Rudall PJ. 2007. Floral morphology and development in Quillajaceae and Surianaceae (Fabales), the species-poor relatives of Leguminosae and Polygalaceae. Annals of Botany 100: 1491–1505. Carlquist S. 1978. Wood anatomy and relationships of Bataceae, Gyrostemonaceae, and Stylobasiaceae. Allertonia 1: 297–330. Crayn DM, Fernando ES, Gadek PA, Quinn CJ. 1995. A reassessment of the familial affinity of the Mexican genus Recchia Moçiño and Sessé ex DC. Brittonia 47: 397–402. Gutzwiller MA. 1961. Die phylogenetische Stellung von Suriana maritima L. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 81: 1–49. Weberling F, Lörcher H, Böhnke F. 1980. Die Stipeln der Irvingioideae und Recchioideae und ihre systematische Wertung nebst Bemerkungen zur Holzanatomie und Palynologie. Plant Systematics and Evolution 133: 261–283. 178. POLYGALACEAE MILKWORT FAMILY Abbott JR. 2011. Notes on the disintegration of Polygala (Polygalaceae), with four new genera for the Flora of North America. Journal of the Botanical Research Institute of Texas 5: 125–137.

Bello MA, Hawkins JA, Rudall PJ. 2010. Floral ontogeny in Polygalaceae and its bearing on the homologies of keeled f lowers in Fabales. International Journal of Plant Sciences 171: 482–498. Brantjes NBM, Van der Pijl L. 1980. Pollination mechanisms in Polygalaceae. Acta Botanica Neerlandica 29: 56–57. Bridgewater S, Baas P. 1982. Wood anatomy of Xanthophyllum Roxb. IAWA Bulletin new series 3: 115–125. Eriksen B. 1993. Floral anatomy and morphology in the Polygalaceae. Plant Systematics and Evolution 186: 17–32. Eriksen B. 1993. Phylogeny of the Polygalaceae and its taxonomic implications. Plant Systematics and Evolution 186: 33–55. Forest F, Manning JC. 2006. Evidence for inclusion of South African endemic Nylandtia in Muraltia (Polygalaceae). Systematic Botany 31: 525–532. Forest F, Chase MW, Persson C, Crane PR, Hawkins JA. 2007. The role of biotic and abiotic factors in evolution of ant dispersal in the milkwort family (Polygalaceae). Evolution 61: 1675–1694. Freire Fierro A, Welling JC. (eds). 2001 onwards. Polygalaceae Web Site: http://www.joethejuggler. com/Polygalaceae/ (accessed April 2014). Prenner G. 2004. Floral development in Polygala myrtifolia (Polygalaceae) and its similarities with Leguminosae. Plant Systematics and Evolution 249: 67–76. 179. ROSACEAE ROSE FAMILY Bortiri E, E, Oh S, Jiang J, Baggett S, Granger A, Weeks C, Buckingham M, Potter D, Parfitt D. 2001. Phylogeny and systematics of Prunus (Rosaceae) as determined by sequences analysis of ITS and the chloroplast trnL-trnF spacer DNA. Systematic Botany 26: 797–807. Campbell CS, Donoghue MJ, Baldwin BG, Wojciechowski MF. 1995. Phylogenetic relationships in Maloideae (Rosaceae): evidence from sequences of the internal transcribed spacer of nuclear ribosomal DNA and its congruence with morphology. American Journal of Botany 82: 903–918. Christenhusz MJM, Väre H. 2012. New combinations in Potentilla (Rosaceae) for the Nordic Flora. Phytotaxa 57: 1–5. Dickinson TA, Lo E, Talent N. 2007. Polyploidy, reproductive biology, and Rosaceae: understanding evolution and making classifications. Plant Systematics and Evolution 266: 59–78. Dobes C, Paule J. 2010. A comprehensive chloroplast DNA-based phylogeny of the genus Potentilla (Rosaceae): implications for its geographic origin, phylogeography and generic circumscription. Molecular Phylogenetics and Evolution 56: 156–175. Eriksson T, Donoghue MJ, Hibbs MS. 1998. Phylogenetic analysis of Potentilla using DNA sequences of nuclear ribosomal internal transcribed spacers (ITS), and implications for the classification of Rosoideae (Rosaceae). Plant Systematics and Evolution 211: 155–179. Hancock JF. 1999. Strawberries. CABI publishing, Wallingford. Juniper BE, Mabberley DJ. 2006. The story of the apple. Timber Press, Portland. Kalkman C. 1988. The phylogeny of the Rosaceae. Botanical Journal of the Linnean Society 98: 37–59. Koopman WJM, Wissemann V, De Cock K, Van

Huylenbroeck J, De Riek J, Sabatino GJH, Visser D, Vosman B, Ritz CM, Maes B, Werlemark G, Nybom H, Debener T, Linde N, Smulders MJM. 2008. AFLP markers as a tool to reconstruct complex relationships: a case study in Rosa (Rosaceae). American Journal of Botany 95: 353–366. Lo EY, Donoghue MJ. 2012. Expanded phylogenetic dating analysis of the apples and their relatives (Pyreae, Rosaceae). Molecular Phylogenetics and Evolution 63: 230–243. Mabberley DJ. 2002. Potentilla and Fragaria (Rosaceae) reunited. Telopea 9: 793–801. Morgan DR, Soltis DE, Robertson KR. 1994. Systematic and evolutionary implications of rbcL sequence variation in Rosaceae. American Journal of Botany 81: 890–903. Potter D. Gao F, Bortiri PE, Oh S-H, Baggett S. 2002. Phylogenetic relationships in Rosaceae inferred from chloroplast matK and trnL-trnF nucleotide sequence data. Plant Systematics and Evolution 231: 77–89. Potter D, Eriksson T, Evans RC, Oh S, Smedmark JEE, Morgan DR, Kerr M, Robertson KR, Arsenault M, Dickinson TA, Campbell CS. 2007. Phylogeny and classification of Rosaceae. Plant Systematics and Evolution 266: 5–43. Robertson KR, Phipps JB, Rohrer JR. 1991. A synopsis of genera in Maloideae (Rosaceae). Systematic Botany 16: 376–394. Smedmark JEE. 2006. Recircumscription of Geum L. (Colurieae: Rosaceae). Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 126: 409–417. Soják J. 2008. Notes on Potentilla XXI. A new division of the tribe Potentilleae (Rosaceae) and notes on generic delimitation. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 127: 349–358. Vamosi JC, Dickinson TA. 2006. Polyploidy and diversification: a phylogenetic investigation in Rosaceae. International Journal of Plant Sciences 167: 349–358. Wallaart RAM. 1980. Distribution of sorbitol in Rosaceae. Phytochemistry 19: 989–1000. Zhang SD, Jin JJ, Chen SY, Chase MW, Soltis DE, Li HT, Yang JB, Li DZ, Yi TS. 2017. Diversification of Rosaceae since the late Cretaceous based on plastid phylogenomics. New Phytologist 214: 1355–1367. Zhang SY. 1992. Systematic wood anatomy of the Rosaceae. Blumea 37: 81–158. 180. BARBEYACEAE ELM-OLIVE FAMILY Dickison WC, Sweitzer EM. 1970. The morphology and relationship of Barbeya oleoides. American Journal of Botany 57: 468–476. Thulin M, Bremer B, Richardson J, Niklasson J. Fay MF, Chase MW. 1998. Family relationships of the enigmatic rosid genera Barbeya and Dirachma from the Horn of Africa region. Plant Systematics and Evolution 213: 103–119. Tobe H, Takahashi M. 1990. Trichome and pollen morphology of Barbeya (Barbeyaceae) and its relationships. Taxon 39: 561–567. 181. DIRACHMACEAE RACHMAN FAMILY Bazara’a M, Guarino L, Miller A, Obadi N. 1991. Dirachma socotrana — back from the brink? Oryx 25: 229–232. Boesewinkel FD, Bouman F. 1997. Ovules and seeds of Dirachma socotrana (Dirachmaceae). Plant Systematics and Evolution 205: 195–204.

Plants of the World

705

FURTHER READING Link DA. 1991. Dirachma somalensis D.A. Link sp. nov. A new species of a remarkable and highly endangered monogeneric family. Bulletin du Jardin Botanique National de Belgique 61: 3–13. Thulin M, Bremer B, Richardson J, Niklasson J, Fay MF, Chase MW. 1998. Family relationships of the enigmatic rosid genera Barbeya and Dirachma from the Horn of Africa region. Plant Systematics and Evolution 213: 103–119. 182. ELAEAGNACEAE OLEASTER FAMILY Bartish IV, Jeppson N, Nybom H, Swenson U. 2002. Phylogeny of Hippophaë (Rhamnaceae) inferred from parsimony analysis of chloroplast DNA and morphology. Systematic Botany 27: 41–54. Gardner IC. 1958. Nitrogen fixation in Elaeagnus root nodules. Nature 181: 717–718. Jansen S, Piesschaert F, Smets E. 2000. Wood anatomy of Elaeagnaceae, with comments on vestured pits, helical thickenings, and systematic relationships. American Journal of Botany 87: 20–28. Xu M. 1994. The medical research and exploitation of sea buckthorn. Hippophaë 7: 32–34. 183. RHAMNACEAE BUCKTHORN FAMILY Fay MF, Lledó MD, Richardson JE, Rye BL, Hopper SD. 2001. Molecular data confirm the affinities of the southwest Australian endemic Granitites with Alphitonia (Rhamnaceae). Kew Bulletin 56: 669–675. Pillans NS. 1942. The genus Phylica Linn. Journal of South African Botany 8: 1–164. Richardson JE, Fay MF, Cronk QCB, Bowman D, Chase MW. 2000. A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. American Journal of Botany 87: 1309–1324. Richardson JE, Fay MF, Cronk QCB, Bowman D, Chase MW. A revision of the tribal classification of Rhamnaceae. Kew Bulletin 55: 311–340. 184. ULMACEAE ELM FAMILY Buisman C. 1928. De oorzaak van de iepenziekte. Tijdschrift der Nederlandsche Heidemaatschappij 40: 338–345. Holmes FW, Heybroek HM. 1990. Dutch elm disease: the early papers: selected works of seven Dutch women phytopathologists. American Phytopathological Society Press, St. Paul. Sweitzer EM. 1971. Comparative anatomy of the Ulmaceae. Journal of the Arnold Arboretum 52: 523–585. Todzia CA. 1989. A revision of Ampelocera (Ulmaceae). Annals of the Missouri Botanical Garden 76: 1087–1102. Ueda K, Kosuge K, Tobe H. 1997. A molecular phylogeny of Celtidaceae and Ulmaceae (Urticales) based on rbcL nucleotide sequences. Journal of Plant Research 110: 171–178. Wang Q, Manchester SR, Li C, Geng B. 2010. Fruits and leaves of Ulmus from the Paleogene of Fushun, northeastern China. International Journal of Plant Sciences 171: 221–226. Wiegrefe SJ, Sytsma KJ, Guries RP. 1998. The Ulmaceae, one family or two? Evidence from chloroplast DNA restriction site mapping. Plant Systematics and Evolution 210: 249–270. Zavada M. 1983. Pollen morphology of Ulmaceae. Grana 22: 23–30. 185. CANNABACEAE HEMP FAMILY Appleby CA, Tjepkema JD, Trinick MJ. 1983. Hemoglobin in a nonleguminous plant, Parasponia:

706

Christenhusz, Fay & Chase

possible genetic origin and function in nitrogen fixation. Science 220: 951–953. Bourrie M. 2003. Hemp culture: a short history of the most misunderstood plants and its uses and abuses. Key Porter Books, Toronto. Burgess AH. 1964. Hops. Botany, cultivation, and utilization. Interscience Publishers, New York. Clarke RC. 1981. Marijuana botany. And/or Press, Berkeley. Godwin H. 1967. Pollen-analytic evidence for the cultivation of Cannabis in England. Review of Palaeobotany and Palynology 4: 71–80. Small ES, Cronquist A. 1976. A practical and natural taxonomy for Cannabis. Taxon 25: 405–435. Sytsma KJ, Morawetz J, Pires JC, Nepokroeff M, Conti E, Zjhra M, Hall JC, Chase MW. 2002. Urticalean rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89: 1531–1546. Yang MQ, Van Velzen R, Bakker FT, Sattarian A, Li DZ, Yi TS. 2013. Molecular phylogenetics and character evolution of Cannabaceae. Taxon 62: 473–485. 186. MORACEAE MULBERRY FAMILY Berg CC. 1973. Some remarks on the classification and differentiation of Moraceae. Mededeelingen van het Botanisch Museum en Herbarium van de Rijks Universiteit te Utrecht 386: 1–10. Berg CC. 1990. Reproduction and evolution in Ficus (Moraceae): traits connected with the adequate rearing of pollinators. Memoirs of the New York Botanical Garden 55: 169–185. Berg CC, Hijman MEE. 1999. The genus Dorstenia (Moraceae). Ilicifolia 2: 1–211. Chase MW, Thijs KW, Kamau P, Fay MF. 2013. Dorstenia christenhuszii (Moraceae), a new species from the Taita Hills, Kenya. Phytotaxa 81: 45–48. Clement WL, Weiblen GD. 2009. Morphological evolution in the mulberry family (Moraceae). Systematic Botany 34: 530–552. Corner EJH. 1962. The classification of Moraceae. Gardens’ Bulletin 19: 187–252. Cruaud A, Rønsted N, Chantarasuwan B, Chou LS, Clement WL, Couloux A, Cousins B, Genson G, Harrison RD, Hanson PE, Hossaert-McKey M, Jabbour-Zahab R, Jousselin E, Kerdelhué C, Kjellberg F, Lopez-Vaamonde C, Peebles J, Peng YQ, Pereira RAS, Schramm T, Ubaidillah R, Van Noort S, Weiblen GD, Yan DR, Yodpinyanee A, Libeskind-Hadas R, Cook JM, Rasplus JY, Savonlainen V. 2012. An extreme case of plantinsect codiversification: figs and fig-pollinating wasps. Systematic Biology 61: 1029–1047. Misiewicz TM, Zerega NC. 2012. Phylogeny, biogeography and character evolution of Dorstenia (Moraceae). Edinburgh Journal of Botany 69: 413–440. Taylor PE, Card G, House J, Dickinson MH, Flagan RC. 2006. High-speed pollen release in the white mulberry, Morus alba L. Sexual Plant Reproduction 19: 19–24. Zerega NJC, Clement WL, Datwyler SL, Weiblen GD. 2005. Biogeography and divergence times in the mulberry family (Moraceae). Molecular Phylogenetics and Evolution 37: 402–416. Zerega NJC, Supardi MNN, Motley TJ. 2010. Phylogeny and recircumscription of Artocarpeae (Moraceae) with a focus on Artocarpus. Systematic Botany 35: 766–782.

187. URTICACEAE NETTLE FAMILY Bassett IJ, Crompton CW, Woodland DW. 1974. The family Urticaceae in Canada. Canadian Journal of Botany 52: 503–516. Berg CC. 1977. Urticales, their differentiation and systematic position. Plant Systematics and Evolution, supplement 1: 349–374. Berg CC. 1978. Cecropiaceae, a new family of the Urticales. Taxon 27 : 39–44. Collinson ME. 1989. The fossil history of Moraceae, Urticaceae (including Cecropiaceae), and Cannabaceae. In: Crane PR, Blackmore S (eds). Evolution, systematics, and fossil history of the Hamamelidae, vol 2. Clarendon Press, Oxford. Emmelin N, Feldberg W. 1947. The mechanism of the sting of the common nettle (Urtica urens). Journal of Physiology 106: 440–455. Friis I. 1983. A synopsis of Obetia (Urticaceae). Kew Bulletin 38: 221–228. Friis I. 1988. Distribution patterns and biological observations in the Urticaceae of Sub-Saharan Africa, Madagascar and the Mascarenes. Monographs in Systematic Botany from the Missouri Botanical Garden 25: 527–543. Hadiah JT, Conn BJ, Quinn CJ. 2008. Infrafamilial phylogeny of Urticaceae, using chloroplast sequence data. Australian Systematic Botany 21: 375–385. Kim CK, Deng T, Chase MW, Zhang DG, Nie ZL, Hang S. 2015. Generic phylogeny and character evolution in Urticeae (Urticaceae) inferred from nuclear and plastid regions. Taxon 64: 65–78. Miller NG. 1971. The genera of the Urticaceae in the southeastern United States. Journal of the Arnold Arboretum 52: 40–68. Monro AK. 2006. The revision of species-rich genera: a phylogenetic framework for the strategic revision of Pilea (Urticaceae) based on cpDNA, nrDNA, and morphology. American Journal of Botany 93: 426–441. Rickson FR. 1976. Anatomical development of the leaf trichilium and Mullerian bodies of Cecropia peltata L. American Journal of Botany 64: 1266–1271. Schöter H, Winkler H. 1935. Monographie der Gattung Elatostema s.l. Allgemeiner Teil. Feddes Repertorium Beihefte 83, 1: 1–56. Sorsa P, Huttunen P. 1975. On the pollen morphology of the Urticaceae. Annales Botanici Fennici 12: 165–182. Sytsma KJ, Morawetz J, Pires JC, Nepokroeff M, Conti E, Zjhra M, Hall JC, Chase MW. 2002. Urticalean rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. American Journal of Botany 89: 1531–1546. Wu ZY, Monro A, Milne RI, Wang H, Yi TS, Liu J, Li DZ. 2013. Molecular phylogeny of the nettle family (Urticaceae) inferred from multiple loci of three genomes and extensive generic sampling. Molecular Phylogenetics and Evolution 69: 814–827. 188. NOTHOFAGACEAE ROBLE FAMILY Cook LG, Crisp MD. 2005. Not so ancient: the extant crown group of Nothofagus represents a postGondwanan radiation. Proceedings of the Royal Society B 272: 2535–2544. Heenan PG, Smissen RB. 2013. Revised circumscription of Nothofagus and recognition of the segregate genera Fuscospora, Lophozonia, and Trisyngyne (Nothofagaceae). Phytotaxa 146: 1–31.

FURTHER READING Jordan GJ, Hill RS. 1999. The phylogenetic affinities of Nothofagus (Nothofagaceae) leaf fossils based on combined molecular and morphological data. International Journal of Plant Sciences 160: 1177–1188. Knapp M, Stockler K, Havell D, Delsuc F, Sebastiani F, Lockhart PJ. 2005. Relaxed molecular clock provides evidence for long-distance dispersal of Nothofagus (southern beech). PLoS Biology 3: 38–43. Manos PS. 1997. Systematics of Nothofagus (Nothofagaceae) based on rDNA spacer sequences (ITS): taxonomic congruence with morphology and plastid sequences. American Journal of Botany 84: 1137–1155. Peterson KR, Pfister DH, Bell CD. 2010. Cophylogeny and biogeography of the fungal parasite Cyttaria and its host Nothofagus, southern beech. Mycologia 102: 1417–1425. Sauquet HE, Ho SYW, Gandolfo MA, Jordan GJ, Wilf P, Cantrill DJ, Bayly MJ, Bromham L, Brown GK, Carpenter RJ, Lee DM, Murphy DJ, Sniderman JMK, Udovicic F. 2012. Testing the impact of calibration on molecular divergence times using a fossil-rich group: the case of Nothofagus (Fagales). Systematic Biology 61: 289–313. Van Steenis CGGJ. 1971. Nothofagus, key genus of plant geography, in time and space, living and fossil, ecology and phylogeny. Blumea 19: 65–98. 189. FAGACEAE BEECH FAMILY Axelrod DI. 1983. Biogeography of oaks in the ArctoTertiary province. Annals of the Missouri Botanical Garden 70: 629–657. Denk T, Grimsson F, Zetter R. 2012. Fagaceae from the early Oligocene of Central Europe: persisting New World and emerging Old World biogeographic links. Review of Palaeobotany and Palynology 169: 7–20. Govaerts R, Frodin DG. 1998. World checklist and bibliography of Fagales (Betulaceae, Corylaceae, Fagaceae and Ticodendraceae). Royal Botanic Gardens, Kew. Koening WD, Knops JMH. 2005. The mystery of masting in trees. BioScience 93: 340–347. Manos PS, Stanford AM. 2001. The historical biogeography of Fagaceae: tracking the Tertiary history of temperate and subtropical forests of the Northern Hemisphere. International Journal of Plant Sciences 162: S77–S93. Manos PS, Doyle JJ, Nixon KC. 1999. Phylogeny, biogeography, and processes of molecular differentiation of Quercus subgenus Quercus (Fagaceae). Molecular Phylogenetics and Evolution 12: 333–349. Manos PS, Zhou ZK, Cannon CH. 2001. Systematics of Fagaceae: phylogenetic tests of reproductive trait evolution. International Journal of Plant Sciences 162: 1361–1379. Manos PS, Cannon CH, Oh SH. 2008. Phylogenetic relationships and taxonomic status of the paleoendemic Fagaceae of western North America: recognition of a new genus, Notholithocarpus. Madroño 55: 181–190. Oh SH, Manos PS. 2008. Molecular phylogenetics and cupule evolution in Fagaceae as inferred from nuclear CRABS CLAW sequences. Taxon 57: 434–451. 190. MYRICACEAE BAYBERRY FAMILY Bond G. 1951. The fixation of nitrogen associated with the root nodules of Myrica gale L., with special

reference to its pH relation and ecological significance. Annals of Botany (London) II 15: 447–459. Carlquist S. 2002. Wood and bark anatomy of Myricaceae: relationships, generic definitions, and ecological interpretations. Aliso 21: 7–29. Chevalier A. 1901. Monographie des Myricacées; anatomie et histologie, organographie, classification et déscription des espèces, distribution géographique. Mémoires de la Société des Sciences Naturelles de Cherbourg 32: 85–340. Hurd TM, Schwintzer CR. 1997. Formation of cluster roots and mycorrhizal status of Comptonia pere– grina and Myrica pensylvanica in Maine, U.S.A. Physiologia Plantarum 99: 680–689. Liang XQ, Wilde V, Ferguson DK, Kvacek Z, Ablaev AG, Wang YF, Li CS. 2010. Comptonia naumannii (Myricaceae) from the early Miocene of Weichang, China, and the palaeobiogeographical implication of the genus. Review of Palaeobotany and Palynology 163: 52–63. Lloyd DG. 1981. The distribution of sex in Myrica gale. Plant Systematics and Evolution 138: 29–45. 191. JUGLANDACEAE WALNUT FAMILY Heimsch C, Wetmore RH. 1939. The significance of wood anatomy in taxonomy of the Juglandaceae. American Journal of Botany 26: 651–660. Manchester SR. 1987. The fossil history of the Juglandaceae. Monographs in Systematic Botany from the Missouri Botanical Garden 21: 1–137. Manning WE. 1960. The genus Juglans in South America and the West Indies. Brittonia 12: 1–26. Manning WE. 1978. The classification within the Juglandaceae. Annals of the Missouri Botanical Garden 65: 1058–1987. Manos PS, Soltis PS, Soltis DE, Manchester SR, Oh SH, Bell CD, Dilcher DL, Stone DE. 2007. Phylogeny of extant and fossil Juglandaceae inferred from the integration of molecular and morphological data sets. Systematic Biology 56: 412–430. Manos PS. Stone DE. 2001. Evolution, phylogeny and systematics of the Juglandaceae. Annals of the Missouri Botanical Garden 88: 231–269. Wing SL, Hickey LJ. 1984. The Platycarya perplex and the evolution of Juglandaceae. American Journal of Botany 71: 388–411. 192. CASUARINACEAE SHE-OAK FAMILY Campbell JD, Holden AM. 1984. Miocene casuarinacean fossils from Southland and Central Otago. New Zealand Journal of Botany 22: 159–167. Dilcher DL, Christophel DC, Bhagwandin Jr HO, Scriven LJ. 1990. Evolution of the Casuarinaceae: morphological comparisons of some extant species. American Journal of Botany 77: 338–355. Flores EM. 1980. Shoot vascular system and phyllotaxis of Casuarina (Casuarinaceae). American Journal of Botany 67: 131–140. Kershaw AP. 1970. Pollen morphological variation within the Casuarinaceae. Pollen and Spores 12: 145–161. Moseley MF. 1948. Comparative anatomy and phylogeny of the Casuarinaceae. Botanical Gazette 110: 231–280. Steane DA, Wilson KL, Hill RS. 2003. Using matK sequence data to unravel the phylogeny of Casuarinaceae. Molecular Phylogenetics and Evolution 28: 47–59. Swamy BGL. 1948. A contribution to the life history of Casuarina. Proceedings of the American Academy of Arts and Sciences 77: 1–32.

Torrey JG, Berg RH. 1988. Some morphological features for generic characterisation among the Casuarinaceae. American Journal of Botany 75: 864–874. 193. TICODENDRACEAE TICO-TREE FAMILY Carlquist S. 1991. Wood and bark anatomy of Ticodendron: comments on relationships. Annals of the Missouri Botanical Garden 78: 96–104. Feuer S. 1991. Pollen morphology and the systematic relationships of Ticodendron incognitum. Annals of the Missouri Botanical Garden 78: 143–151. Gómez-Laurito J, Gómez PL. 1989. Ticodendron: a new tree from Central America. Annals of the Missouri Botanical Garden 76: 1148–1151. Hammel B, Burger W. 1991. Neither oak nor alder, but nearly: the history of Ticodendraceae. Annals of the Missouri Botanical Garden 78: 89–95. Manchester SR. 2011. Fruits of Ticodendraceae (Fagales) from the Eocene of Europe and North America. International Journal of Plant Sciences 172: 1179–1187. Sogo A, Tobe H. 2008. Mode of pollen tube growth in pistils of Ticodendron incognitum (Ticodendraceae, Fagales) and the evolution of chalazogamy. Botanical Journal of the Linnean Society 157: 621–631. Tobe H. 1991. Reproductive morphology, anatomy, and relationships of Ticodendron. Annals of the Missouri Botanical Garden 78: 135–142. 194. BETULACEAE BIRCH FAMILY Ashburner K, McAllister H. 2013. The genus Betula: a taxonomic revision of birches. Royal Botanic Gardens, Kew. Chen ZD, Manchester SR, Sun HY. 1999. Phylogeny and evolution of Betulaceae as inferred from DNA sequences, morphology, and paleobotany. American Journal of Botany 86: 1168–1181. Forest F, Savolainen V, Chase MW, Lupia R, Bruneau A, Crane PR. 2005. Teasing apart molecular- versus fossil-based error estimates when dating phylogenetic trees: a case study in the birch family (Betulaceae). Systematic Botany 30: 118–133. Furlow JJ. 1979. The systematics of the American species of Alnus (Betulaceae). Rhodora 81: 1–121, 151–248. Grimm G, Renner SS. 2013. Harvesting Betulaceae sequences from GenBank to generate a new chronogram for the family. Botanical Journal of the Linnean Society 172: 465–477. Hall JW. 1952. The comparative anatomy and phylogeny of the Betulaceae. Botanical Gazette 113: 235–270. 195. APODANTHACEAE STEMSUCKER FAMILY Barkman TJ, McNeal JR, Lim S-H, Coat G, Croom HB, Young ND, dePamphilis CW. 2007. Mitochondrial DNA suggests at least 11 origins of parasitism in angiosperms and reveals genomic chimerism in parasitic plants. BMC Evolutionary Biology 7: 248. Bellot S, Renner SS. 2013. Pollination and mating systems of Apodanthaceae and the distribution of reproductive traits in parasitic angiosperms. American Journal of Botany 100: 1083–1094. Blarer A, Nickrent DL, Endress PK. 2004. Comparative floral structure and systematics in Apodanthaceae (Rafflesiales). Plant Systematics and Evolution 245: 119–142. Bouman F, Meijer W. 1994. Comparative structure of ovules and seeds in Rafflesiaceae. Plant Systematics and Evolution 193: 187–212.

Plants of the World

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FURTHER READING Filipowicz N, Renner SS. 2010. The worldwide holoparasitic Apodanthaceae confidently placed in the Cucurbitales by nuclear and mitochondrial gene trees. BMC Evolutionary Biology 10: 219. Gomes AL, Fernandes GW. 1994. Influence of parasitism by Pilostyles ingae (Rafflesiaceae) on its host plant, Mimosa naguirei (Leguminosae). Annals of Botany 74: 205–208. Kuijt J. 1969. The biology of parasitic flowering plants. University of California, Berkeley. Milanello do Amaral M, Ceccantini G. 2011. The endoparasite Pilostyles ulei (Apodanthaceae — Cucurbitales) influences wood structure in three host species of Mimosa. IAWA Journal 32: 1–13. 196. ANISOPHYLLEACEAE LEECHWOOD FAMILY Anderson JAR, Muller J. 1975. Palynological study of a Holocene peat and a Miocene coal deposit from NW Borneo. Review of Palaeobotany and Palynology 19: 291–351. Dahlgren RMT. 1988. Rhizophoraceae and Anisophylleaceae: summary statement, relationships. Annals of the Missouri Botanical Garden 75: 1259–1277. Matthews ML, Endress PK, Schönenberger J, Friis EM. 2001. A comparison of f loral structures of Anisophylleaceae and Cunoniaceae and the problem of their systematic position. Annals of Botany 88: 439–455. Raven PH, Tomlinson PB. 1988. RhizophoraceaeAnisophylleaceae: a symposium. Annals of the Missouri Botanical Garden 75: 1258. (see also papers in this volume). Schwarzbach AE, Ricklefs RE. 2000. Systematic affinities of Rhizophoraceae and Anisophylleaceae, and intergeneric relationships within Rhizophoraceae, based on chloroplast DNA, nuclear ribosomal DNA and morphology. American Journal of Botany 87: 547–564. Setogushi H, Kosuge H, Tobe H. 1999. Molecular phylogeny of Rhizophoraceae based on rbcL gene sequences. Journal of Plant Research 112: 443–455. Tobe H, Raven PH. 1987. Systematic embryology of the Anisophylleaceae. Annals of the Missouri Botanical Garden 74: 1–26. Tobe H, Raven PH. 1988. Floral morphology and evolution in Anisophylleaceae. Botanical Journal of the Linnean Society 98: 1–25. Zhang LB, Simmons MP, Renner SS. 2007. A phylogeny of Anisophylleaceae based on six nuclear and plastid loci: ancient disjunctions and recent dispersal between South America, Africa and Asia. Molecular Phylogenetics and Evolution 44: 1057–1067. 197. CORYNOCARPACEAE CRIBWOOD FAMILY Carlquist S, Miller RB. 2001. Wood anatomy of Corynocarpus is consistent with cucurbitalean placement. Systematic Botany 26: 54–65. Garnock-Jones PJ, Brockie RE, FitzJohn RG. 2007. Gynodioecy, sexual dimorphism and erratic fruiting in Corynocarpus laevigatus (Corynocarpaceae). Australian Journal of Botany 55: 803–808. Novicke JW, Skvarla JJ. 1983. Pollen morphology and the relationships of the Corynocarpaceae. Taxon 32: 176–183. Patel RN. 1975. Wood anatomy of the dicotyledons indigenous to New Zealand. 9, Corynocarpus. New Zealand Journal of Botany 13: 19–29.

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Christenhusz, Fay & Chase

Philipson WR. 1987. Corynocarpus J.R. and G.Forst. — an isolated genus. Botanical Journal of the Linnean Society 95: 9–18. Stevenson G. 1978. Botanical evidence linking the New Zealand Maoris with New Caledonia and the New Hebrides. Nature 276: 704–705. Wagstaff SJ, Dawson MI. 2000. Classification, origin, and patterns of diversification of Corynocarpus (Corynocarpaceae) inferred from DNA sequences. Systematic Botany 25: 134–149. 198. CORIARIACEAE TANNER-BUSH FAMILY Carlquist S. 1985. Wood anatomy of Coriariaceae: phylogenetic and ecological implications. Systematic Botany 10: 174–183. Garg M. 1981. Pollen morphology and systematic position of Coriaria. Phytomorphology 30: 5–10. Gregor H-J. 1980. Seeds of the genus Coriaria Linné (Coriariaceae) in the European Neogene. Tertiary Research 3: 61–69. Good RDO. 1930. The geography of the genus Coriaria. New Phytologist 29: 170–198. Skog LE. 1972. The genus Coriaria in the Western Hemisphere. Rhodora 74: 242–253. Thomson PN, Gornall RJ. 1995. Breeding systems in Coriaria (Coriariaceae). Botanical Journal of the Linnean Society 117: 293–304. Yokoyama J, Suziki M, Iwatsuki K, Hasebe M. 2000. Molecular phylogeny of Coriaria, with special emphasis on the disjunct distribution. Molecular Phylogenetics and Evolution 14: 11–19. 199. CUCURBITACEAE CUCUMBER FAMILY Carlquist S. 1992. Wood anatomy of selected Cucurbitaceae and its relationship to habit and systematics. Nordic Journal of Botany 12: 347–355. Chandler MEJ. 1964. The Lower Tertiary floras of southern England IV: a summary and survey of findings in the light of recent botanical observations. British Museum (Natural History), London. Clarke AC, Burtenshaw MK, McLenachan PA, Erickson DL, Penny D. 2006. Reconstructing the origins and dispersal of the Polynesian bottle gourd (Lagenaria siceraria). Molecular Biology and Evolution 23: 893–900. Dillehay TD, Rossen J, Andres TC, Williams DE. 2007. Preceramic adoption of peanut, squash, and cotton in northern Peru. Science 316: 1890–1893. Duchen P, Renner SS. 2010. The evolution of Cayaponia (Cucurbitaceae): repeated shifts from bat to bee pollination and long-distance dispersal to Africa 2–5 million years ago. American Journal of Botany 97: 1129–1141. Erickson DL, Smith BD, Clarke AC, Sandweiss DH, Tuross N. 2005. Asian origin for a 10,000-year-old domesticated plant in the Americas. Proceedings of the National Academy of Sciences of the USA 102: 18315–18320. Gentry AH, Wettach RH. 1986. Fevillea — a new oil seed from Amazonian Peru. Economic Botany 40: 177–185. Goldman A. 2014. The complete squash: a passionate grower’s guide to pumpkins, squashes and gourds. Workman, New York. Heiser CB, Schilling EE. 1988. Phylogeny and distribution of Luffa (Cucurbitaceae). Biotropica 20: 185–191. Jeffrey C. 1980. A review of Cucurbitaceae. Botanical Journal of the Linnean Society 81: 233–247. Jeffrey C. 2005. A new system of Cucurbitaceae. Botanicheskii Zhurnal 90: 332–335. Kirkbride JH Jr. 1993. Biosystematic monograph of the

genus Cucumis. Parkway, Boone. Kocyan A, Zhang LB, Schaefer H, Renner SS. 2007. A multi-locus chloroplast phylogeny for the Cucurbitaceae and its implications for character evolution and classification. Molecular Phylogenetics and Evolution 44: 553–577. Monro AK, Stafford PJ. 1998. A synopsis of the genus Echinopepon (Cucurbitaceae: Sicyoeae), including three new taxa. Annals of the Missouri Botanical Garden 85: 257–272. Olson ME. 2003. Stem and leaf anatomy of the arborescent Cucurbitaceae Dendrosicyos socotrana with comments on the evolution of pachycauls from lianas. Plant Systematics and Evolution 239: 199–214. Renner SS, Schaefer H, Kocyan A. 2007. Phylogenetics of Cucumis (Cucurbitaceae): cucumber (C. sativus) belongs in an Australian/Asian clade far from African melon (C. melo). BMC Evolutionary Biology 7: 58. Sanjur OI, Piperno DR, Andres TC, Wessel-Beaver L. 2002. Phylogenetic relationships among domesticated and wild species of Cucurbita (Cucurbitaceae) inferred from a mitochondrial gene: implications for crop plant evolution and areas of origin. Proceedings of the National Academy of Sciences of the USA 99: 535–540. Schaefer H, Heibl C, Renner SS. 2009. Gourds afloat: a dated phylogeny reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous oversea dispersal events. Proceedings of the Royal Society B 276: 843–851. Schaefer H, Renner SS. 2011. Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60: 122–138. Sebastian PM, Schaefer H, Telford IRH, Renner SS. 2010. Cucumber and melon have their wild progenitors in India, and the sister species of Cucumis melo is from Australia. Proceedings of the National Academy of Sciences of the USA 107: 14269–14273. Smith BD. 1997. The initial domestication of Cucurbita pepo in the Americas 10,000 years ago. Science 276: 932–934. Van der Ham RWJM, Van Heuven BJ. 2003. A new type of Old World Cucurbitaceae pollen. Grana 45: 241–248. Whitaker TW, Davis GN. 1962. Cucurbits — botany, cultivation and utilization. Leonard Hill, London. Zhang LB, Simmons MP, Kocyan A, Renner SS. 2006. Phylogeny of the Cucurbitales based on DNA sequences of nine loci from three genomes: implications for morphological and sexual system evolution. Molecular Phylogenetics and Evolution 39: 305–322. 200. TETRAMELACEAE FALSE-HEMP-TREE FAMILY Lakhanpal RN, Verma JK. 1965. Fossil wood of Tetrameles from the Deccan Intertrappean beds of Mohgaonkalan, Madhya Pradesh. Paleobotanist 14: 209–213. Swensen SM, Mullin BC, Chase MW. 1994. Phylogenetic affinities of Datiscaceae based on an analysis of nucleotide sequences from the plastid rbcL gene. Systematic Botany 19: 157–168. 201. DATISCACEAE DURANGO-ROOT FAMILY Boesewinkel FD. 1984. Ovule and seed structure in Datiscaceae. Acta Botanica Neerlandica 32: 417–425.

FURTHER READING Davidson C. 1976. Anatomy of xylem and phloem of Datiscaceae. Natural History Museum of Los Angeles County, Contributions to Science 280: 1–28. Holmes WC, Blizzard HJ. 2010. The derivation of the name Datisca (Datiscaceae). Phytoneuron 2010-6: 1–2. Liston A, Rieseberg LH, Elias TS. 1989. Morphological stasis and molecular divergence in the intercontinental disjunct genus Datisca (Datiscaceae). Aliso 12: 525–542. Swensen SM, Luthi JN, Rieseberg LH. 1998. Datiscaceae revisited: monophyly and the sequence of breeding system evolution. Systematic Botany 23: 157–169. Swensen SM, Mullin BC, Chase MW. 1994. Phylogenetic affinities of Datiscaceae based on an analysis of nucleotide sequences from the plastid rbcL gene. Systematic Botany 19: 157–168. 202. BEGONIACEAE BEGONIA FAMILY Clement WL, Tebbitt MC, Forrest LL, Blair JE, Bouillet L, Eriksson T, Swensen SM. 2004. Phylogenetic position and biogeography of Hillebrandia sandwicensis (Begoniaceae): a rare Hawaiian relict. American Journal of Botany 91: 905–917. Cuerrier A, Brouillet L, Barabé D. 1990. Numerical taxonomic study of the Begoniaceae using the Mantel test on leaf microcharacters. Taxon 39: 549–560. Doorenbos J. 1985. Domestication of Begonia. Acta Botanica Neerlandica 34: 230–231. Forrest LL, Hollingsworth PM. 2003. A recircumscription of Begonia based on nuclear ribosomal sequences. Plant Systematics and Evolution 241: 193–211. Forrest LL, Hughes M, Hollingsworth PM. 2005. A phylogeny of Begonia using nuclear ribosomal sequence data and non-molecular characters. Systematic Botany 30: 671–682. Goodall-Copestake WP, Harris DJ, Hollingsworth PM. 2009. The origin of a mega-diverse genus: dating Begonia (Begoniaceae) using alternative datasets, calibrations, and relaxed clock methods. Botanical Journal of the Linnean Society 159: 363–380. Hughes M, Hollingsworth PM. 2008. Population genetic divergence corresponds with species-level biodiversity patterns in the large genus Begonia. Molecular Ecology 17: 2643–2651. Lee YS. 1974. A study of stem anatomy in Begonia L. Phytologia 27: 464–489. Panda S, De Wilde JJFE. 1995. Diversity and taxonomic value of stigmatic surfaces in Begoniaceae: SEM analysis. Acta Botanica Neerlandica 44: 139–150. Plana V, Gascoigne A, Forrest LL, Harris D, Pennington RT. 2004. Pleistocene and pre-Pleistocene Begonia speciation in Africa. Molecular Phylogenetics and Evolution 31: 449–461. Rajbhandary S, Hughes M, Phuttai T, Thomas DC, Shrestha KK. 2011. Asian Begonia: out of Africa via the Himalayas? Gardens’ Bulletin Singapore 63: 277–286. Rubite RR, Hughes M, Alejandro GJD, Peng C-I. 2013. Recircumscription of Begonia sect. Baryandra (Begoniaceae): evidence from molecular data. Botanical Studies 54: 38. Sosef MSM. 1994. Refuge begonias. Taxonomy, phylogeny and historical biogeography of Begonia sect. Loasibegonia and sect. Scutobegonia in relation to glacial rain forest refuges in Africa. Wageningen Agricultural University Papers 94-1: 1–306.

Tebbitt MC. 2005. Begonias, cultivation, identification, and natural history. Timber Press, Portland. Thomas DC, Hughes M, Phutthai T, Rajbhandary S, Rubite R, Ardi WH, Richardson JE. 2011. A noncoding plastid DNA phylogeny of Asian Begonia (Begoniaceae): evidence for morphological homoplasy and sectional polyphyly. Molecular Phylogenetics and Evolution 60: 428–444. Thompson ML, Thompson EJ. 1981. Begonias. The complete reference guide. Times Books, New York. 203. LEPIDOBOTRYACEAE SNAIL-CEDAR FAMILY Hammel BE, Zamora NA. 1993. Ruptiliocarpon (Lepidobotryaceae): a new arborescent genus and tropical American link to Africa, with a reconsideration of the family. Novon 3: 408–417. Léonard J. 1950. Lepidobotrys Engl., type d’une famille nouvelle des spermatophytes: les Lepidobotryaceae. Bulletin du Jardin Botanique National de la Belgique 20: 31–40. Mennega AMW. 1993. Comparative wood anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3: 418–422. Tobe H, Hammel B. 1993. Floral morphology, embryology, and seed anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3: 423–428. Van Welzen P, Baas P. 1984. A leaf anatomical contribution to the classification of the Linaceae complex. Blumea 29: 453–479. 204. CELASTRACEAE SPINDLE FAMILY Airy Shaw HK, Cutler DF, Nilsson S. 1973. Pottingeria, its taxonomic position, anatomy and palynology. Kew Bulletin 28: 97–104. Alvarez M, Martín JS, Deil U. 2012. Nanism and ephemerism as reasons for hidden abundance in vernal pool plants: the example of Lepuropetalon spathulatum in Chile. Feddes Repertorium 123: 55–66. Baas P, Geesink R, Van Heel W, Muller J. 1979. The affinities of Plagiopteron suaveolens Griff. (Plagiopteraceae). Grana 18: 69–89. Blakelock RA. 1951. A synopsis of the genus Euonymus L. Kew Bulletin 2: 210–290. Carlquist S. 1987. Wood anatomy and relationships of Stackhousiaceae. Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 108: 473–480. Coughenour JM, Simmons MP, Lombardi JA, Cappa JJ. 2010. Phylogeny of Celastraceae subfamily Salacioideae and tribe Lophopetaleae inferred from morphological characters and nulear and plastid genes. Systematic Botany 35: 358–367. Coughenour JM, Simmons MP, Lombardi JA, Yakobson K, Archer RH. 2011. Phylogeny of Celastraceae subfamily Hippocrateoideae inferred from morphological characters and nuclear and plastid loci. Molecular Phylogenetics and Evolution 59: 320–330. Getahun A, Krikorian AD. 1973. Chat: coffee’s rival from Harar, Ethiopia. I. Botany, cultivation and use. Economic Botany 27: 353–377. Goldblatt P, Tobe H, Carlquist S, Patel VC. 1985. Familial position of the Cape genus Empleuridium. Annals of the Missouri Botanical Garden 72: 167–183. Islam MB, Simmonds MP, Archer RH. 2006. Phylogeny of the Elaeodendron group (Celastraceae) inferred from morphological characters and nuclear and plastid genes. Systematic Botany 31: 512–524.

Martens P. 1936. Pollination et biologie florale chez Parnassa palustris. Bulletin du Société Royale Botanique de la Belgique 68: 183–221. Matthews ML, Endress PK. 2005. Comparative floral structure and systematics in Celastrales (Celastraceae, Parnassiaceae, Lepidobotryaceae). Botanical Journal of the Linnean Society 149: 129–194. McKenna MJ, Simmons MP, Bacon CD, Lombardi JA. 2011. Delimitation of segregate genera of Maytenus s.l. (Celastraceae) based on molecular and morphological characters. Systematic Botany 36: 922–932. Mennega AMW. 1997. Wood anatomy of the Hippocrateoideae (Celastraceae). IAWA Journal 18: 331–368. Perrier de la Bâthie H. 1942. Au sujet des affinités des Brexia et des Célastreacées et de deux Brexia nouveaux de Madagascar. Bulletin de la Société Botanique de France 89: 219–221. Savolainen C, Spichiger R, Manen JF. 1997. Polyphyletism of Celastrales deduced from a chloroplast noncoding DNA region. Molecular Phylogenetics and Evolution 7: 145–157. Simmons MP, Savolainen V, Clevinger CC, Archer RH, Davis JI. 2001. Phylogeny of the Celastraceae inferred from 26S nuclear DNA, phytochrome B, rbcL, atpB, and morphology. Molecular Phylogenetics and Evolution 19: 353–366. Simmons MP, Cappa JJ, Archer RH, Ford AJ, Eichstedt D, Clevinger CC. 2008. Phylogeny of the Celastreae (Celastraceae) and the relationships of Catha edulis (qat) inferred from morphological characters and nuclear and plastid genes. Molecular Phylogenetics and Evolution 48: 745–757. Smith AC. 1941. The American species of Hippocrateaceae. Brittonia 3: 341–555. Vasudeva Rao MK, Chakrabarty T. 1985. Nicobariodendron Vasud., T. Chakrab. (Celastraceae): a new genus from the Nicobar Islands, India. Journal of Economic and Taxonomic Botany 7: 513–516. Zhang LB, Simmonds MP. 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes. Systematic Botany 31: 122–137. 205. HUACEAE CAMEROON-GARLIC FAMILY Baas P. 1972. Anatomical contributions to plant taxonomy, II. The affinities of Hua Pierre and Afrostyrax Perkins et Gilg. Blumea 20: 161–192. Mildbraed J. 1913. Über die Gattungen Afrostyrax Perk et Gilg und Hua Pierre und die “KnoblauchRinden” Westafrikas. Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 49: 552–559. 206. CONNARACEAE ZEBRAWOOD FAMILY Jongkind CCH, Lemmens RHMJ. 1989. The Connaraceae: a taxonomic study with special emphasis on Africa. Landbouwuniversiteit, Wageningen. Lemmens RHMJ. 1992. A reconsideration of Ellipanthus (Connaraceae) in Madagascar and continental Africa and a comparison with the species in Asia. Adansonia 14: 99–108. 207. OXALIDACEAE WOOD-SORREL FAMILY Emshwiller E, Doyle JJ. 2002. Origin of domestication and polyploidy in oca (Oxalis tuberosa, Oxalidaceae). American Journal of Botany 89: 1042–1056. Heibl C, Renner SS. 2012. Distribution models and a dated phylogeny for Chilean Oxalis species reveal occupation of new habitats by different lineages,

Plants of the World

709

FURTHER READING not rapid adaptive radiation. Systematic Biology 61: 823–834. King SR, Gershoff SN. 1987. Nutritional evaluation of three underexploited Andean tubers: Oxalis tuberosa (Oxalidaceae), Ullucus tuberosus (Basellaceae), and Tropaeolum tuberosum (Tropaeolaceae). Economic Botany 41: 503–511. Lack AJ, Kevan PG. 1987. The reproductive biology of a distylous tree, Sarcotheca celebica (Oxalidaceae) in Sulawesi, Indonesia. Botanical Journal of the Linnean Society 95: 1–8. Lourteig A. 2000. Oxalis L. subgéneros Monotaxis (Small) Lourt., Oxalis y Trifidus Lourt. Bradea 7: 201–629. Oberlander KC, Dreyer LL, Bellstedt DU. 2011. Molecular phylogenetics and origins of southern African Oxalis. Taxon 60: 1667–1677. Veldkamp JF. 1967. A revision of Sarcotheca Bl. and Dapania Korth. (Oxalidaceae). Blumea 15: 519–543. 208. CUNONIACEAE BUTTERSPOON-TREE FAMILY Barnes RW, Jordan GJ. 2000. Eucryphia (Cunoniaceae) reproductive and leaf macrofossils from Australian Cainozoic sediments. Australian Systematic Botany 13: 373–394. Barnes RW, Hill RS, Bradford JC. 2001. The history of Cunoniaceae in Australia from macrofossil evidence. Australian Journal of Botany 49: 301–320. Bradford JC. 1998. A cladistic analysis of speciesgroups in Weinmannia (Cunoniaceae) based on morphology and inflorescence architecture. Annals of the Missouri Botanical Garden 85: 565–593. Bradford JC. 2002. Molecular phylogenetics and morphological evolution in Cunonieae (Cunoniaceae). Annals of the Missouri Botanical Garden 89: 491–503. Bradford JC, Barnes RW. 2001. Phylogenetics and classification of Cunoniaceae (Oxalidales) using chloroplast DNA sequences and morphology. Systematic Botany 26: 356–385. Chambers KL, Poinar G Jr, Buckley R. 2010. Tropidogyne, a new genus of early Cretaceous eudicots (Angiospermae) from Burmese amber. Novon 20: 23–29. Dickison WC. 1980. Comparative wood anatomy and evolution of Cunoniaceae. Allertonia 2: 281–321. Doweld AB. 1998. The carpology and taxonomic relationships of Davidsonia (Davidsoniaceae). Edinburgh Journal of Botany 55: 13–25. Hopkins HCF, Rozefelds AC, Pillon Y. 2013. Karrabina gen. nov. (Cunoniaceae), for the Australian species previously placed in Geissois, and a synopsis of genera in the tribe Geissoieae. Australian Systematic Botany 26: 167–185. Hufford L, Dickison WC. 1992. A phylogenetic analysis of Cunoniaceae. Systematic Botany 17: 181–200. Matthews ML, Endress PK, Schönenberger J, Friis EM. 2001. A comparison of f loral structures of Anisophylleaceae and Cunoniaceae and the problem of their systematic position. Annals of Botany 88: 439–455. Moody ML, Hufford L. 2000. Floral development and structure of Davidsonia (Cunoniaceae). Canadian Journal of Botany 78: 1034–1043. Sweeney PW, Bradford JC, Lowry II PP. 2004. Phylogenetic position of the New Caledonian endemic genus Hooglandia as determined by maximum parsimony analysis of chloroplast

710

Christenhusz, Fay & Chase

DNA. Annals of the Missouri Botanical Garden 91: 266–274. 209. ELAEOCARPACEAE FAIRY-PETTICOATS FAMILY Bricker JS. 1992. Pollination biology of the genus Crinodendron (Elaeocarpaceae). Journal of the Arizona-Nevada Academy of Sciences 24–25: 51–54. Carlquist S. 1977. Wood anatomy of Tremandraceae: phylogenetic and ecological implications. American Journal of Botany 64: 704–713. Coode MJE. 1983. A conspectus of Sloanea (Elaeocarpaceae) in the Old World. Kew Bulletin 38: 347–427. Crayn DM, Rossetto M, Maynard DJ. 2006. Molecular phylogeny and dating reveals and OligoMiocene radiation of dry-adapted shrubs (former Tremandraceae) from rainforest tree progenitors (Elaeocarpaceae) in Australia. American Journal of Botany 93: 1328–1342. Earle Smith C. 1954. The New World species of Sloanea (Elaeocarpaceae). Contributions from the Gray Herbarium 175: 1–114. Gasson P. 1996. Wood anatomy of the Elaeocarpaceae. Pp. 47–71. In: Donaldson LA, Singh AP, Butteheld BG, Whitehouse LJ (eds) Recent advances in wood anatomy. New Zealand Forest Research Institute, Rotorua. Manchester SR, Kvacek Z. 2009. Fruits of Sloanea (Elaeocarpaceae) in the Paleogene of North America and Greenland. International Journal of Plant Sciences 170: 941–950. Matthews ML, Endress PK. 2002. Comparative floral morphology and systematics in Oxalidales (Oxalidaceae, Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae, Elaeocarpaceae, Tremandraceae). Botanical Journal of the Linnean Society 140: 321–381. Thompson J. 1976. A revision of the genus Tetratheca (Tremandraceae). Telopia 1: 139–215. Van Balgooy MMJ. 1982. A revision of Sericolea Schlechter (Elaeocarpaceae). Blumea 28: 103–141. 210. CEPHALOTACEAE ALBANYPITCHERPLANT FAMILY Adlassnig W. Peroutka M, Lendl T. 2011. Traps of carnivorous pitcher plants as a habitat: composition of the fluid, biodiversity and mutualistic activities. Annals of Botany 107: 181–194. Albert VA, Williams SE, Chase MW. 1992. Carnivorous plants: phylogeny and structural evolution. Science 257: 1491–1495. Conran JG, Denton MD. 1996. Germination in the Western Australian pitcher plant Cephalotus follicularis and its unusual early seedly development. Western Australian Naturalist 21: 37–42. Froebe HA, Bauer N. 1988. Die Morphogenese der Kannenblätter von Cephalotus follicularis Labill. Akademie der Wissenschaften und der Literatur, Mainz. Abhandlungen der mathematisch-naturwissenschaftlichen Klasse 3: 1–19. McPherson S. 2009. Pitcher plants of the Old World (2 vols). Redfern Natural History Publications, Poole. Luffitz F. 1966. The West Australian pitcher plant (Cephalotus follicularis Labill.) Australian Plants 12: 34–35. 211. BRUNELLIACEAE PALO-BOBO FAMILY Cuatrecasas, J. 1970; 1985. Brunelliaceae. Flora Neotropica Monographs 2: 1–189; Flora Neotropica Monographs 2 (supplement): 29–103.

Orozco CI. 1997. Sobre la posición sistematica de Brunellia Ruiz, Pavón. Caldasia 19: 145–164. Orozco CI. 2001. Pollen morphology of Brunellia (Br unelliaceae) and related taxa in the Cunoniaceae. Grana 40: 245–255. Orozco CI, Weberling F. 1999. A comparative study of inflorescences in Brunellia Ruiz, Pav. (Brunelliaceae) and related taxa. Beiträge zur Biologie der Pflanzen 71: 261–279. 212. PANDACEAE KANA-NUT FAMILY Forman LL. 1966. The reinstatement of Galearia Zoll., Mor. and Microdesmis Hook.f. in the Pandaceae, with appendices by C. R. Metcalfe and N. Parameswaran. Kew Bulletin 20: 309–321. Hayden WC, Hayden SM. 2000. Wood anatomy of Acalyphoideae (Euphorbiaceae). IAWA Journal 21: 213–235. Nowicke JW. 1984. A palynological study of the Pandaceae. Pollen and Spores 26: 31–42. Xi Z, Ruhfel BR, Schaefer H, Amorim AM, Sugumaran M, Wurdack KJ, Endress PK, Matthews M, Stevens PF, Mathews S, Davis III CC. 2012. Phylogenomics and a posteriori data partitioning resolve the Cretaceous angiosperm radiation in Malpighiales. Proceedings of the National Academy of Sciences of the USA 109: 17519–17524. 213. IRVINGIACEAE OGBONO-NUT FAMILY Byng JW, Bernardini B, Christenhusz MJM, Chase MW. 2016. Systematics of Irvingiaceae and Ixonanthaceae (Malpighiales): phylogenetic analysis based on three plastid DNA loci. Phytotaxa 260: 157–166. Den Hartog RM, Baas P. 1978. Epidermal characters of the Celastraceae sensu lato. Acta Botanica Neerlandica 27: 355–388. Furness CA. 2011. Comparative structure and development of pollen and tapetum in Malpighiales, with a focus on the parietal clade. International Journal of Plant Sciences 172: 980–1011. Nooteboom HP. 1967. The taxonomic position of Irvingioideae, Allantospermum Forman, and Cyrillopsis Kuhlm. Adansonia II, 7: 161–168. Weberling F, Lörcher H, Böhnke F. 1980. Die Stipeln der Irvingioideae und Recchioideae und ihre systematische Wertung nebst Bemerkungen zur Holzanatomie und Palynologie. Plant Systematics and Evolution 133: 261–283. 214. CTENOLOPHONACEAE LITOH FAMILY Krutsch W. 1989. Paleogeography and historical phytogeography (paleochorology) in the Neophyticum. Plant Systematics and Evolution 162: 5–61. Link DA. 1992. Floral nectaries of the Geraniales and their systematic implication. IV. Ctenolophonaceae Badre. Flora 187: 103–107. Saad SI. 1962. Pollen morphology of Ctenolophon. Botaniska Notiser 115: 49–57. Van der Ham RWJM. 1989. New observations on the pollen of Ctenolophon Oliver (Ctenolophonaceae), with remarks in the evolutionary history of the genus. Review of Palaeobotany and Palynology 59: 153–160. 215. RHIZOPHORACEAE MANGROVE FAMILY Alston AHG. 1925. Revision of Cassipourea. Kew Bulletin 1925: 241–276. Duke NC. 2010. Overlap of eastern and western mangroves in the southwestern Pacific: hybridisation of all three Rhizophora (Rhizophoraceae) combinations in New Caledonia. Blumea 55: 171–188.

FURTHER READING Graham A. 2006. Paleobotanical evidence and molecular data in reconstructing the historical phytogeography of Rhizophoraceae. Annals of the Missouri Botanical Garden 93: 327–334. Juncosa AM, Tomlinson PB. 1987. Floral development in mangrove Rhizophoraceae. American Journal of Botany 74: 1263–1279. Levin GA. 1992. Systematics of Paradrypetes (Euphorbiaceae). Systematic Botany 17: 74–83. Raven PH, Tomlinson PB. 1988. RhizophoraceaeAnisophylleaceae: a symposium. Annals of the Missouri Botanical Garden 75: 1258. [See also other papers in this volume.] Schwarzbach AE, Ricklefs RE. 2000. Systematic affinities of Rhizophoraceae and Anisophylleaceae, and intergeneric relationships within Rhizophoraceae, based on chloroplast DNA, nuclear ribosomal DNA, and morphology. American Journal of Botany 87: 547–564. Tomlinson PB. 1986. The botany of mangroves. Cambridge University Press, Cambridge. Tomlinson PB, Cox PA. 2000. Systematic and functional anatomy of seedlings in mangrove Rhizophoraceae: vivipary explained? Botanical Journal of the Linnean Society 134: 215–231. Wurdack KJ, Davies CC. 2008. Malpighiales phylogenetics: gaining ground on one of the most recalcitrant clades in the angiosperm tree of life. American Journal of Botany 96: 1551–1570. 216. ERYTHROXYLACEAE COCA FAMILY Barros MAG. 1998. Sistemas reproductivos e polinização en espécies simpátricas de Erythroxylum P. Br. (Erythroxylaceae) do Brasil. Revista Brasileira de Botânica 21: 159–166. Bohm BA, Ganders FR, Plowman T. 1982. Biosystematics and evolution of cultivated coca (Erythroxylaceae). Systematic Botany 7: 121–133. Davis W. 1997. One river — science, adventure and hallucinogenics in the Amazon Basin. Simon, Schuster, London. Ganders FR. 1979. Heterostyly in Erythroxylum coca (Erythroxylaceae). Botanical Journal of the Linnean Society 78: 11–20. Johnson EL, Saunders JA, Mischke S, Helling CS, Emche SD. 2003. Identification of Erythroxylum taxa by AFLP DNA analysis. Phytochemistry 64: 187–197. Plowman T. 1979. Botanical perspectives on coca. Journal of Psychedelic Drugs 11: 103–117. Rury PM. Systematic anatomy of Erythroxylum P. Browne: practical and evolutionary implications for the cultivated cocas. Journal of Ethnopharmacology 3: 229–263. Rury PM. 1985. Systematic and ecological wood anatomy of the Erythroxylaceae. IAWA Bulletin new series 6: 365–397. Rury PM, Plowman T. 1983. Morphological studies of archeological and recent coca leaves (Erythroxylum spp.). Botanical Museum Leaflets of Harvard University 29: 297–341. United Natuons Drug Control, Transnational Institute. Coca leaf: myth and reality. Website: ht t p://w w w.u nd r u gc ont r ol.i n fo/e n /i s s ue s / unscheduling-the-coca-leaf/item/262-coca-leafmyths-and-reality (accessed May 2014). 217. OCHNACEAE MICKEY-MOUSE-PLANT FAMILY Amaral MdCE. 1991. Phylogenetische Systematik der Ochnaceae. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 113: 105–196.

Decker JM. 1966. Wood anatomy and phylogeny of Luxemburgieae (Ochnaceae). Phytomorphology 16: 39–55. Dickison WC. 1990. The morphology and relationships of Medusagyne (Medusagynaceae). Plant Systematics and Evolution 171: 27–55. Doweld B. 1998. On the phylogenetic relationships of Medusagyne (Medusagynaceae) as evidenced by the structure of its fruits and seeds. Botanicheskii Zhurnal 83: 54–68. Fay MF, Swensen SM, Chase MW. 1997. Taxonomic affinities of Medusagyne oppositifolia (Medusagynaceae). Kew Bulletin 52: 111–120. Foster AS. 1951. Heterophylly and foliar venation in Lacunaria. Bulletin of the Torrey Botanical Club 78: 382–400. Guédès M, Sastre C. 1981. Morphology of the gynoecium and systematic position of the Ochnaceae. Botanical Journal of the Linnean Society 82: 121–138. Kubitzki K, Amaral MdCE. 1991. Transference of function in the pollination system of the Ochnaceae. Plant Systematics and Evolution 177: 77–80. Roberson A, Wise R, White F. 1989. Medusagyne oppositifolia. Kew Magazine 6: 166–171. Schneider JV, Swenson U, Samuel R, Stuessy T, Zizka G. 2006. Phylogenetics of Quiinaceae (Malpighiales): evidence from trnL-trnF sequence data and morphology. Plant Systematics and Evolution 257: 189–203. Schneider JV, Bissiengou P, Amaral MdCE, Tahir A, Fay MF, Thines M, Sosef MSM, Zizka G, Chatrou LW. (2014). Phylogenetics, ancestral state reconstruction and a new infrafamilial classification of the pantropical Ochnaceae (Medusagynaceae, Ochnaceae s.str., Quiinaceae) based on five DNA regions. Molecular Phylogenetics and Evolution 78: 199–214. 218. BONNETIACEAE CASCARILLA FAMILY Carvalho MP, Lima MMC, Santos MG, Rocha LM, Kuster RM. 2013. Anthraquinones and xanthone from Bonnetia stricta and their chemosystematic significance. Biochemical Systematics and Ecology 48: 73–75. Dickison WC, Weitzman AL. 1996. Comparative anatomy of the young stem, node and leaf of Bonnetiaceae, including observations on a foliar endodermis. American Journal of Botany 83: 405–418. Dickison WC, Weitzman AL. 1998. Floral morphology and anatomy of Bonnetiaceae. Journal of the Torrey Botanical Society 125: 258–286. Weitzman AL, Stevens PF. 1997. Notes on the circumscription of Bonnetiaceae and Clusiaceae, with taxa and new combinations. BioLlania, Edición Especial 6: 551–564. 219. CLUSIACEAE MANGOSTEEN FAMILY Bennett GJ, Lee H-H. 1989. Xanthones from Guttiferae. Phytochemistry 28: 967–998. Crepet WL, Nixon KC. 1998. Fossil Clusiaceae from the Late Cretaceous (Turonian) of New Jersey and implications regarding the history of bee pollination. American Journal of Botany 85: 1122–1133. Dick CW, Abdul-Salim K, Bermingham E. 2003. Molecular systematic analysis reveals cryptic Tertiary diversification of a widespread tropical rainforest tree. American Naturalist 162: 691–703. Gustafsson MHG. 2000. Floral morphology and relationships of Clusia gundlachii with a discussion

of floral organ identity and diversity of the genus Clusia. International Journal of Plant Sciences 161: 43–53. Gustafsson MHG, Bittrich V, Stevens PF. 2002. Phylogeny of Clusiaceae based on rbcL sequences. International Journal of Plant Sciences 163: 1045–1054. Lüttge U. 2002. The genus Clusia L.: molecular evidence for independent evolution of photosynthetic flexibility. Plant Biology 4: 86–93. Mourão KSM, Beltrati CM. 1995. Morphologia dos frutos, sementes e plântulas de Platonia insignis Mart. (Clusiaceae). I. Aspectos anatômicos dos frutos e sementes em desenvolvimento. Acta Amazônica 25: 11–32. Mourão KSM, Marzinek J. 2009. Fruit ontogenesis in Clusia parviflora Humb., Bonpl. ex Willd. (Clusiaceae). Acta Botanica Brasilica 23: 797–804. Planchon JE, Triana J. 1860–1862. Mémoire sur la famille des Guttifères. Annales des Sciences Naturelles, IV, Botanique 13: 306–376; 16: 263–308. Richards AJ. 1990. Studies in Garcinia, dioecious tropical forest trees: the origin of the mangosteen (G. mangostana L.). Botanical Journal of the Linnean Society 103: 301–308. Ruhfel BR, Bittrich V, Bove CP, Gustafsson MHG, Philbrick CT, Rutishauser R, Xi Z, Davis CC. 2011. Phylogeny of the clusioid clade (Malpighiales): evidence from plastid and mitochondrial genomes. American Journal of Botany 98: 306–325. Ruhfel BR, Stevens PF, Davis CC. 2013. Combined morphological and molecular phylogeny of the clusioid clade (Malpighiales) and the placement of the ancient rosid macrofossil Paleoclusia. International Journal of Plant Sciences 174: 910–936. Sweeney PW. 2008. Phylogeny and floral diversification in the genus Garcinia (Clusiaceae) and relatives. International Journal of Plant Sciences 169: 1288–1303. 220. CALOPHYLLACEAE TAKAMAKA FAMILY De Oliveira PEAM, Sazima K. 1990. Pollination biology of two species of Kielmeyera (Guttiferae) from Brazilian cerrado vegetation. Plant Systematics and Evolution 172: 35–49. Dunthorn MS. 2004. Cryptic dioecy in Mammea (Clusiaceae). Plant Systematics and Evolution 249: 191–196. Dunthorn MS. 2009. Foliar anatomy and fiber motifs in Mammea (Clusiaceae, Kielmeyeroideae). Plant Systematics and Evolution 280: 153–166. Stevens PF. 1980. A revision of the Old World species of Calophyllum (Guttiferae). Journal of the Arnold Arboretum 61: 117–699. 221. PODOSTEMACEAE RIVERWEED FAMILY Burkhardt G, Becker H, Grubert M, Thomas J, Eicher T. 1994. Bioactive chromenes from Rhyncholacis penicillata. Phytochemistry 37: 1593–1597. Jäger-Zürn I, Grubert M. 2000. Podostemaceae depend on sticky biofilms with respect to attachment to rocks in waterfalls. International Journal of Plant Sciences 161: 599–607. Kita Y, Kato M. 2001. Infrafamilial phylogeny of the aquatic angiosperm Podostemaceae inferred from the nucleotide sequence of the matK gene. Plant Biology 3: 156–163. Kita Y. 2002. Molecular phylogeny, morphological evolution and biogeography of Podostemaceae. Bunrui 2: 19–26.

Plants of the World

711

FURTHER READING Koi S, Kita Y, Hirayama Y, Rutishauser R, Huber KA, Kato M. 2012. Molecular phylogenetic analysis of Podostemaceae: implications for taxonomy of major groups. Botanical Journal of the Linnean Society 169: 461–492. Les DH, Philbrick CT, Novelo RA. 1997. The phylogenetic position of river-weeds (Podostemaceae): insights from rbcL sequence data. Aquatic Botany 57: 5–27. Philbrick CT, Novelo RA. 2004. Monograph of Podostemum (Podostemaceae). Systematic Botany Monographs 70: 1–106. 222. HYPERICACEAE ST. JOHN’S-WORT FAMILY Baas P. 1970. Floral and vegetative anatomy of Eliaea from Madagascar and Cratoxylum from Indomalesia (Guttiferae). Blumea 18: 369–391. Crockett SL, Robson NKB. 2011. Taxonomy and chemotaxonomy of the genus Hypericum. Medicinal and Aromatic Plant Science and Biotechnology 5: 1–13. Ernst E (ed). 2003. Hypericum: the genus Hypericum. Taylor, Francis, New York. Gogelein AJF. 1967, A revision of the genus Cratoxylum Bl. (Guttiferae). Blumea 15: 453–475. Meseguer AS, Sanmartin I. 2012. Paleobiology of the genus Hypericum (Hypericaceae): a survey of the fossil record and its paleogeographic implications. Anales del Jardín Botánico de Madrid 69: 97–106. Meseguer AS, Aldadoro JJ, Sanmartin I. 2013. Bayesian inference of phylogeny, morphology and range evolution reveals a complex evolutionary history in St. John’s wort (Hypericum). Molecular Phylogenetics and Evolution 67: 379–403. Nürk NM, Blattner FR. 2010. Cladistic analysis of morphological characters in Hypericum (Hypericaceae). Taxon 59: 1495–1507. Nürk, NM, Madriñán S, Carine MA, Chase MW, Blattner FR. 2013. Molecular phylogenetics and morphological evolution of St. John’s wort (Hypericum; Hypericaceae). Molecular Phylogenetics and Evolution 66: 1–16. Robson NKB. 1981. Studies in the genus Hypericum L. (Guttiferae), 2. Characters of the genus. Bulletin of the British Museum of Natural History (Botany) 8: 55–226. Robson NKB. 2012. Studies in the genus Hypericum L. (Hypericaceae) 9. Addenda, corrigenda, keys, lists and general discussion. Phytotaxa 72: 1–111. [and other papers by Robson cited here] Sirvent TM, Krasnoff CB, Gibson DM. 2003. Induction of hypericins and hyperforins in Hypericum perforatum in response to damage by herbivores. Journal of Chemical Ecology 29: 2667–2681. 223. CARYOCARACEAE SOUARI-TREE FAMILY Araújo PA de M, De Mattos Filho A. 1973. Estrutura das madeiras de Caryocaraceae, Archivos do Jardim Botânico do Rio de Janeiro 19: 5–47. Dickison WC. 1990. A study of the floral morphology and anatomy of the Caryocaraceae. Bulletin of the Torrey Botanical Club 117: 123–137. Prance GT. 1990. The genus Caryocar L. (Caryocaraceae): an underexploited tropical resource. Advances in Economic Botany 8: 177–188. Prance GT, Da Silva MF. 1973. Monograph of Caryocaraceae. Flora Neotropica 12: 1–75. 224. LOPHOPYXIDACEAE KOTEB FAMILY Pfeiffer H. 1951. Lophopyxis als Typus einer eigenen Familie. Revista Sudamericana de Botánica 10: 3–6.

712

Christenhusz, Fay & Chase

Sleumer H. 1968. The genus Lophopyxis Hook.f. (Lophopyxidaceae). Blumea 16: 321–323. Sleumer H. 1971. Lophopyxidaceae. Flora Malesiana I, 7: 89–92. 225. PUTRANJIVACEAE CHILDLIFE-TREE FAMILY Forster PI. 1997. A taxonomic revision of Drypetes Vahl (Euphorbiaceae) in Australia. Austrobaileya 4: 477–494. Tokuoka T, Tobe H. 1999. Embryology of tribe Drypeteae, an enigmatic taxon of Euphorbiaceae. Plant Systematics and Evolution 215: 189–208. Wurdack KJ, Hoffmann P, Samuel R, De Bruijn A, Van der Bank M, Chase MW. 2004. Molecular phylogenetic analysis of Phyllanthaceae (Phyllanthoideae pro parte, Euphorbiaceae sensu lato) using plastid rbcL DNA sequences. American Journal of Botany 91: 1882–1900. 226. CENTROPLACACEAE BIKU-BIKU FAMILY Hou D. 1958. A conspectus of the genus Bhesa (Celastraceae). Blumea 4: 149–153. Zhang LB, Simmonds MP. 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes. Systematic Botany 31: 122–137. Zhang XC, Baas P, Mennega AMW. 1990. Wood anatomy of Bhesa sinica (Celastraceae). IAWA Bulletin 11: 57–60. 227. ELATINACEAE WATERWORT FAMILY Carlquist S. 1984. Wood and stem anatomy of Bergia suffruticosa: relationships of Elatinaceae and broader significance of vascular tracheids and fibriform vessel elements. Annals of the Missouri Botanical Garden 71: 232–242. Davis CC, Chase MW. 2004. Elatinaceae are sister to Malpighiaceae; Peridiscaceae belong to Saxifragales. American Journal of Botany 91: 262–273. Frisedahl A. 1927. Über die Entwicklung chasmo- und kleistogamer Blüten bei der Gattung Elatine. Acta Horti Gothoburgensis 3: 99–142. Salisbury EJ. 1967. On the reproduction and biology of Elatine hexandra (Lapierre) DC. (Elatinaceae); a typical species of exposed mud. Kew Bulletin 21: 139–149. 228. MALPIGHIACEAE ACEROLA FAMILY Anderson C. 2011. Revision of Ryssopterys and transfer to Stigmatophyllon (Malpighiaceae). Blumea 56: 73–104. Anderson C, Anderson WR, Davies CC. 2006 (onwards). Malpighiaceae website. http://herbarium.lsa.umich.edu/malpigh/ Anderson WR. 1979. Floral conservatism in Neotropical Malpighiaceae. Biotropica 11: 219–223. Anderson WR. 1980. Cryptic self-fertilisation in the Malpighiaceae. Science 207: 892–893. Cameron KM, Chase MW, Anderson WR, Hills HG. 2001. Molecular systematics of Malpighiaceae: evidence from plastid rbcL and matK sequences. American Journal of Botany 88: 1847–1862. Cardinal S, Danforth BN. 2013. Bees diversified in the age of eudicots. Proceedings of the Royal Society B, 280: 20122686. Davis CC, Anderson WR. 2010. A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology. American Journal of Botany 97: 2031–2046.

Davis CC, Chase MW. 2004. Elatinaceae are sister to Malpighiaceae; Peridiscaceae belong to Saxifragales. American Journal of Botany 91: 262–273. Davis CC, Bell CD, Mathews S, Donoghue MJ. 2002. Laurasian migration explains Gondwana disjunctions: evidence from Malpighiaceae. Proceedings of the National Academy of Sciences of the USA 99: 6833–6837. Davis CC, Webb CO, Wurdack KJ, Jaramillo CA, Donoghue MJ. 2005. Explosive radiation of Malpighiales supports a Mid-Cretaceous origin of modern tropical rain forests. American Naturalist 165: E36–E65. Davis CC, Schaefer H, Xi Z, Baum DA, Donoghue MJ, Harmon LJ. 2014. Long-term morphological stasis maintained by a plant-pollinator mutualism. Proceedings of the National Academy of Sciences of the USA 111: 5914–5919. Meyer FK. 2000. Revision der Gattung Malpighia L. (Malpighiaceae). Phanerogamarum Monographiae 23: 1–630. Papadopulos AS, Powell MP, Pupulin F, Warner J, Hawkins JA, Salamin N, Chittka L, Williams NH, Whitten WM, Loader D, Valente LM, Chase MW, Savolainen V. 2013. Convergent evolution of floral signals underlies the success of Neotropical orchids. Proceedings of the Royal Society B, 280: 20130960 Taylor DW, Crepet WL. 1987. Fossil floral evidence of Malpighiaceae and an early plant-pollinator relationship. American Journal of Botany 74: 274–286. 229. BALANOPACEAE PIMPLEBARK FAMILY Carlquist S. 1980. Anatomy and systematics of Balanopaceae. Allertonia 2: 191–246. Litt A, Chase MW. 1999. The systematic position of Euphronia, with comments on the position of Balanops: an analysis based on rbcL sequence data. Systematic Botany 23: 401–409. Sutter DM, Endress PK. 2003. Female flower and cupule structure in Balanopaceae, an enigmatic rosid family. Annals of Botany 92: 459–469. 230. TRIGONIACEAE TRIANGLE-VINE FAMILY Boesewinkel FD. 1987. Ovules and seeds of Trigoniaceae. Acta Botanica Neerlandica 36: 81–91. Lleras E. 1978. Trigoniaceae. Flora Neotropica Monograph 19: 1–73. Litt A, Chase MW. 1998. The systematic position of Euphronia, with comments on the position of Balanops: an analysis based on rbcL sequence data. Systematic Botany 23: 401–409. Matthews ML, Endress PK. 2008. Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chr ysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales). Botanical Journal of the Linnean Society 157: 249–309. 231. DICHAPETALACEAE RATBANE FAMILY Litt A, Chase MW. 1998. The systematic position of Euphronia, with comments on the position of Balanops: an analysis based on rbcL sequence data. Systematic Botany 23: 401–409. Matthews ML, Endress PK. 2008. Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chr ysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales). Botanical Journal of the Linnean Society 157: 249–309.

FURTHER READING Nandi OI, Chase MW, Endress PK. 1998. A combined cladistic analysis of angiosperms using rbcL and non-molecular data sets. Annals of the Missouri Botanical Garden 85: 137–212. Prance GT. 1972. Dichapetalaceae. Flora Neotropica Monograph 10: 1–84. Prance GT. 1995. A synopsis of the genus Staphanopodium (Dichapetalaceae). Kew Bulletin 50: 295–305. 232. EUPHRONIACEAE EUPHRONIA FAMILY Litt A, Chase MW. 1998. The systematic position of Euphronia, with comments on the position of Balanops: an analysis based on rbcL sequence data. Systematic Botany 23: 401–409. Marcano-Berti L. 1989. Euphroniaceae: una nueva familia. Pittieria 18: 15–19. Matthews ML, Endress PK. 2008. Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales). Botanical Journal of the Linnean Society 157: 249–309. 233. CHRYSOBALANACEAE COCOPLUM FAMILY Bardon L, Champagne J, Dexter K, Sothers C, Prance G, Chave J. 2012. Origin and evolution of the Chrysobalanaceae family: insights into the evolution of plants in the Neotropics. Botanical Journal of the Linnean Society 171: 19–37. Bardon L, Sothers C, Prance GT, Male PJG, Xi Z, Davis CC, Murienne J, Garcia-Villacorta R, Coissac E, Lavergne S, Chave J. 2016. Unraveling the biogeographical history of Chrysobalanaceae from plastid genomes. American Journal of Botany 103: 1089–1102. Matthews ML, Endress PK. 2008. Comparative floral structure ad systematics in Chrysobalanaceae s.l. (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales). Botanical Journal of the Linnean Society 157: 249–309. Patel VC, Skvarla JJ, Raven PH. 1983. Pollen ultrastructure of Chrysobalanaceae. Vidya 26: 1–10. Prance GT, White F. 1988. The genera of Chrysobalanaceae: a study in practical and theoretical taxonomy and its relevance to evolutionary biology. Philosophical Transactions of the Royal Society of London B 320: 1–148. Sothers C, Prance GT, Buerki S, De Kok R, Chase MW. 2014. Taxonomic novelties in Neotropical Chrysobalanaceae: towards a monophyletic Couepia. Phytotaxa 172: 176–200. Yakandawala D, Morton C, Prance GT. 2010. Phylogenetic relationships of the Chrysobalanaceae inferred from chloroplast, nuclear and morphological data. Annals of the Missouri Botanical Garden 97: 259–281. 234. HUMIRIACEAE UMIRI FAMILY Boesewinkel H-D. 1985. The ovule and seed of Humiria balsamifera (Aubl.) St. Hil. Acta Botanica Neerlandica 34: 183–191. Bove CP. 1997. Phylogenetic analysis of Humiriaceae with notes on the monophyly of Ixonanthaceae. Journal of Comparative Biology 2: 19–24. Cuatrecasas J. 1961. A taxonomic revision of Humiriaceae. Contributions of the United States National Herbarium 35, 2: 24–214. Herrera F, Manchester SR, Jaramillo C, MacFadden B, da Silva-Caminha SA. 2010. Phytogeographic history and phylogeny of the Humiriaceae. International Journal of Plant Sciences 171:

392–408 Schultes RE. 1979. Interesting uses of the Humiriaceae in the northwestern Amazon. Journal of Ethnopharmacology 1: 89–94. 235. ACHARIACEAE CHAULMOOGRA FAMILY Chase MW, Zmarzty S, Lledó MD, Wurdack KJ, Swensen SM, Fay MF. 2002. When in doubt, put it in Flacourtiaceae: A molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bulletin 57: 141–181. Groppo M, Fiaschi P, Salatino MLF, Cecchantini GCT, De Assis Ribeiro dos Santos F, Verola CF, Antonelli A. 2010. Placement of Kuhlmanniodendron Fiaschi, Groppo in Lindackerieae (Achariaceae, Malpighiales) comfirmed by analyses of rbcL sequences, with notes on pollen morphology and wood anatomy. Plant Systematics and Evolution 286: 27–37. Obama C, Breteler FJ. 2004. A synopsis of Dasylepis Oliv. (Achariaceae) with a description of a new species from Lower Guinea. Kew Bulletin 59: 585–591. Sosa V, Chase MW, Barcenas C. 2003. Chiangiodendron (Achariaceae): an example of the Laurasian f lora of tropical forests of Central America. Taxon 52: 519–524. 236. VIOLACEAE VIOLET FAMILY Ballard HE Jr, Sytsma KJ, Kowal RR. 1999. Shrinking the violets: phylogenetic relationships of infrageneric groups in Viola (Violaceae) based on internal transcribed spacer DNA sequences. Systematic Botany 23: 439–458. Beattie AJ. 1970. Pollination mechanisms in Viola. New Phytologist 69: 343–360. Hodges SA, Ballard HE Jr, Arnold ML, Chase MW. 1995. Genetic relationships in the Violaceae: data from morphology, anatomy, chromosome numbers and rbcL sequences. American Journal of Botany 82: 136. Munzinger JK, Ballard HE Jr. 2003. Hekkingia (Violaceae), a new arborescent violet from French Guiana, with a key to genera in the family. Systematic Botany 28: 345–351. Tokuoka T. 2008. Molecular phylogenetic analysis of Violaceae (Malpighiales) based on plastic and nuclear DNA sequences. Journal of Plant Research 121: 253–260. Wahlert GA, Marcussen T, De Paula-Souza J, Feng M, Ballard HE Jr. 2014. A phylogeny of the Violaceae (Malpighiales) inferred from plastid DNA sequences: implications for generic diversity and intrafamilial classification. Systematic Botany 39: 239–252. 237. GOUPIACEAE KOPI FAMILY Araujo PAM, Filho AM. 1973. Estrutura de madeira de Goupia glabra Aubl. (Goupiaceae). Archivos do Jardim Botânico do Rio de Janeiro 19: 149–153. Furness CA. 2011. Comparative structure and development of pollen and tapetum in Malpighiales, with a focus on the parietal clade. International Journal of Plant Sciences 172: 980–1011. Lundell CL. Goupia guatemalensis (Celastraceae), a genus and species new to Mesoamerica. Phytologia 57: 238–239. 238. PASSIFLORACEAE PASSIONFRUIT FAMILY Arbo MM, Espert SM. 2009. Morphology, phylogeny and biogeography of Turnera L. (Turneraceae). Taxon 58: 457–467.

Barrett SCH. 1978. Heterostyly in a tropical weed: the reproductive biology of the Turnera ulmifolia complex (Turneraceae). Canadian Journal of Botany 56: 1713–1725. Benitez-Vieyra S, Hempel de Ibarra N, Wertlen AM, Cocucci AA. 2007. How to look like a mallow: evidence of floral mimicry between Turneraceae and Malvaceae. Proceedings of the Royal Society B, 274: 2239–2248. Benson WW, Brown KS Jr, Gilbert LE. 1976. Coevolution of plants and herbivores: passion flower butterflies. Evolution 29: 659–680. De Wilde WJJO. 1971. A monograph of the genus Adenia Forssk. (Passifloraceae). Mededelingen van de Langbouwhogeschool Wageningen 71-18: 1–281. De Wilde WJJO. 1972. The indigenous Old World passifloras. Blumea 20: 227–250. Feuillet C, MacDougal JM. 2004. A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora 13: 34–38. Fordyce JA. 2010. Host shifts and evolutionary radiations of butterflies. Proceedings of the Royal Society B, 277: 3735–3743. Gengler-Nowak K. 2003. Molecular phylogeny and taxonomy of Malesherbiaceae. Systematic Botany 28: 333–444. Hearn DJ. 2006. Adenia (Passif loraceae) and its adaptive radiation: phylogeny and growth form diversification. Systematic Botany 31: 805–821. Thulin M, Razafimandimbison SG, Chafe P, Heidari N, Kool A, Shore JK. 2012. Phylogeny of the Turneraceae clade (Passifloraceae s.l.): transAtlantic disjunctions and two new genera in Africa. Taxon 61: 308–323. Tokuoka T. 2012. Molecular phylogenetic analysis of Passifloraceae sensu lato (Malpighiales) based on plastid and nuclear DNA sequences. Journal of Plant Research 125: 489–497. Ulmer T, MacDougal JM. 2004. Passiflora: passionflowers of the world. Timber Press, Portland. Vanderplank RJR. 2007. Plant/insect mimicry in Passiflora. Passiflora 16: 26–27. 239. LACISTEMATACEAE CEMP-WOOD FAMILY Agostini G. 1973. El genero Lozania Mutis (Lacistemataceae). Acta Botanica Venezuelica 8: 167–175. Alford MH. 2005. Lacistemataceae in Tree of life web project. http://tolweb.org/Lacistemataceae. Chirtoiü M. 1918. Observations sur les Lacistema et la situation systématique de ce genre. Bulletin de la Société Botanique de Genève, séries 2, 10: 317–349. Sleumer HO. 1980. Flacourtiaceae. Flora Neotropica Monograph 22: 182–206. Young FE. 2008 (onwards). Lacistemataceae holistic database. https://www.lacistemataceae.org/ 240. SALICACEAE WILLOW FAMILY Azuma T, Kajita T, Yokoyana J, Ohashi H. 2000. Phylogenetic relationships of Salix (Salicaceae) based on rbcL sequence data. American Journal of Botany 87: 67–75. Boucher LD, Manchester SR, Judd WS. 2003. An extinct genus of Salicaceae based on twigs with attached flowers, fruits and foliage from the Eocene Green River formation of Utah and Colorado, USA. American Journal of Botany 90: 1389–1399. Chase MW, Zmarzty S, Lledó MD, Wurdack KJ, Swensen SM, Fay MF. 2002. When in doubt, put

Plants of the World

713

FURTHER READING it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bulletin 57: 141–181. Chen JH, Sun H, Wen J, Yang YP. 2010. Molecular phylogeny of Salix L. (Salicaceae) inferred from three chloroplast datasets and its systematic implications. Taxon 59: 29–37. Golysheva MD. 1975. Leaf anatomy of Idesia polycarpa Maxim. and other Flacourtiaceae in connection with the problem of affinitive interrelations between the families Salicaceae and Flacourtiaceae. Botanicheskii Zhurnal 60: 787–799. Halle N, De Wilde JJFE. 1978. Trichostephanus acuminatus Gilg. (Flacourtiaceae), un approche biosystematique. Adansonia 18: 167–182. Judd WS. 1997. The Flacourtiaceae in the southeastern United States. Harvard Papers in Botany 1: 65–79. Lemke DE. 1988. A synopsis of Flacourtiaceae. Aliso 12: 29–43. Leskinen E, Alström-Rapaport C. 1999. Molecular phylogeny of Salicaceae and closely related Flacourtiaceae: evidence from 5.8 S, ITS 1 and ITS 2 of the rDNA. Plant Systematics and Evolution 215: 209–227. Meeuse ADJ. 1975. Taxonomic relationships of Salicaceae and Flacourtiaceae, their bearing on interpretaion of floral morphology and dilleniid phylogeny. Acta Botanica Neerlandica 24: 437–457. Newsholme C. 2003. Willows: the genus Salix. Timber Press, Portland. 241. PERACEAE LIGHTNING-BUSH FAMILY Hayden WC, Hayden SM. 2000. Wood anatomy of Acalyphoideae (Euphorbiaceae). IAWA Journal 21: 213–235. Tokuoka T, Tobe H. 2003. Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications. Journal of Plant Research 115: 361–374. Wurdack KL, Davis CC. 2009. Malpighiales phylogenetics: gaining ground on one of the most recalcitrant clades in the angiosperm tree of life. American Journal of Botany 96: 1551–1570. 242. RAFFLESIACEAE CORPSE-FLOWER FAMILY Barkman TJ, McNeal JR, Lim SH, Coat G, Croom HB, Young ND, dePamphilis CW. 2007. Mitochondrial DNA suggests at least 11 origins of parasitism in angiosperms and reveals genomic chimerism in parasitic plants. BMC Evolutionary Biology 7: 248. Bänziger H, Hansen B. 2000. A new taxonomic revision of a deceptive f lower, Rhizanthes Dumortier (Raff lesiaceae). Natural History Bulletin of the Siam Society 48: 117–143. Beaman RS, Decker PJ, Beaman JH. 1988. Pollination of Rafflesia (Rafflesiaceae). American Journal of Botany 75: 1148–1162. Bouman F, Meijer W. 1994. Comparative structure of ovules and seeds in Rafflesiaceae. Plant Systematics and Evolution 193: 187–212. Davis CC, Latvis M, Nickrent DL, Wurdack KL, Baum DA. 2007. Floral gigantism in Rafflesiaceae. Science 315: 1812. Davis CC, Wurdack KJ. 2004. Host-to-parasite gene transfer in flowering plants: phylogenetic evidence from Malpighiales. Science 305: 676–678. Kuijt J. 1969. The biology of parasitic flowering plants. University of California Press, Berkeley. Patiño S, Grace J, Bänziger H. 2000. Endothermy

714

Christenhusz, Fay & Chase

by flowers of Rhizanthes lowii (Rafflesiaceae). Oecologia 124: 149–155. Seymour RS. 2001. Biophysics and physiology of temperature regulation in thermogenic flowers. Bioscience Reports 21: 223–236. 243. EUPHORBIACEAE SPURGE FAMILY Blattner FR, Weising K, Bänfer G, Mascwitz U, Fiala B. 2001. Molecular analysis of phylogenetic relationships among myrmecophytic Macaranga species (Euphorbiaceae). Molecular Phylogenetics and Evolution 19: 331–344. Bollendorff SM, Van Welzen PC, Slik JFW. 2000. A taxonomic revision of Mallotus section Polyadenii (Euphorbiaceae). Blumea 45: 319–340. Bruyns PV, Mapaya RJ, Hedderson T. 2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) in southern Africa based on ITS and psbA-trnH sequence data. Taxon 55: 397–420. Calvin M. 1987. Fuel oils from euphorbs and other plants. Botanical Journal of the Linnean Society 94: 97–110. Carter S. 1994. A preliminary classification of Euphorbia subgenus Euphorbia. Annals of the Missouri Botanical Garden 81: 368–379. Crepet EL, Daglian CP. 1982. Euphorbioid inflorescences from the Middle Eocene Claiborne formation. American Journal of Botany 69: 258–266. Dressler RL. 1957. The genus Pedilanthus (Euphorbiaceae). Contributions from the Gray Herbarium 182: 1–188. Esser HJ, Van Welzen P, Djarwaningsih T. 1997. A phylogenetic classification of the Malesian Hippomaneae (Euphorbiaceae). Systematic Botany 22: 617–628. Gilbert MG. 1994. The relationships of the Euphorbieae. Annals of the Missouri Botanical Garden 81: 283–288. Gillespie LJ. 1994. Pollen morphology and phylogeny of the tribe Plukenetieae (Euphorbiaceae). Annals of the Missouri Botanical Garden 81: 317–348. Horn JW, Van Ee BW, Morawetz JJ, Riina R, Steinmann V, Berry PE, Wurdack KJ. 2012. Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae). Molecular Phylogenetics and Evolution 63: 305–326. Krähenbühl M, Yuan YM, Küpfer P. 2002. Chromosome and breeding system evolution of the genus Mercurialis (Euphorbiaceae): implications of ITS molecular phylogeny. Plant Systematics and Evolution 234: 155–169. Kruijt RC. 1996. A taxonomic monograph of Sapium Jacq., Anomostachys (Baill.) Hurus., Duvigneudia J.Léonard and Scerocroton Hochst. (Euphorbiaceae tribe Hippomaneae). Bibliotheca Botanica 146: 1–109. Kulju KKM, Van der Ham WJM, Breteler FJ. 2008. Rediscovery and phylogenetic position of the incertae sedis genus Afrotrewia (Euphorbiaceae): morphological, pollen and molecular evidence. Taxon 57: 137–143. Prenner G, Rudall PJ. 2007. Comparative ontogeny of the cyathium in Euphorbia (Euphorbiaceae) and its allies: exploring the organ-flower-inflorescence boundary. American Journal of Botany 94: 1612–1629. Schultes R. 1970. The history of taxonomic studies in Hevea. Botanical Review 36: 197–276. Seigler DS. 1994. Phytochemistry and systematics of the Euphorbiaceae. Annals of the Missouri

Botanical Garden 81: 380–401. Tokuoka T. 2007. Molecular phylogenetic analysis of Euphorbiaceae sensu stricto based on plastid and nuclear DNA sequences and ovule and seed character evolution. Journal of Plant Research 120: 511–522. Van Welzen PC, Stuppy W. 1999. Phylogenetic considerations of Euphorbiaceae tribe Aleuritidae. Annals of the Missouri Botanical Garden 86: 894–903. Webster GL. 1994. Classification of the Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 3–32. Westra LYT, Koek-Noorman JK. 2004. Wood atlas of the Euphorbiaceae s.l. IAWA Journal, Supplement 4. Nationaal Herbarium Nederland, Utrecht. Wurdack KJ, Hoffmann P, Chase MW. 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences. American Journal of Botany 92: 1397–1420. 244. LINACEAE FLAX FAMILY Boesewinkel FD. 1984. A comparative SEM study of the seed coat of recent and of 900–1100 years old subfossil linseed. Berichte der Deutche Botanische Gesellschaft 97: 443–450. Dulberger R. 1974. Structural dimorphism of stigmatic papillae in distylous Linum species. American Journal of Botany 61: 238–243. McDill JR, Repplinger M, Simpson BB, Kadereit JW. 2009. The phylogeny of Linum and Linaceae subfamily Linoideae, with implications for their systematics, biogeography, and evolution of heterostyly. Systematic Botany 34: 386–405. McDill JR, Simpson BB. 2011. Molecular phylogenetics of Linaceae with complete sampling and data from two plastid genes. Botanical Journal of the Linnean Society 165: 64–83. Muir AD, Westcott ND (eds). 2003. Flax — the genus Linum. Taylor, Francis, London, New York. Saad SI. 1962. Palynological studies in the Linaceae. Pollen and Spores 4: 65–82. Thompson JD, Pailler T, Strasberg D, Manicacci D. 1996. Tristyly in the endangered Mascarene Island endemic Hugonia serrata (Linaceae). American Journal of Botany 83: 1160–1167. Van Welzen PC, Baas P. 1984. A leaf anatomical contribution to the classification of the Linaceae complex. Blumea 29: 453–479. 245. IXONANTHACEAE TWENTYMEN-TREE FAMILY Byng JW, Bernardini B, Christenhusz MJM, Chase MW. 2016. Systematics of Irvingiaceae and Ixonanthaceae (Malpighiales): phylogenetic analysis based on three plastid DNA loci. Phytotaxa 260: 157–166. Forman LL. 1965. A new genus of Ixonanthaceae with notes on the family. Kew Bulletin 19: 517–526. Kool R. 1988. A taxonomic revision of the genus Ixonanthes (Linaceae). Blumea 26: 191–204. Nooteboom HP. 1967. The taxonomic position of Irvingioideae, Allantospermum Forman, and Cyrillopsis Kuhlm. Adansonia II, 7: 161–168. Van Welzen PC, Baas P. 1984. A leaf anatomical contribution to the classification of the Linaceae complex. Blumea 29: 453–479. 246. PICRODENDRACEAE HOLLYSPURGE FAMILY Berg R. 1975. Fruit, seed and myrmecochorous dispersal in Micrantheum (Euphorbiaceae). Norwegian Journal of Botany 22: 173–194.

FURTHER READING Hayden WJ. 1994. Systematic anatomy of Euphorbiaceae subfamily Oldfieldioideae. I. Overview. Annals of the Missouri Botanical Garden 81: 180–202. Hayden WJ, Gillis WT, Stone DE, Broome CR, Webster GL. 1984. Systematics and palynology of Picrodendron: further evidence for relationshops with the Oldfieldioideae (Euphorbiaceae). Journal of the Arnold Arboretum 65: 105–127. Levin GA. 1992. Systematics of Paradrypetes (Euphorbiaceae). Systematic Botany 17: 74–83. Levin GA, Simpson MG. 1994. Phylogenetic implications of pollen ultrastructure in the Oldfieldioideae (Euphorbiaceae). Annals of the Missouri Botanical Garden 81: 239–244. Merino Sutter D, Forster PI, Endress PK. 2006. Female flowers and systematic position of Picrodendraceae (Euphorbiaceae s.l., Malpighiales). Plant Systematics and Evolution 261: 187–215. Van Welzen PC, Forster PI. 2011. Picrodendraceae (formerly Euphorbiaceae s.l. subfam. Oldfieldioideae). Flora Malesiana ser. 1, vol. 20: 45–61. 247. PHYLLANTHACEAE LEAFFLOWER FAMILY Hoffmann P, Kathriarachchi H, Wurdack KJ. 2006. A phylogenetic classification of Phyllanthaceae (Malpighiales; Euphorbiaceae sensu lato). Kew Bulletin 61: 37–53. Hoffmann P, McPherson G. 2007. Revision of Wielandia, including Blotia and Petalodiscus (Phyllanthaceae; Euphorbiaceae s.l.). Annals of the Missouri Botanical Garden 94: 519–553. Kathriarachchi H, Hoffmann P, Samuel R, Wurdack KJ, Chase MW. 2005. Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3’ndhF, rbcL, and nuclear PHYC). Molecular Phylogenetics and Evolution 36: 112–134. Kawakita A, Kato M. 2004. Evolution of obligate pollination mutualism in New Caledonian Phyllanthus (Euphorbiaceae). American Journal of Botany 91: 410–415. Li Y, Dressler S, Zhang D, Renner SS. 2009. More Miocene dispersal between Africa and Asia — the case of Bridelia (Phyllanthaceae). Systematic Botany 34: 521–529. Vorontsova MS, Hoffmann P, Maurin O, Chase MW. 2007. Molecular phylogenetics of tribe Poranthereae (Phyllanthaceae; Euphorbiaceae sensu lato). American Journal of Botany 94: 2026–2040. Wurdack KJ, Hoffmann P, Samuel R, De Bruijn A, Van der Bank M, Chase MW. 2004. Molecular phylogenetic analysis of Phyllanthaceae (Phyllanthoideae pro parte, Euphorbiaceae sensu lato) using plastid rbcL DNA sequences. American Journal of Botany 91: 1882–1900. 248. GERANIACEAE CRANE’S-BILL FAMILY Albers F. 1996. The taxonomic status of Sarcocaulon (Geraniaceae). South African Journal of Botany 62: 345–347. Aldasoro JJ, Navarro C, Vargas P, Saez L, Aedo C. 2002. California, a new genus of Geraniaceae endemic to the southwest of North America. Anales del Jardín Botánico de Madrid 59: 209–216. Bakker FT, Culham A, Hettiarachi P, Touloumenidou T, Gibby M. 2004. Phylogeny of Pelargonium (Geraniaceae) based on DNA sequences from three genomes. Taxon 53: 17–28. Boesewinkel FD. 1988. The seed structure and

taxonomic relationships of Hypseocharis Remy. Acta Botanica Neerlandica 37: 111–120. Fiz O, Vargas P, Alarcón ML, Aedo M, Garcia JL, Aldasoro JJ. 2008. Phylogeny and historical biogeography of Geraniaceae in relation to climate change and pollination ecology. Systematic Botany 33: 326–342. Gama-Arachchige NS, Baskin JM, Geneve RL, Baskin CC. 2010. Identification and characterisation of the water gap in physically dormant seeds of Geraniaceae, with special reference to Geranium carolinianum. Annals of Botany 105: 977–990. Price RA, Palmer JD. 1993. Phylogenetic relationships of the Geraniaceae and Geraniales from rbcL sequence comparisons. Annals of the Missouri Botanical Garden 80: 661–571. Spomer GG. 1999. Evidence of protocarnivorous capabilities in Geranium viscosissimum and Potentilla arguta and other sticky plants. International Journal of Plant Sciences 160: 98–101. Struck M. 1997. Floral divergence and convergence in the genus Pelargonium (Geraniaceae) in southern Africa: ecological and evolutionary considerations. Plant Systematics and Evolution 71: 79–109. Van der Walt JJA, Vorster PJ. 1983. Phytogeography of Pelargonium. Bothalia 14: 517–523. Vorster PJ (ed.). 1990. Proceedings of the 1st international Geraniaceae symposium. Stellenbosch University, Cape Town. 249. FRANCOACEAE BRIDAL-WREATH FAMILY Boesewinkel FD. 1999. Seed structure and phylogenetic relationships of the Geraniales. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 119: 277–291. Carlquist S. 1986. Wood anatomy and familial status of Viviania. Aliso 11: 159–165. Palazzesi L, Gottschling M, Barreda V, Weigend M. 2012. First Miocene fossils of Vivianiaceae shed new light on phylogeny, divergence times, and historical biogeography of Geraniales. Biological Journal of the Linnean Society 107: 67–85. Ronse Decraene LP, Smets EF. 1999. Similarities in floral ontogeny and anatomy between the genera Francoa (Francoaceae) and Greyia (Greyiaceae). International Journal of Plant Sciences 160: 377–393. Ronse Decraene LP, Linder HP, Dlamini T, Smets EF. 2001. Evolution and development of floral diversity of Melanthiaceae, an enigmatic family from southern Africa. International Journal of Plant Sciences 162: 59–82. Sytsma KJ, Spalink D, Berger B. 2014. Calibrated chronograms, fossils, outgroup relationships, and root priors: re-examining the historical biogeography of Geraniales. Biological Journal of the Linnean Society 113: 29–49. Weigend M. 2005. Notes on the floral morphology in Vivianiaceae. Plant Systematics and Evolution 253: 125–131. 250. COMBRETACEAE BUSHWILLOW FAMILY Exell AW, Stace CA. 1966. Revision of the Combretaceae. Boletim da Sociedade Broteriana II 40: 5–25. Friis EM, Pedersen KR, Crane PR. 1992. Esgueiria gen. nov., fossil f lowers with combretaceous features from the Late Cretaceous of Portugal. Biologiske Skrifter 41: 1–45. Jongkind CCH. 1995. Review of the genus Strephonema (Combretaceae). Annals of the Missouri Botanical Garden 82: 535–541.

Manchester SR, O’Leary EL. 2010. Phylogenetic distribution and identification of fin-winged fruits. Botanical Review 76: 1–82. Maurin O, Chase MW, Jordaan M, Van der Bank M. 2010. Phylogenetic relationships of Combretaceae inferred from nuclear and plastid DNA sequence data: implications for generic classification. Botanical Journal of the Linnean Society 162: 453–476. Ohri D. 1966. Genome size and polyploidy variation in the tropical hardwood genus Terminalia (Combretaceae). Plant Systematics and Evolution 200: 225–232. Stace C. 1981. The significance of the leaf epidermis in the taxonomy of the Combretaceae: conclusions. Botanical Journal of the Linnean Society 81: 327–339. Stace C. 2010. Combretaceae. Flora Neotropica monograph 107. New York Botanical Garden, New York. Tan FX, Shi SH, Zhong Y, Gong X, Wang YG. 2002. Phylogenetic relationships of Combretoideae (Combretaceae) inferred from plastid, nuclear gene and spacer sequences. Journal of Plant Research 115: 475–481. Van Vliet CJCM. 1979. Wood anatomy of the Combretaceae. Blumea 25: 141–223. 251. LYTHRACEAE POMEGRANATE FAMILY Baas P, Zweypfenning RCVJ. 1979. Wood anatomy of the Lythraceae. Acta Botanica Neerlandica 28: 117–155. Dulberger R. 1970. Tristyly in Lythrum junceum. New Phytologist 69: 751–759. Graham A, Graham SA. 1971. The geologic history of the Lythraceae. Brittonia 23: 335–346. Graham SA. 1964. The genera of Lythraceae in the southeastern United States. Journal of the Arnold Arboretum 45: 235–250. Graham SA. 2013. Fossil records in the Lythraceae. Botanical Review 79: 48–145. Graham SA, Hall J, Sytsma K, Shi SH. 2005. Phylogenetic analysis of the Lythraceae based on four gene regions and morphology. International Journal of Plant Sciences 166: 995–1017. Malone MH, Rother A. 1994. Heimia salicifolia: a phytochemical and phytopharmacologic review. Journal of Ethnopharmacology 42: 135–159. 252. ONAGRACEAE FUCHSIA FAMILY Eyde RH. 1982. Evolution and systematics of Onagraceae: floral anatomy. Annals of the Missouri Botanical Garden 69: 735–747. Ford VS, Gottlieb LD. 2007. Tribal relationships within Onagraceae inferred from PgiC sequences. Systematic Botany 32: 348–356. Levin RA, Wagner WL, Hoch PC, Nepokroeff M, Pires JC, Zimmer EA, Sytsma KJ. 2003. Familylevel relationships of Onagraceae based on chloroplast rbcL and ndhF data. American Journal of Botany 90: 107–115. Leven RA, Wagner WL, Hoch PC, Hahn WJ, Rodriguez A, Baum DA, Katinas L, Zimmer EA, Sytsma KJ. 2004. Paraphyly in tribe Onagraceae: insights into phylogenetic relationships of Onagraceae based on nuclear and chloroplast sequence data. Systematic Botany 29: 147–164. Wagner WL, Hoch PC. 2005 (onwards). Onagraceae, the evening primrose family website. http://botany. si.edu/onagraceae/index.cfm Wagner WL, Hoch PC, Raven PH. 2007. Revised classification of the Onagraceae. Systematic Botany Monographs 83.

Plants of the World

715

FURTHER READING 253. VOCHYSIACEAE QUARUBA FAMILY Litt A. 1996. Phylogeny of the Vochysiaceae: implications of molecular data for floral evolution. American Journal of Botany 83: 175. Quirk JT. 1980. Wood anatomy of the Vochysiaceae. IAWA Bulletin 1, 4: 172–179. Stafleu FA. 1948, 1952, 1953. A monograph of the Vochysiaceae. I, II, III, IV. Recueil des Travaux Botaniques Néerlandais 41: 398–540; Acta Botanica Neerlandica 1: 222–242; 2: 144–217; 3: 459–480. Sytsma KJ, Litt A, Zjhra ML, Pires JC, Nepokroeff M, Conti E, Walker J, Wilson PG. 2004. Clades, clocks, and continents: historical and biogeographical analysis of Myrtaceae, Vochysiaceae, and relatives in the Southern Hemisphere. International Journal of Plant Sciences 165: S85–S105. 254. MYRTACEAE MYRTLE FAMILY Basinger JF, Greenwood DR, Wilson PG, Christophel DC. 2007. Fossil flowers and fruits of capsular Myrtaceae from the Eocene of South Australia. Canadian Journal of Botany 85: 204–215. Bayly MJ, Rigault P, Spokevicius A, Ladiges PY, Ades PK, Anderson C, Bossinger G, Merchant A, Udovicic F, Woodrow IE, Tibbits J. 2013. Chloroplast genome analysis of Australian eucalypts — Eucalyptus, Corymbia, Angophora and Stockwellia (Myrtaceae). Molecular Phylogenetics and Evolution 69: 704–716. Biffin E, Craven LA, Crisp MD, Gadek PA. 2006. Molecular systematics of Syzygium and allied genera (Myrtaceae): evidence from the chloroplast genome. Taxon 55: 79–94. Biffin E, Lucas EJ, Craven LA. Ribeiro da Costa I, Harrington MG, Crisp MD. 2010. Evolution of exceptional species richness among lineages of fleshy-fruited Myrtaceae. Annals of Botany 106: 79–93. Briggs BG, Johnson LAS. 1979. Evolution in the Myrtaceae–evidence from inflorescence structure. Proceedings of the Linnean Society of New South Wales 102: 157–256. Brokker MIH, Nicolle D. 2013. Atlas of leaf venation and oil gland patterns in eucalypts. CSIRO Publishing, Collingwood. Coppen JJW (ed.) 2002. Eucalyptus: The genus Eucalyptus. Taylor and Francis, London. Craven LA. 2006. New combinations in Melaleuca for Australian species of Callistemon (Myrtaceae). Novon 16: 468–475. Edwards RD, Craven LA, Crisp MD, Cook LG. 2010. Melaleuca revisited: cpDNA and morphological data confirm that Melaleuca L. is not monophyletic. Taxon 59: 744–754. Gandolfo MA, Hermsen EJ, Zamaloa MC, Nixon KC, González CC, Wilf P, Cúneo NR, Johnson KR. 2011. Oldest known Eucalyptus macrofossils are from South America. PLoS ONE 6: e21084. Hermsen EJ, Gandolfo MA, Zamaloa MC. 2012. The fossil record of Eucalyptus in Patagonia. American Journal of Botany 99: 1356–1374. Johnson LAS, Briggs BG. 1985. Myrtales and Myrtaceae — a phylogenetic analysis. Annals of the Missouri Botanical Garden 71: 700–756. Ladiges P, Parra-O. C, Gibbs A, Udovicic F, Nelson G, Bayly M. 2011. Historical biogeographical patterns in continental Australia: congruence among areas of endemism of two major clades of eucalypts. Cladistics 27: 29–41. Landrum LR, Kawasaki ML. 1997. The genera of Myrtaceae in Brazil: an illustrated synoptic treatment and identification keys. Brittonia 49:

716

Christenhusz, Fay & Chase

508–536. Lucas EJ, Harris SA, Mazine FF, Belsham SR, Nic Lughadha EM, Telford A, Gasson PE, Chase MW. 2007. Suprageneric phylogenetics of Myrteae, the generically richest tribe in Myrtaceae (Myrtales). Taxon 56: 1105–1128. Lucas EJ, Matsumoto K, Harris SA, Nic Lughadha EM, Bernadini N, Chase MW. 2011. Phylogenetics, morphology, and evolution of the large genus Myrcia s.l. (Myrtaceae). International Journal of Plant Sciences 172: 915–934. Manchester SR, Dilcher DL, Wing SL. 1998. Attached leaves and fruits of myrtaceous affinity from the Middle Eocene of Colorado. Review of Palaeobotany and Palynology 102: 153–163. McKinnon GE, Vaillancourt RE, Steane DA, Potts BM. 2008. An AFLP marker approach to lowerlevel systematics in Eucalyptus (Myrtaceae). American Journal of Botany 95: 368–380. Parra-O C, Bayly MJ, Drinnan A, Udivicic F, Ladiges P. 2009. Phylogeny, major clades and infrageneric classification of Corymbia (Myrtaceae), based on nuclear ribosomal DNA and morphology. Australian Systematic Botany 22: 384–399. Rye BL, James SH. 1992. The relationship between dysploidy and reproductive capacity in Myrtaceae. Australian Journal of Botany 40: 829–848. Schmid R. 1980. Comparative anatomy and morphology of Psiloxylon and Heteropyxis, and the subfamilial and tribal classification of Myrtaceae. Taxon 29: 559–595. Schmid R, Baas P. 1984. The occurrence of scalariform perforation plates and helical vessel wall thickenings in wood of Myrtaceae. IAWA Bulletin 5: 197–215. Sytsma KJ, Litt A, Zjhra ML, Pires JC, Nepokroeff M, Conti E, Walker J, Wilson PG. 2004. Clades, clocks, and continents: historical and biogeographical analysis of Myrtaceae, Vochysiaceae, and relatives in the Southern Hemisphere. International Journal of Plant Sciences 165: S85–S105. Thornhill AH, Popple LW, Carter RJ, Ho SYW, Crisp MD. 2012. Are pollen fossils useful for calibrating relaxed molecular clock dating of phylogenies? A comparative study using Myrtaceae. Molecular Phylogenetics and Evolution 63: 15–27. Thornhill AH, Macphail M. 2012. Fossil myrtaceous pollen as evidence for the evolutionary history of Myrtaceae: a review of fossil Myrtaceidites species. Review of Palaeobotany and Palynology 176–177: 1–23. Van der Merwe MM, Van Wyk AE, Botha AM. 2005. Molecular phylogenetic analysis of Eugenia L. (Myrtaceae) with emphasis on southern African taxa. Plant Systematics and Evolution 251: 21–34. Wilson PG, O’Brien MM, Heslewood MM, Quinn CJ. 2005. Relationships within Myrtaceae sensu lato based on a matK phylogeny. Plant Systematics and Evolution 251: 3–19. Wright SD, Yong CG, Dawson JW, Whittaker DJ, Gardner RC. 2000. Riding the ice age El Niño? Pacific biogeography and evolution of Metrosideros subg. Metrosideros (Myrtaceae) inferred from nuclear ribosomal DNA. Proceedings of the National Academy of Sciences of the USA 97: 4118–4123.

Botany 88: 486–498. Goldenberg R, Penneys DS, Almeda F, Judd WS, Michelangeli FA. 2008. Phylogeny of Miconia (Melastomataceae): patterns of stamen diversification in a megadiverse Neotropical genus. International Journal of Plant Sciences 26: 963–978. Groenendijk JP, Bouman F, Cleef AM. 1996. An exploratory study on the seed morphology of Miconia Ruiz and Pavon (Melastomataceae), with taxonomic and ecological implications. Acta Botanica Neerlandica 45: 323–344. Martin CV, Little D, Goldenberg R, Michelangeli FA. 2008. A phylogenetic evaluation of Leandra (Miconieae, Melastomataceae): a polyphyletic genus where the seeds tell the story, not the petals. Cladistics 24: 315–327. Martin CV, Michelangeli FA. 2009. Comparative seed morphology of Leandra (Miconieae, Melastomataceae). Brittonia 61: 175–188. Melastomataceae.Net. 2007–2014. A site with information on the biodiversity of Melastomataceae. https://melastomataceae.net/ Michelangeli FA, Penneys DS, Giza J, Soltis D, Hils MH, Skean JD Jr. 2004. A preliminary phylogeny of the tribe Miconieae (Melastomataceae) based on nrITS sequence data and its implications on inflorescence position. Taxon 53: 279–290. Michelangeli FA, Guimaraes PJF, Penneys DS, Almeda F, Kriebel R. 2012. Phylogenetic relationships and distribution of New World Melastomeae (Melastomataceae). Botanical Journal of the Linnean Society 171: 38–60. Morley T. 1976. Memecyleae (Melastomataceae). Flora Neotropica Monograph 15. Penneys DS, Judd WS. 2005. A systematic revision and cladistic analysis of Charianthus (Melastomataceae) using morphological and molecular characters. Systematic Botany 30: 559–584. Penneys DS, Judd WS. 2013. Combined molecular and morphological phylogenetic analysis of the Blakeeae (Melastomataceae). International Journal of Plant Sciences 174: 802–817. Renner SS. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic Journal of Botany 13: 519–540. Renner SS. 2004. Multiple Miocene Melastomataceae dispersals between Madagascar, Africa and India. Philosophical Transactions of the Royal Society B 359: 1485–1494. Renner SS, Meyer K. 2001. Melastomataceae come full circle: biogeographic reconstruction and molecular clock dating. Evolution 55: 1315–1324. Ruokolainen K, Linna A, Tuomisto H. 1997. Use of Melastomataceae and pteridophytes for revealing phytogeographical patterns in Amazonian rain forests. Journal of Tropical Ecology 13: 243–256. Schulman L, Hyvönen J. 2003. A cladistic analysis of Adelobotrys (Melastomataceae) based on morphology, with notes on generic limits within the tribe Merianieae. Systematic Botany 28: 738–756. Stone RD, Andreasen K. 2010. The Afro-Madagascan genus Warneckea (Melastomataceae): molecular systematics and revised infrageneric classification. Taxon 59: 83–92.

255. MELASTOMATACEAE SENDUDUK FAMILY Clausing G, Renner SS. 2001. Molecular phylogenetics of Melastomataceae and Memecylaceae: implications for character evolution. American Journal of

256. CRYPTERONIACEAE BEKOI FAMILY Conti E, Rutschmann F, Eriksson T, Sytsma K, Baum D. 2004. Calibration of molecular clocks and the biogeographic history of Crypteroniaceae: a reply to Robert G. Moyle. Evolution 58: 1874–1876.

FURTHER READING Moyle RG. 2004. Calibration of molecular clocks and the biogeographic history of Crypteroniaceae. Evolution 58: 1869–1871. Rutschman F, Eriksson T, Conti E. 2004. Did Crypteroniaceae really disperse out of India? Molecular dating evidence from rbcL, ndhF and rpl16 intron sequences. International Journal of Plant Sciences 165: S69–S83. Tone H, Raven PH. 1983. The embryology of Axinandra zeylanica (Crypteroniaceae) and the relationships of the genus. Botanical Gazette 144: 426–432. Tone H, Raven PH. 1987. The embryology and relationships of Crypteronia (Crypteroniaceae). Botanical Gazette 148: 96–102. Tone H, Raven PH. 1987. The embryology and relationships of Dactylocladus (Crypteroniaceae) and a discussion of the family. Botanical Gazette 148: 103–111. Van Beusekom-Osinga RJ, Van Beusekom CF. 1975. Delimitation and subdivision of the Crypteroniaceae (Myrtales). Blumea 22: 255–266. Van Vliet GJCM. 1975. Wood anatomy of Crypteroniaceae sensu lato. Journal of Microscopy 104: 65–82. 257. ALZATEACEAE WANTSUM FAMILY Baas P. 1979. The anatomy of Alzatea Ruiz, Pav. (Myrtales). Acta Botanica Neerlandica 28: 156–158. Graham SA. 1984. Alzateaceae, a new family of Myrtales in the American tropics. Annals of the Missouri Botanical Garden 71: 757–779. Lourteig A. 1965. On the systematic position of Alzatea verticillata R., P. Annals of the Missouri Botanical Garden 52: 371–378. Si l ve r s t o n e - S o p k i n PA , Graham SA . 1986. Alzateaceae, a plant family new to Colombia. Brittonia 38: 340–343. 258. PENAEACEAE CAPE-FELLWORT FAMILY Carlquist S, Debuhr L. 1977. Wood anatomy of Penaeaceae (Myrtales): comparative, phylogenetic, and ecological implications. Botanical Journal of the Linnean Society 75: 211–227. Mújica MB, Cutler DF. 1974. Taxonomic implications of anatomical studies on the Oliniaceae. Kew Bulletin 29: 93–123. Rao VS, Dahlgren R. 1969. The floral anatomy and relationships of Oliniaceae. Botaniska Notiser 122: 160–171. Schönenberger J, Conti E. 2003. Molecular phylogeny and floral evolution of Pennaeaceae, Oliniaceae, Rhynchocalycaceae, and Alzateaceae (Myrtales). American Journal of Botany 90: 293–309. Strey RG, Leistner OA. 1968. The rediscovery of Rhynchocalyx lawsonioides Oliv. Journal of South African Botany 34: 9–13. Tobe H, Raven PH. 1984. The embryology and relationships of Rhynchocalyx Oliv. (Rhynchocalycaceae). Annals of the Missouri Botanical Garden 71: 836–843. 259. APHLOIACEAE MOUNTAIN-PEACH FAMILY Matthews ML, Endress PK. 2005. Comparative floral structure and systematics in Crossosomatales (Crossosomataceae, Stachyuraceae, Staphyleaceae, Aphloiaceae, Geissolomataceae, Ixerbaceae, Strasburgeriaceae. Botanical Journal of the Linnean Society 147: 1–46. Miller RB. 1975. Systematic anatomy of the xylem and comments on the relationships of Flacourtiaceae.

Journal of the Arnold Arboretum 56: 20–102. Oh SH. 2010. Phylogeny and systematics of Crossosomatales as inferred from chloroplast atpB, matK, and rbcL sequences. Korean Journal of Plant Taxonomy 40: 208–217. 260. GEISSOLOMATACEAE CAPE-CUPS FAMILY Carlquist S. 1975. Wood anatomy and relationships of the Geissolomataceae. Bulletin of the Torrey Botanical Club 102: 128–134. Carlquist S. 1990. Leaf anatomy of Geissolomataceae and Myriothamnaceae as possible indicator of relationships to Bruniaceae. Bulletin of the Torrey Botanical Club 117: 420–428. Dahlgren R, Rao VS. 1969. A study of the family Geissolomataceae. Botaniska Notiser 122: 207–227. McDonald DJ. 1998. The enigma of the Geissolomataceae. Veld, Flora 84: 122–123. 261. STRASBURGERIACEAE TAWARI FAMILY Cameron KM. 2003. On the phylogenetic position of the New Caledonian endemic families Pa r a c r y ph ia c e a e, O nc ot he ca c e a e, a nd Strasburgeriaceae: a comparison of molecules and morphology. Botanical Review 68: 428–443. Dickison WC. 1981. Contributions to the morphology and anatomy of Strasburgeria and a discussion of the taxonomic position of the Strasburgeriaceae. Brittonia 33: 564–580. Jarzen DM, Pocknall DT. 1993. Tertiary Bluffopollis scabratus (Cooper) Pocknall, Mildenhall, 1984, and modern Strasburgeria pollen: a botanical comparison. New Zealand Journal of Botany 31: 185–192. Matthews ML, Endress PK. 2005. Comparative floral structure and systematics in Crossosomatales (Crossosomataceae, Stachyuraceae, Staphyleaceae, Aphloiaceae, Geissolomataceae, Ixerbaceae, Strasburgeriaceae. Botanical Journal of the Linnean Society 147: 1–46. 262. STAPHYLEACEAE BLADDERNUT FAMILY Carlquist SA, Hoekman DA. 1985. Wood anatomy of Staphyleaceae: ecology, statistical correlations, and systematics. Flora 177: 195–216. Dickison WC. 1986. Floral morphology and anatomy of Staphyleaceae. Botanical Gazette 147: 312–326. Dickison WC. 1987. Leaf and nodal anatomy and systematics of Staphyleaceae. Botanical Gazette 148: 475–489. Gadek PA, Fernando ES, Quinn CJ, Hoot BS, Terrazas T, Sheahan MC, Chase MW. 1996. Sapindales: molecular delimitation and infraordinal groups. American Journal of Botany 83: 802–811. Simmons SL, Panero JL. 2000. Phylogeny and biogeography of Staphyleaceae (DC.) Lindl. American Journal of Botany 87 (6, suppl.): 157. Tiffney BH. 1979. Fruits and seeds of the Brandon Lignite III. Turpinia (Staphyleaceae). Brittonia 31: 39–51. Weaver RE. 1980. The bladdernuts. Arnoldia 40: 76–93. 263. GUAMATELACEAE GUATEMALANBRAMBLE FAMILY Oh SH, Potter D. 2006. Description and phylogenetic position of a new angiosperm family, Guamatelaceae, inferred from chloroplast rbcL, atpB, and matK sequences. Systematic Botany 31: 730–738.

264. STACHYURACEAE SPIKETAIL FAMILY Chen SK. 1981. A study on the Stachyuraceae from China. Acta Botanica Yunnanica 3: 125–137. Sosa V, Chase MW. 2003. Phylogenetics of Crossosomataceae based on rbcL sequence data. Systematic Botany 28: 96–105. Tang YC, Cao YL, Xi YZ, He J. 1983. Systematic studies of Chinese Stachyuraceae, I. phytogeographical, cytological, palynological. Acta Phytotaxonomica Sinica 21: 236–253. Zhu YP, Wen J, Zhang ZY, Chen ZD. 2006. Evolutionary relationships and diversification of Stachyuraceae based on sequences of four chloroplast markers and the nuclear ribosomal ITS region. Taxon 55: 931–940. 265. CROSSOSOMATACEAE ROCKFLOWER FAMILY Calderón G, Rzedowski J. 1997. Velascoa (Crossosomataceae), un género nuevo de la Sierra Madre Oriental de México. Acta Botanica Mexicana 39: 53–59. DeBuhr L. 1978. Wood anatomy of Forsellesia (Glossopetalon) and Crossosoma (Crossosomataceae, Rosales). Aliso 9: 179–184. Holmgren NH. 1988. Glossopetalon (Crossosomataceae) and a new variety of G. spinescens from the Great Basin, U.S.A. Brittonia 40: 269–274. Mason CT. 1992. Crossosomataceae: Crossosoma family. Journal of the Arizona and Nevada Academy of Science 26: 7–9. Richardson PE. 1970. The morphology of the Crossosomataceae. I. Leaf, stem, and node. Bulletin of the Torrey Botanical Club 97: 34–39. Sosa V. Chase MW. 2003. Phylogenetics of Crossosomataceae based on rbcL sequence data. Systematic Botany 28: 96–105. 266. PICRAMNIACEAE BITTERBUSH FAMILY Fernando ES, Quinn CJ. 1995. Picramniaceae, a new family, and a recircumscription of Simaroubaceae. Taxon 44: 177–181. Jacobs H. 2003. Comparative phytochemistry of Picramnia and Alvaradoa, genera of the newly established family Picramniaceae. Biochemical Systematics and Ecology 31: 773–783. Pirani JR. 1990. As espécies de Picramnia Sw. (Simaroubaceae) do Brasil: uma sinopse. Boletim da Universidade de São Paulo. Botanica 12: 115–180. Pirani JR. 1993. Inflorescence morphology and evolution in the genus Picramnia (Simaroubaceae). Candollea 48: 119–135. Thomas WW. 2011. Nothotalisia, a new genus of Picramniaceae from tropical America. Brittonia 63: 51–61. 267. BIEBERSTEINIACEAE KHARDUG FAMILY Bakker FT, Vassiliades DD, Morton C. 1998. Phylogenetic position of Biebersteinia orphanidis Boiss. (Geraniaceae) inferred from rbcL and atpB sequence comparisons. Botanical Journal of the Linnean Society 127: 149–158. Greenham J, Vassiliades DD, Harborne JB, Williams CA, Eagles J, Grayer RJ, Veitch NC. 2001. A distinctive flavonoid chemistry for the anomalous genus Biebersteinia. Phytochemistry 56: 87–91. Liu JQ, Ho TN, Chen SL, Lu AM. 2001. Karyomorphology of Biebersteinia Stephan (Geraniaceae) and its systematic and taxonomic significance. Botanical Bulletin of Academia Sinica (Taipei) 42: 61–66.

Plants of the World

717

FURTHER READING Muellner AN, Vassiliades DD, Renner SS. 2007. Placing Biebersteiniaceae, a herbaceous clade of Sapindales, in a temporal and geographic context. Plant Systematics and Evolution 266: 233–252. Zhang XF, Hu BL, Zhou BN. 1995. Studies on the active constituents of Tibetan herb Biebersteinia heterostemon Maxim. Acta Pharmaceutica Sinica 30: 211–214. 268. NITRARIACEAE NITREBUSH FAMILY Chevalier A. 1949. Les Nitraria, plantes utiles des déserts salés. Revue de Botanique Appliquée et d’Agriculture Tropicale 29: 595–601. Gadek P. Fernando ES, Quinn CJ, Hoot SB, Terrazas T, Sheahan MC, Chase MW. 1996. Sapindales: molecular delimitation and infraordinal groups. American Journal of Botany 83: 802–811. Elgin N. 1950. Les caractères morphologiques et anatomiques du Peganum harmala L. (Zygophyllaceae). Revue de la Faculté des sciences de l’Université d’Istanbul B 15: 333–361. Li S, Tu L. 1994. The embryology and its systematic significance of Nitraria. Bulletin of Botanical Research, Harbin 14: 255–262. Kamelina OP. 1994. Embryology and systematic position of Tetradiclis (Tetradiclidaceae). Botanicheskii Zhurnal 79: 11–27. Pan XL, Shen GM, Chen P. 1999. A preliminary research on taxonomy and systematics of genus Nitraria. Acta Botanica Yunnanica 21: 287–295. Ronse Decraene LP, De Laet J, Smets EF. 1996. Morphological studies in Zygophyllaceae II. The floral development and vascular anatomy of Peganum harmala. American Journal of Botany 83: 201–215. Sheahan MC, Chase MW. 1996. A phylogenetic analysis of Zygophyllaceae R.Br. based on morphological, anatomical and rbcL DNA sequence data. Botanical Journal of the Linnean Society 113: 227–262. Singh BP, Kaur I. 1998. Systematic position of the genus Peganum. Journal of Economic and Taxonomic Botany 22: 705–708. 269. KIRKIACEAE WHITE-SERINGA FAMILY Bachelier JB, Endress PK. 2008. Floral structure of Kirkia (Kirkiaceae) and its position in Sapindales. Annals of Botany 102: 539–550. Fernando ES, Gadek PA, Quinn CJ. 1995. Simaroubaceae, an artificial construct: evidence from rbcL sequence variation. American Journal of Botany 82: 92–103. Jadin F. 1901. Contribution à l’étude des Simaroubacées. Annales des Sciences Naturelles, Botanique Séries 8: 201–304. Stannard BL. 1981. A revision of Kirkia (Simaroubaceae). Kew Bulletin 35: 829–839. Stannard BL. 2007. The inclusion of Pleiokirkia in Kirkia (Kirkiaceae), and corresponding combination. Kew Bulletin 62: 151–152. 270. BURSERACEAE FRANKINCENSE-ANDMYRRH FAMILY Bachelier JB, Endress, PK. 2009. Comparative floral morphology and anatomy of Anacardiaceae and Burseraceae (Sapindales), with special focus on gynoecium structure and evolution. Botanical Journal of the Linnean Society 159: 499–571. Becerra JX, Venable DL, Evans PH, Bowers WS. 2001. Interactions between chemical and mechanical defences in the plant genus Bursera and their implications for herbivores. American Zoologist

718

Christenhusz, Fay & Chase

41: 865–876. Becerra JX, Noge K, Olivier S, Venable DL. 2012. The monophyly of Bursera and its impact for divergence times of Burseraceae. Taxon 61: 333–343. Clarkson JJ, Chase MW, Harley MM. 2002. Phylogenetic relationships in Burseraceae based on plastid rps16 intron sequences. Kew Bulletin 57: 183–193. Daly DC. 1989. Studies in Neotropical Burseraceae II. Generic limits in Neotropical Protieae and Canarieae. Brittonia 41: 17–27. Daly DC. 1999. Notes on Trattinnickia, including a synopsis in eastern Brazil’s Atlantic forest complex. Studies in Neotropical Burseraceae IX. Kew Bulletin 54: 129–137. De-Nova JA, Medina R, Montero JC, Weeks A, Rosell JA, Olson ME, Eguiarte LE, Magallón S. 2012. Insights into the historical construction of speciesrich Mesoamerican seasonally dry tropical forests: the diversification of Bursera (Sapindaceae). New Phytologist 193: 276–287. Dolara P, Luceri C, Ghelardini C, Monserrat C, Aiolli S, Luceri F, Lodovici M, Menichetti S, Romanelli MN. 1996. Analgesic effects of myrrh. Nature 379: 29. Fine PVA, Daly DC, Villa G, Mesones I, Cameron KM. 2005. The contribution of edaphic heterogeneity to the evolution and diversity of Burseraceae trees in the western Amazon. Evolution 59: 1464–1478. Fine PVA, Zapata F, Daly DC. 2014. Investigating processes of Neotropical rain forest tree diversification by examining the evolution and historical biogeography of the Protieae (Burseraceae). Evolution 68: 1988–2004. Forman LL, Van der Ham RWJM, Harley MM, Lawrence TJ. 1994. Rosselia, a new genus of Burseraceae from the Louisiade Archipelago, Papua New Guinea. Kew Bulletin 49: 601–621. Groom N, 1981. Frankincense and myrrh: a study of the Arabian incense trade. Longman, London. Harley MM, Song U, Banks H. 2005. Pollen morphology and systematics of Burseraceae. Grana 44: 282–289. Lambert JB, Donnelly EW, Heckenbach EA, Johnson CL, Kozminski MA, Wu Y, Santiago-Blay JA. 2013. Molecular classification of natural exudates in rosids. Phytochemistry 94: 171–183. Pernet R. 1972. Phytochimie des Burseracées. Lloydia 35: 280–287. Swart JJ. 1942. A monograph of the genus Protium and some allied genera, Burseraceae. Koch en Knuttel, Gouda. Thulin M, Beier BA, Razafimandimbison SG, Banks HI. 2008. Ambilobea, a new genus from Madagascar, the position of Aucoumea, and comments on the tribal classification of the frankincense and myrrh family. Nordic Journal of Botany 26: 218–229. Tucker AO. 1986. Frankincense and myrrh. Economic Botany 40: 425–433. Webber IE. 1941. Systematic anatomy of the woods of the “Burseraceae”. Lilloa 6: 441–465. Weeks A. 2009. Evolution of the pili nut genus Canarium L., Burseraceae, and its cultivated species. Genetic Resources and Crop Evolution 56: 765–781. Weeks A, Daly DC, Simpson BB. 2005. The phylogenetic history and biogeography of the frankincense and myrrh family (Burseraceae) based on nuclear and chloroplast sequence data. Molecular Phylogenetics and Evolution 35: 85–101. Weeks A, Simpson BB. 2007. Molecular phylogenetic

analysis of Commiphora (Burseraceae) yields insight on the evolution and historical biogeography of an “impossible” genus. Molecular Phylogenetics and Evolution 42: 62–79. 271. ANACARDIACEAE CASHEW FAMILY Aguilar-Ortigoza CJ, Sosa V. 2004. The evolution of toxic phenolic compounds in a group of Anacardiaceae genera. Taxon 53: 357–364. Bachelier JB, Endress, PK. 2009. Comparative floral morphology and anatomy of Anacardiaceae and Burseraceae (Sapindales), with special focus on gynoecium structure and evolution. Botanical Journal of the Linnean Society 159: 499–571. Barfod A. 1988. Inflorescence morphology of some South American Anacardiaceae and the possible phylogenetic trends. Nordic Journal of Botany 8: 3–11. Behrens R. 1996. Cashew as an agroforestry crop: prospects and potentials. Margraf, Weikersheim. Epstein WL. 1994. Occupational poison ivy and oak dermatitis. Dermatologic Clinics 12: 511–516. Gadek P, Fernando ES, Quinn CJ, Hoot SB, Terrazas T, Sheahan MC, Chase MW. 1996. Sapindales: molecular delimitation and infraordinal groups. American Journal of Botany 83: 802–811. Gambaro V, Chamy MC, Von Brand E, Gambarino JA. 1986. 3-(pentadec-10-enyl)-catechol, a new allergenic compound from Lithraea caustica (Anacardiaceae). Planta Medica 44: 20–22. Gillis WT. 1971. The systematics and ecology of poison-ivy and poison-oaks. Rhodora 73: 71–159, 161–237, 370–443, 465–540. Hall JB, O’Nrien EM, Sinclair FL. 2002. Sclerocarya birrea: a monograph. University of Wales, Bangor. Heimsch CH Jr. 1940. Wood anatomy and pollen morphology of Rhus and allied genera. Journal of the Arnold Arboretum 21: 279–291. Herrera F, Manchester SR, Jaramillo C. 2012. Permineralized fruits from the late Eocene of Panama give clues of the composition of forests established early in the uplift of Central America. Review of Palaeobotany and Palynology 175: 10–24. Hou D. 1978. Anacardiaceae. Flora Malesiana I, 8(3): 395–548. Kostermans AJGH, Bompard J-M. 1993. The mangoes, their botany, nomenclature, horticulture and utilization. Academic Press, London. Kullavanijaya P, Ophaswongse S. 1997. A study of dermatitis in the lacquerware industry. Contact Dermatitis 36: 244–246. Lambert JB, Donnelly EW, Heckenbach EA, Johnson CL, Kozminski MA, Wu Y, Santiago-Blay JA. 2013. Molecular classification of natural exudates in rosids. Phytochemistry 94: 171–183. Manchester SR, Wilde V, Collinson ME. 2007. Fossil cashew nuts from the Eocene of Europe: biogeographic links between Africa and South America. International Journal of Plant Sciences 168: 1199–1206. Miller AJ, Young DA, Wen J. 2001. Phylogeny and biogeography of Rhus (Anacardiaceae) based on ITS sequences. International Journal of Plant Sciences 162: 1401–1407. Mitchell JD. 1990. The poisonous Anacardiaceae genera of the world. Advances in Economic Botany 8: 103–129. Mitchell JD, Daly D, Pell SK, Randrianasolo A. 2006. Poupartiopsis gen. nov. and its context in Anacardiaceae classification. Systematic Botany 31: 337–348.

FURTHER READING Moffett RO. 2007. Name changes in Old World Rhus and recognition of Searsia (Anacardiaceae). Bothalia 37: 165–175. Pell SK, Urbatsch L. 2000. Evaluation of evolutionary relationships in Anacardiaceae using matK sequence data. American Journal of Botany 87(6, suppl.): 149. Prendergast HDV, Jaeschke HF, Rumball N. 2001. A lacquer legacy at Kew. Royal Botanic Gardens, Kew. Venning FD. 1948. The ontogeny of the laticiferous canals in the Anacardiaceae. American Journal of Botany 35: 637–644. Yi T, Miller AJ, Wen J. 2007. Phylogeny of Rhus (Anacardiaceae) based on sequences of nuclear Nia-i3 intron and chloroplast trnC-trnD. Systematic Botany 32: 379–391. 272. SAPINDACEAE MAPLE FAMILY Buerki S, Forest F, Acevedo-Rodríguez P, Callmander MW, Nylander JAA, Harrington M, Sanmartín I, Küpfer F, Alvarez N. 2009. Plastid and nuclear DNA markers reveal intricate relationships at subfamilial and tribal levels in the soapberry family (Sapindaceae). Molecular Phylogenetics and Evolution 51: 238–258. Buerki S, Lowry II PP, Alvarez N, Razafimandimbison SG, Küpfer P, Callmander MW. 2010. Phylogeny and circumscription of Sapindaceae revisited: molecular sequence data, morphology, and biogeography support recognition of a new family, Xanthoceraceae. Plant Ecology and Evolution 143: 148–159. Buerki S, Forest F, Alvarez N, Nylander JAA, Arrigo N, Sanmartín I. 2010. An evaluation of new parsimony-based versus parametric inference methods in biogeography: a case study using the globally distributed plant family Sapindaceae. Journal of Biogeography 38: 531–550. Buerki S, Lowry II PP, Andriambololonera S, Phillipson PB, Vary L, Callmander MW. 2011. How to kill two genera with one tree: clarifying generic circumscriptions in an endemic Malagasy clade of Sapindaceae. Botanical Journal of the Linnean Society 165: 223–234. Buerki S, Forest F, Salamin N, Alvarez N. 2010. Comparative performance of supertree algorithms in large data sets using the soapberry family (Sapindaceae) as a case study. Systematic Biology 60: 32–44. Buerki S, Callmander MW, Lowry II PP, Devey DS, Munzinger J. 2012. Phylogenetic inference of New Caledonian lineages of Sapindaceae: molecular evidence requires a reassessment of generic circumscriptions. Taxon 61: 109–119. Buerki S, Forest F, Alvarez N. 2013. The abrupt climate change at the Eocene-Oligocene boundary and the emergence of Southeast Asia triggered the spread of sapindaceous lineages. Annals of Botany 112: 151–160. Erwin DM, Stockey RA. 1990. Sapindaceous flowers from the Middle Eocene Princeton chert (Allenby Formation) of British Columbia, Canada. Canadian Journal of Botany 68: 2025–2034. Gadek P. Fernando ES, Quinn CJ, Hoot SB, Terrazas T, Sheahan MC, Chase MW. 1996. Sapindales: molecular delimitation and infraordinal groups. American Journal of Botany 83: 802–811. Harrington MG, Gadek PA. 2009. A species well travelled — the Dodonaea viscosa (Sapindaceae) complex based on phylogenetic analysis of nuclear ribosomal ITS and ETSf sequences. Journal of

Biogeography 36: 2313–2323. Harrington MG, Gadek PA. 2010. Phylogenetics of hipbushes and pepperflowers (Dodonaea, Diplopeltis — Sapindaceae), based on nuclear ribosomal ITS and partial ETS sequences incorporating secondary-structure models. Australian Systematic Botany 23: 431–442. Harris AJ, Xiang QY, Thomas DT. 2009. Phylogeny, origin, and biogeographic history of Aesculus L. (Sapindales) — an update from combined analysis of DNA sequences, morphology, and fossils. Taxon 58: 108–126. Manchester SR, Chen Z-D, Lu A-M, Uemura K. 2009. Eastern Asian endemic seed plant genera and their paleogeographic history throughout the Northern Hemisphere. Journal of Systematics and Evolution 47: 1–41. Pelegrini DD, Tsuzuki JK, Amado CAB, Cortez DAG, Ferreira ICP. 2008. Biological activity and isolated compounds in Sapindus saponaria L. and other plants of the genus Sapindus. Latin American Journal of Pharmacy 27: 922–927. Renner SS, Beenken L, Grimm GW, Schaefer H, Ricklefs RE. 2007. The evolution of dioecy, heterodichogamy, and labile sex expression in Acer. Evolution 61: 2701–2719. Thulin M. 2004. A new genus of Sapindaceae from Somalia. Nordic Journal of Botany 24: 509–511. Van Gelderen DM, Oterboom HJ, Jong PC. 2010. Maples of the world. Timber Press, Portland. 273. RUTACEAE CITRUS FAMILY Appelhans MS, Smets E, Razamfimandimbison SG, Haevermans R, Van Marle EJ, Couloux A, Rabarison J, Randrianarivelojosia M, Keßler PJA. 2011. Phylogeny, evolutionary trends and classification of the Spathelia-Ptaeroxylon clade: morphological and molecular insights. Annals of Botany 107: 1259–1277. Appelhans MS, Van Heuven BJ, Lens F, Baas P. 2012. Phylogenetic and ecological signals in the wood of Spathelioideae (Rutaceae). IAWA Journal 33: 337–353. Appelhans MS, Wen J, Wood KR, Allan GJ, Zimmer EA, Wagner WL. 2014. Molecular phylogenetic analysis of Hawaiian Rutaceae (Melicope, Platydesma and Zanthoxylum) and their different colonisation patterns. Botanical Journal of the Linnean Society 174: 425–448. Bayer RJ, Mabberley DJ, Morton C, Miller CH, Sharma IK, Pfeil BE, Rich S, Hitchcock R, Sykes S. 2009. A molecular phylogeny of the orange subfamily (Rutaceae-Aurantioideae). American Journal of Botany 96: 668–685. Boesewinkel FD. 1977. Development of ovule and testa in Rutaceae I. Ruta, Xanthoxylum, and Skimmia. Acta Botanica Neerlandica 26: 193–211. Brückner C. 1991. Fruchtanatomische Studien an Dictamnus albus L, Zanthoxylum simulans Hance, Ptelea trifoliata L. und Ruta graveolens L. (Rutaceae). Feddes Repertorium 102: 541–570. Caris P, Smets E, De Koster K, Ronse Decraene LP. 2006. Floral ontogeny of Cneorum tricoccum L. (Rutaceae). Plant Systematics and Evolution 257: 223–232. Chase MW, Morton CM, Kallunki JA. 1999. Phylogenetic relationships of Rutaceae: a cladistic analysis of the subfamilies using evidence from rbcL and atpB sequence variation. American Journal of Botany 86: 1191–1199. Dugo G, Di Giacomo A (eds) 2002. Citrus. Taylor, Francis, London.

Engler A. 1931. Cneoraceae, Rutaceae. Die natürlichen Pflanzenfamilien ed. 2, 19a: 184–359. Gregor HJ. 1989. Aspects of the fossil record and phylogeny of the family Rutaceae (Zanthoxyleae, Toddalioideae). Plant Systematics and Evolution 162: 251–265. Groppo M, Pirani JR, Salatino MLF, Blanco SR, Kallunki JA. 2008. Phylogeny of Rutaceae based on two noncoding regions from cpDNA. American Journal of Botany 95: 985–1005. Harbough DT, Wagner WL, Allan GJ, Zimmer EA. 2009. The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: an example from the plant genus Melicope (Rutaceae). Journal of Biogeography 36: 230–241. Hartley TG. 1997. Five new rain forest genera of Australasian Rutaceae. Adansonia III, 19: 189–212. Mabberley DJ. 1997. A classification for edible Citrus. Telopea 7: 167–172. Mabberley DJ. 2004. Citrus (Rutaceae): a review of recent advances in etymology, systematics and medical applications. Blumea 49: 481–498. Mabberley DJ. 2010. The species of Citrus (Rutaceae) with pinnate leaves. Blumea 55: 73–74. Mole B, Udovicic F, Ladiges PY, Duretto MF. 2004. Molecular phylogeny of Phebalium (Rutaceae: Boronieae) and related genera based on ITS 1+2 regions of rDNA. Plant Systematics and Evolution 249: 197–212. Morton CM, Grant M, Blackmore S. 2003. Phylogenetic relationships of the Aurantioideae inferred from chloroplast DNA sequence data. American Journal of Botany 90: 1463–1469. Pfeil BE, Crisp MD. 2008. The age and biogeography of Citrus and the orange subfamily (Rutaceae: Aurantioideae) in Australasia and New Caledonia. American Journal of Botany 95: 1621–1631. Poon WS, Shaw PC, Simmons MP, But PPH. 2007. Congruence of molecular, morphological, and biochemical profiles in Rutaceae: a cladistic analysis of the subfamilies Rutoideae and Toddalioideae. Systematic Botany 32: 837–846. Reuther W, Webber HJ, Batchelor LD (eds). 1967–89. The citrus industry, 2nd ed., 5 vols. University of California, Riverside. Salvo G, Bachetta G, Ghahremaninejad F, Conti E. 2008. Phylogenetic relationships of Ruteae (Rutaceae): new evidence from the chloroplast genome and comparisons with non-molecular data. Molecular Phylgenetics and Evolutions 49: 736–748. Salvo G, Ho SYW, Rosenbaum G, Ree R, Conti E. 2010. Tracing the temporal and spatial origins of island endemics in the Mediterranean region: a case study from the citrus family (Ruta L., Rutaceae). Systematic Biology 59: 705–722. Samuel R, Ehrendorfer F, Chase MW, Greger H. 2001. Phylogenetic analysis of Aurantioideae (Rutaceae) based on non-coding plastid DNA sequences and phytochemical features. Plant Biology 3: 77–87. Scott KD, McIntyre CL, Playford J. 2000. Molecular analyses suggest a need for a significant rearrangement of Rutaceae subfamilies and a minor reassessment of species relationships within Flindersia. Plant Systematics and Evolution 223: 15–27. Stace HM, Armstrong JA, James SH. 1993. Cytoevolutionary patterns in Rutaceae. Plant Systematics and Evolution 187: 1–28. Tolkowsky S. 1938. Hesperides: a history of the culture and use of citrus fruits. John Bales, Sons, Curnow, London.

Plants of the World

719

FURTHER READING Trinder-Smith TH, Linder HP, Van der Niet T, Verboom GA, Nowell TL. 2007. Plastid DNA sequences reveal generic paraphyly within Diosmeae (Rutoideae, Rutaceae). Systematic Botany 32: 847–855. Waterman PG, Grundon MF (eds) 1983. Chemistry and chemical taxonomy of the Rutales. Academic Press, London. Wei L, Wang YZ, Li ZY. 2012. Floral ontogeny of Ruteae (Rutaceae) and its systematic implications. Plant Biology 14: 190–197. Wilson PG. 1998. A taxonomic review of the genera Eriostemon and Philotheca (Rutaceae: Boronieae). Nuytsia 12: 239–265. Zhou Q, Wang Y, Jin X. 2002. Ontogeny of floral organs and morphology of f loral apex in Phellodendron amurense (Rutaceae). Australian Journal of Botany 50: 633–644. 274. SIMAROUBACEAE TREE-OF-HEAVEN FAMILY Abbe EC, Earle TT. 1940. Inf lorescence, f loral anatomy and morphology of Leitneria floridana. Bulletin of the Torrey Botanical Club 67: 173–193. Channell RB, Wood CE Jr. 1962. The Leitneriaceae in the southeastern United States. Journal of the Arnold Arboretum 43: 435–438. Clayton JW, Fernando ES, Soltis PM, Soltis DE. 2007. Molecular phylogeny of the tree-of-heaven family (Simaroubaceae) based on chloroplast and nuclear markers. International Journal of Plant Sciences 168: 1325–1339. Clayton JW, Soltis PS, Soltis DE. 2009. Recent long-distance dispersal overshadows ancient biogeographical patterns in a pantropical angiosperm family (Simaroubaceae, Sapindales). Systematic Biology 58: 395–410. Corbett SL, Manchester SR. 2004. Phytogeography and fossil history of Ailanthus (Simaroubaceae). International Journal of Plant Sciences 165: 671–690. Fernando ES, Quinn CJ. 1992. Pericarp anatomy and systematics of the Simaroubaceae s.l. Australian Journal of Botany 40: 263–289. Fernando ES, Gadek PA, Quinn CJ. 1995. Simaroubaceae, an artificial construct: evidence from rbcL sequence variation. American Journal of Botany 82: 92–103. Fernando ES, Quinn CJ. 1995. Picramniaceae, a new family, and a recircumscription of Simaroubaceae. Taxon 44: 177–181. Petersen FP, Fairbrothers DE. 1983. A serotaxonomic appraisal of Amphipterygium and Leitneria– two amentiferous taxa of Rutiflorae (Rosidae). Systematic Botany 8: 134–148. Webber IE. 1936. Systematic anatomy of the woods of Simaroubaceae. American Journal of Botany 23: 577–587. 275. MELIACEAE NEEM FAMILY Cheek MR. 1992. The wood anatomy of Pseudobersama mossambicensis and Trichilia capitata (Meliaceae) compared. Kew Bulletin 47: 753–758. Fisher JB, Rutishauser R. 1990. Leaves and epiphyllous shoots in Chisocheton (Meliaceae): a continuum of woody leaf and stem axes. Canadian Journal of Botany 68: 2316–2328. Grudinksi M, Wanntorp L, Pannell CM, MuellnerRiehl AN. 2014. West to east dispersal in a widespread animal-dispersed woody angiosperm genus (Aglaia, Meliaceae) across the

720

Christenhusz, Fay & Chase

Indo-Australian Archipelago. Journal of Biogeography 41: 1149–1159. Kribs DA. 1930. Comparative anatomy of the woods of the Meliaceae. American Journal of Botany 17: 724–738. Mabberley DJ. 1979. The species of Chisocheton (Meliaceae). Bulletin of the British Museum of Natural History, Botany 6: 301–386. Muellner AN, Mabberley DJ. 2008. Phylogenetic position and taxonomic disposition of Turraea brevif lora (Meliaceae), a hitherto enigmatic species. Blumea 53: 607–616. Muellner AN, Samuel R, Johnson SA, Cheek M, Pennington TD, Chase MW. 2003. Molecular phylogenetics of Meliaceae (Sapindales) based on nuclear and plastid DNA sequences. American Journal of Botany 90: 471–480. Muellner AN, Savolainen V, Samuel R, Chase MW. 2006. The mahogany family “out-of-Africa”: divergence time estimation, global biogeographic patterns inferred from plastid rbcL DNA sequences, extant, and fossil distribution of diversity. Molecular Phylogenetics and Evolution 40: 236–250. Muellner AN, Samuel R, Chase MW, Coleman A, Stuessy TF. 2008. An evaluation of tribes and generic relationships in Melioideae (Meliaceae) based on nuclear ITS ribosomal DNA. Taxon 57: 98–108. Pannell CM. 1992. A taxonomic monograph of the genus Aglaia Lour. (Meliaceae). Kew Bulletin additional series 16. Pannell CM, Koziol MJ. 1989. Ecological and phytochemical diversity of arillate seeds in Aglaia (Meliaceae): a study of vertebrate dispersal in tropical trees. Philosophical Transactions of the Royal Society B 316: 303–333. Pennington TD. 1981. Meliaceae. Flora Neotropica Monograph 28. Pennington TD, Styles BT. 1975. A generic monograph of the Meliaceae. Blumea 22: 419–540. Singh KK, Phogat S, Dhillon RS, Tomar A (eds). 2009. Neem. I.K. International Publishing House, New Delhi. 276. PETENAEACEAE PETÉN-LINDEN FAMILY Christenhusz MJM, Fay MF, Clarkson JJ, Gasson P, Morales Can J, Jiménez Barrios JB, Chase MW. 2010. Petenaeaceae, a new angiosperm family in Huerteales with a distant relationship to Gerrardina (Gerrardinaceae). Botanical Journal of the Linnean Society 164: 16–25. Kukachka BF. 1962. Wood anatomy of Petenaea cordata Lu ndell ( E l a e o c a r p a c e a e) . Wrightia 3: 36–40. Lundell CL. 1962. Plantae Mayanae — V. Petenaea cordata, a new genus and species in the Elaeocarpaceae, and other taxonomic notes. Wrightia 3: 21–35. 277. GERRARDINACEAE BROWN-IRONWOOD FAMILY Alford, M.H. 2006. Gerrardinaceae: a new family of African flowering plants unresolved among Brassicales, Huerteales, Malvales, and Sapindales. Taxon 55: 959–964. Christenhusz MJM, Fay MF, Clarkson JJ, Gasson P, Morales Can J, Jiménez Barrios JB, Chase MW. 2010. Petenaeaceae, a new angiosperm family in Huerteales with a distant relationship to Gerrardina (Gerrardinaceae). Botanical Journal of the Linnean Society 164: 16–25.

Worberg A, Alford MH, Quandt D, Borsch T. 2009. Huerteales sister to Brassicales plus Malvales, and newly circumscribed to include Dipentodon, Gerrardina, Huertea, Perrottetia, and Tapiscia. Taxon 58: 468–478 278. TAPISCIACEAE SILVERPHEASANT-TREE FAMILY Carlquist S, Hoekman DA. 1985. Wood anatomy of Staphyleaceae: ecology, statistical correlations, and systematics. Flora 177: 195–216. Cuatrecasas J. 1953. Huertea, un genre nouveau pour la flore de Colombie. Bulletin de la Société Botanique de France 100: 159–163. Dickison WC. 1987. Leaf and nodal anatomy and systematics of Staphyleaceae. Botanical Gazette 148: 475–489. Liu WZ, Kang HQ, Zheng HC, Feng YZ. 2008. An investigation on the sexual reproductive cycle in Tapiscia sinensis. Journal of Systematics and Evolution 46: 175–182. Liu W, Ni X. 2013. Anatomy and development of gynoecium in Tapiscia sinensis. Frontiers in Plant Evolution and Development 4: 421. Manchester SR. 1988. Fruits and seeds of Tapiscia (Staphyleaceae) from the Middle Eocene of Oregon, U.S.A. Tertiary Research 9: 59–66. Solereder H. 1892. Über die Staphylaceengattung Tapiscia Oliv. Berichte der Deutsche Botanische Gesellschaft 10: 545–551. Xie CP. 2006. A review of research advances in rare and endangered plant Tapiscia sinensis. Subtropical Plant Science 4: 20. 279. DIPENTODONTACEAE SHICHI FAMILY Den Hartog RM, Baas P. 1978. Epidermal characters of the Celastraceae sensu lato. Acta Botanica Neerlandica 27: 355–388. Dunn ST. 1911. Dipentodon. A new genus of uncertain systematic position. Bulletin of Miscellaneous Information (Kew) 7: 310–313. Liu JS, Cheng JR. 1991. On the systematic position of genus Dipentodon Dunn. Journal of Wuhan Botanical Research 9: 29–39. Lundell CJ. Neotropical species of the genus Perrottetia (Celastraceae). Phytologia 57: 231–238. Matthews ML, Endress PK. 2005. Comparative floral structure and systematics in Celastrales (Celastraceae, Parnassiaceae, Lepidobotryaceae). Botanical Journal of the Linnean Society 149: 129–194. Peng Y, Chen Z, Gong X, Zhong Y, Shi S. 2003. Phylogenetic position of Dipentodon sinicus: evidence from DNA sequences of chloroplast rbcL, nuclear ribosomal 18S, and mitochondrial matR genes. Botanical Bulletin of Academia Sinica (Taipei) 44. Yuan QJ, Zhang ZY, Peng H, Ge S. 2008. Chloroplast phylogeny of Dipentodon (Dipentodontaceae) in southwest China and northern Vietnam. Molecular Ecology 17: 1054–1065. Zhang LB, Simmons MP. 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes. Systematic Botany 31: 122–137. 280. CYTINACEAE ROCKROSE-RAPE FAMILY Alvarado-Cárdenas LO. 2009. Systematics of the genus Bdallophytum (Cytinaceae). Acta Botanica Mexicana 87: 1–21. Bouman F, Meijer W. 1994. Comparative structure of ovules and seeds in Rafflesiaceae. Plant Systematics and Evolution 193: 187–212.

FURTHER READING Burgoyne PM. 2006. A new species of Cytinus (Cytinaceae) from South Africa and Swaziland with a key to the southern African species. Novon 16: 315–319. De Vega C, Berjano R, Arista M, Ortiz PL, Talavera S, Stuessy TF. 2008. Genetic races associated with the genera and sections of host species in the holoparasitic plant Cytinus (Cytinaceae) in the western Mediterranean basin. New Phytologist 178: 875–887. De Vega C, Ortiz PL, Arista M, Talavera S. 2007. The endophytic system of Mediterranean Cytinus (Cytinaceae) developing on five host Cistaceae species. Annals of Botany 100: 1209–1217. Garcia-Franco JG, Rico-Gray V. 1997. Reproductive biology of the holoparasitic endophyte Bdallophyton bambusarum (Rafflesiaceae). Botanical Journal of the Linnean Society 123: 237–247. Govaerts R, Nickrent DL. Cytinus ruber (Cytinaceae). Curtis’s Botanical Magazine 26: 314–321. Nickrent DL. 2007. Cytinaceae are sister to Muntingiaceae (Malvales). Taxon 56: 1129–1135. Visser J. 1981. South African parasitic flowering plants. Juta, Capetown. 281. MUNTINGIACEAE BAJELLY-TREE FAMILY Bayer C, Chase MW, Fay MF. 1998. Muntingiaceae, a new family of dicotyledons with malvalean affinities. Taxon 47: 37–42. Benn SJ, Lembke DE. 1991. Taxonomy of Neotessmannieae (Tiliaceae). American Journal of Botany Supplement 78: 166–167. Carlquist S. 2005. Wood and bark anatomy of Muntingiaceae: a phylogenetic comparison within Malvales s.l. Brittonia 57: 59–67. Fleming TH, Williams CF, Bonaccorso FJ, Herbst LH. 1985. Phenology, seed dispersal and colonisation in Muntingia calabura, a Neotropical pioneer tree. American Journal of Botany 72: 383–391. Venkata Rao C. 1951. Life history of Muntingia calabura L. Current Science 20: 47–48. 282. NEURADACEAE PIETSNOT FAMILY Alverson WS, Karol KG, Baum DA, Chase MW, Swensen SM; McCourt R, Sytsma KJ. 1998. Circumscription of the Malvales and relationships to other Rosidae: evidence from rbcL sequence data. American Journal of Botany 85: 876–887. Huber H. 1993. Neurada, eine Gattung der Malvales. Sendtnera 1: 7–10. Ronse Decraene LP, Smets EF. 1996. The floral development of Neurada procumbens L. (Neuradaceae). Acta Botanica Neerlandica 45: 228–241. 283. MALVACEAE MALLOW FAMILY Alverson WS, Karol KG, Baum DA, Chase MW, Swensen SM, McCourt R, Sytsma KJ. 1998. Circumscription of the Malvales and relationships to other Rosidae: evidence from rbcL sequence data. American Journal of Botany 85: 876–887. Alverson WS, Whitlock BA, Nyffeler R, Bayer C, Baum DA. 1999. Phylogeny of core Malvales: evidence from ndhF sequence data. American Journal of Botany 86: 1474–1486. Anderson GJ. 1976. The pollination biology of Tilia. American Journal of Botany 63: 1203–1212. Bates DM. 1968. Generic relationships in the Malvaceae, tribe Malveae. Gentes Herbarum 10: 117–135. Baum DA, Alverson WS, Nyffeler RN. 1998. A durian by any other name: taxonomy and nomenclature of the core Malvales. Harvard Papers in Botany 3: 317–332.

Baum DA. 1995. The comparative pollination and f loral biology of baobabs (AdansoniaBombacaceae). Annals of the Missouri Botanical Garden 82: 322–348. Bayer C. 1999. The bicolor unit — homology and transformation of an inflorescence structure unique to core Malvales. Plant Systematics and Evolution 214: 187–198. Bayer C, Fay MF, De Bruijn AY, Savolainen V, Morton CM, Kubitzki K, Alverson WS, Chase MW. 1999. Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atpB and rbcL DNA sequences. Botanical Journal of the Linnean Society 129: 267–303. Brodie S, Cheek M, Staniforth M. 1998. Trochetiopsis ebenus. Sterculiaceae. Curtis’s Botanical Magazine 15: 27–36. Carvalho MR, Herrera FA, Jaramillo CA, Wing SL, Callejas R. 2011. Paleocene Malvaceae from northern South America and their biogeographical implications. American Journal of Botany 98: 1337–1355. Christensen PB. 1986. Pollen morphological studies in the Malvaceae. Grana 25: 95–117. Cronk QCB. 1990. The history of the endemic flora of St Helena: late Miocene ‘Trochetiopsis-like’ pollen from St Helena and the origin of Trochetiopsis. New Phytologist 114: 159–165. Cuatrecasas J. 1964. Cacao and its allies — a taxonomic revision of the genus Theobroma. Contributions from the United States National Herbarium 35: 379–614. Dillehay TD, Rossen J, Andres TC, Williams DE. 2007. Preceramic adoption of peanut, squash, and cotton in northern Peru. Science 316: 1890–1893. Donnell AA, Ballard HE Jr, Cantino PD. 2012. Callianthe (Malvaceae): a new genus of Neotropical Malvaceae. Systematic Botany 37: 212–222. Feeney J. 2001. The red tea of Egypt. Saudi Aramco World 52(5): 36–41. Fryxell PA. 1979. The natural history of the cotton tribe. Texas A&M University Press, College Station. Fuchs H. 1967. Pollen morphology of the family Bombacaceae. Review of Palaeobotany and Palynology 3: 119–132. García PE, Schönswetter P, Aguilar JF, Feliner GN, Schneeweiss GM. 2009. Five molecular markers reveal extensive morphological homoplasy and reticulate evolution in the Malva alliance (Malvaceae). Molecular Phylogenetics and Evolution 50: 226–239. Hinsley SR. 2002. The Malvaceae pages. http://www. malvaceae/info Janka H, Von Balthazar M, Alverson WS, Baum DA, Semir J, Bayer C. 2008. Structure, development and evolution of the androecium in Adansonieae (core Bombacoideae, Malvaceae s.l.). Plant Systematics and Evolution 275: 69–91. Koopman MM, Baum DA. 2008. Phylogeny and biogeography of trube Hibisceae (Malvaceae) on Madagascar. Systematic Botany 33: 364–374. Le Péchon T, Dubuisson J-Y, Haevermans T, Cruaud C, Couloux A, Gigord LDB. 2010. Multiple colonizations from Madagascar and converged acquisition of dioecy in the Mascarene Dombeyoideae (Malvaceae) as inferred from chloroplast and nuclear DNA sequence analysis. Annals of Botany 106: 343–357. Manchester SR. 1994. Inflorescence bracts of fossil and extant Tilia in North America, Europe, and Asia: patterns of morphological divergence and

biogeographic history. American Journal of Botany 81: 1176–1185. Nyffeler R, Bayer C, Alverson WS, Yen A, Whitlock BA, Chase MW, Baum DA. 2005. Phylogenetic analysis of the Malvadendrina clade (Malvacee s.l.) based on plastid DNA sequences. Organisms, Diversity and Evolution 5: 109–123. Paterson AH et al. (74 authors) 2012. Repeated polyploidisation of Gossypium genomes and the evolution of spinnable cotton fibres. Nature 492: 423–427. Pfeil BE, Crisp MD. 2005. What to do with Hibiscus? A proposed nomenclatural resolution for a large and well-known genus of Malvaceae and comments on paraphyly. Australian Systematic Botany 18: 49–60. Skema C. 2012. Towards a new circumscription of Dombeya (Malvales: Dombeyaceae): a molecular phylogenetic and morphological study of Dombeya of Madagascar and a new segregate genus Andringitra. Taxon 61: 612–628. Tate JA, Aguilar JF, Wagstaff SJ, La Duke JC, Slotta TAB, Simpson BB. 2005. Phylogenetic relationships within the tribe Malveae (Malvaceae, subfamily Malvoideae) as inferred from ITS sequence data. American Journal of Botany 92: 584–602. Von Balthazar M, Schönenberger J, Alverson WS, Janka H, Bayer C, Baum DA. 2006. Structure and evolution of the androecium in the Malvatheca clade (Malvaceae s.l.) and implications for Malvaceae and Malvales. Plant Systematics and Evolution 260: 171–197. Whitlock BA, Hale AM. 2011. The phylogeny of Ayenia, Byttneria, and Rayleya (Malvaceae s.l.) and its implications for the evolution of growth forms. Systematic Botany 36: 129–136. Wilkie P, Clark A, Pennington RT, Cheek M, Bayer C, Wilcock CC. 2006. Phylogenetic relationships within the subfamily Sterculioideae (Malvaceae/ Sterculiaceae-Sterculieae) using the chloroplast gene ndhF. Systematic Botany 31: 160–170. 284. SPHAEROSEPALACEAE LOMBIRY FAMILY Birkinshaw E, Edmond R, Rajeriarison C, Ratovoson F, Reza L, Schatz G. 2004. Red lists for Malagasy plants. IV. Sphaerosepalaceae. Missouri Botanical Garden, Antananarivo University, ANGAP. Boureau E. 1958. Contribution a l’etude anatomique des especes actuelles de Rhopalocarpaceae. Bulletin du Muséum Nationale d’Histoire Naturelle de Paris II, 30: 213–221. Den Outer RW, Schütz PR. 1981. Wood anatomy of some Sarcolaenaceae and Rhopalocarpaceae and their systematic position. Mededelingen van de Landbouwhogeschool Wageningen 81: 1–25. Fay MF, Bayer C, Alverson W, De Bruijn AY, Swensen SM, Chase MW. 1998. Plastid rbcL sequences indicate a close affinity between Diegodendron and Bixa. Taxon 47: 43–50. Horn JW, Dickison WS. 1997. Structural biology and phylogenetics of the Sphaerosepalaceae and Diegodendraceae. American Journal of Botany 84: 44. Huard J. 1965. Anatomie des Rhopalocarpacées. Adansonia II, 5: 103–123. Schatz GE, Lowry II P, Wolf AE. 1999. Endemic families of Madagascar II. A synoptic revision of Sphaerosepalaceae. Adansonia III, 21: 107–123. Van Tieghem P. 1900. Sur les Bixacées, les Cochlospermacées et les Sphérosépalacées. Journal de Botanique (L. Morot) 14: 32–54.

Plants of the World

721

FURTHER READING 285. THYMELAEACEAE MEZEREON FAMILY Beaumont A, Edwards TJ, Manning J, Maurin O, Rautenbach M, Motsi MC, Fay MF, Chase MW, Van der Bank M. 2009. Gnidia (Thymelaeaceae) is not monophyletic: taxonomic implications for Thymelaeoideae and a partial new generic taxonomy for Gnidia. Botanical Journal of the Linnean Society 160: 402–417. Eyken PAAF. 1906. Sur l’essence du bois de Gonystylus miquelianus, T., B. Receuil des Travaux Chimiques des Pays-Bas (et de la Belgique): 44–47. Halda JJ. 2001. The genus Daphne. Sen, Dobré. Heinig K. 1951. Studies in the floral morphology of the Thymelaeaceae. American Journal of Botany 38: 113–132. Herber BE. 2002. Pollen morphology of the Thymelaeaceae in relation to its taxonomy. Plant Systematics and Evolution 232: 107–121. Maguire B, Steyermark JA. 1981. Tepuianthaceae, Sapindales. Memoirs of the New York Botanical Garden 32: 4–21. Motsi MC, Moteetee AM, Beaumont AJ, Rye BL, Powell MP, Savolainen V, Van der Bank M. 2010. A phylogenetic study of Pimelea and Thecanthes (Thymelaeaceae): evidence from plastid and nuclear ribosomal DNA sequence data. Australian Systematic Botany 23: 270–284. Nevling LI. 1959. A revision of the genus Daphnopsis. Annals of the Missouri Botanical Garden 46: 257–353. Rogers ZS. 2009. A world checklist of Thymelaeaceae (version 1). Missouri Botanical Garden, St Louis. http://www.tropicos.org/project/Thymelaeaceae Roth I, Lindorf H. 1990. Blatt- und Rindenstruktur von Tepuianthus auyantepuiensis, Tepuianthaceae, einer neueren Familie aus Venezuela. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 111: 403–421. Tan K. 1980. Studies in the Thymelaeaceae I, II. Notes from the Royal Botanic Garden Edinburgh 38: 149–164, 189–246. Van der Bank M, Fay MF, Chase MW. 2002. Molecular phylogenetics of Thymelaeaceae with particular reference to African and Australian genera. Taxon 51: 329–339. Wurdack KJ, Horn JW. 2001. A reevaluation of the affinities of the Tepuianthaceae: molecular and morphological evidence for placement in the Malvales. Botany 2001: plants and people, abstracts: 151. 286. BIXACEAE ANNATTO FAMILY Capuron R. 1963. Diegodendron R. Capuron gen. nov., type de la nouvelle famille Diegodendraceae (Ochnales sensu Hutchinson). Adansonia II, 3: 385–392. Dempsey RE, Garwood NC. 1994. A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character. Bulletin of the Natural History Museum London, Botany 24: 173–179. Dickison WC. 1988. Xylem anatomy of Diegodendron humbertii. IAWA Bulletin II, 9: 332–336. Fay MF, Bayer C, Alverson W, De Bruijn AY, Swensen SM, Chase MW. 1998. Plastid rbcL sequences indicate a close affinity between Diegodendron and Bixa. Taxon 47: 43–50. Horn JW, Dickison WS. 1997. Structural biology and phylogenetics of the Sphaerosepalaceae and Diegodendraceae. American Journal of Botany 84: 44. Ingram JS, Francis BJ. 1969. The annatto tree (Bixa

722

Christenhusz, Fay & Chase

orellana L.) — a guide to its occurrence, cultivation, preparation and uses. Tropical Science 9: 97–102. Keating RC. 1968. Comparative morphology of Cochlospermaceae. I. Synopsis of the family and wood anatomy. Phytomorphology 18: 379–392. Poppendieck H-H. 1980. A monograph of the Cochlospermaceae. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 101: 191–265. Ronse Decraene LP. 1989. Floral development of Cochlospermum tinctorium and Bixa orellana with special emphasis on the androecium. American Journal of Botany 76: 1344–1359. 287. CISTACEAE ROCK-ROSE FAMILY Catalina Londoño A, Alvarez E, Forero E, Morton CM. 1995. A new genus and species of Dipterocarpaceae from the Neotropics. I. Introduction, taxonomy, ecology, and distribution. Brittonia 47: 237–247. Civeyrel L, Leclercq J, Demoly JP, Agnan Y, Quèbre N, Pélissier C, Otto T. 2011. Molecular systematics, character evolution, and pollen morphology of Cistus and Halimium. Plant Systematics and Evolution 295: 23–54. Comadini O, Contu M, Rinaldi AC. 2006. An overview of Cistus ectomycorrhizal fungi. Mycorrhiza 16: 381–395. Dickie IA, Guza RC, Krazewski SE, Reich PB. 2004. Shared ectomycorrhizal fungi between a herbaceous perennial (Helianthemum bicknellii) and oak (Quercus) seedlings. New Phytologist 164: 375–382. Guzmán B, Lledó MD, Vargas P. 2009. Adaptive radiation in Mediterranean Cistus (Cistaceae). PLoS ONE 4: e6362. Guzmán B, Vargas P. 2009. Historical biogeography and character evolution of Cistaceae (Malvales) based on analysis of plastid rbcL and trnL-trnF sequences. Organisms, Diversity and Evolution 9: 83–99. Maguire B, Ashton PS, De Zeeuw C, Giannasi DE, Niklas KJ. 1977. Pakaraimoideae, Dipterocarpaceae of the Western Hemisphere. Taxon 26: 341–385. Nandi 1998. Floral development and systematics of Cistaceae. Plant Systematics and Evolution 212: 107–134. Papaefthimiou D, Papanikolaou A, Falara V, Givanoudi S, Kostas S, Kanellis AK. 2014. Genus Cistus: a model for exploring labdane-type diterpenes’ biosynthesis and a natural source of high value products with biological, aromatic, and pharmacological properties. Frontiers in Chemistry doi: 10.3389/fchem.2014.00035 Ukraintseva VV. 1993. Pollen morphology of the family Cistaceae in relation to its taxonomy. Grana (Suppl.) 2: 33–36. 288. SARCOLAENACEAE TUNIC-BELLS FAMILY Aubriot X, Soulebeau A, Haevermans T, Schatz G, Cruaud C, Lowry PP. 2016. Molecular phylogenetics of Sarcolaenaceae (Malvales), Madagascar’s largest endemic plant family. Botanical Journal of the Linnean Society 182: 729–743. Capuron R. 1970. Observations sur les Sarcolaenacées. Adansonia II, 10: 247–265. Carlquist S. 1964. Pollen morphology and evolution of Sarcolaenaceae (Chlaenaceae). Brittonia 16: 231–254. Coetzee JA, Muller J. 1984. The phytogeographic

significance of some extinct Gondwana pollen types from the Tertiary of the southwestern Cape (South Africa). Annals of the Missouri Botanical Garden 71: 1088–1099. Den Outer RW, Schütz PR. 1981. Wood anatomy of some Sarcolaneaceae and Rhopalocarpaceae and their systematic position. Mededelingen van de Landbouwhogeschool Wageningen 81: 1–25. Ducousso M, Béna D, Bourgeois C, Buyck B, Eyssartier G, Vincelette M, Rabevohitra W, Randrihasipara, Dreyfus B, Prin Y. The last common ancestor of Sarcolaenaceae and Asian dipterocarp trees was ectomycorrhizal before the India-Madagascar separation, about 88 million years ago. Molecular Ecology 13: 231–236. Lowry II PP, Schatz GE, Leroy JF, Wolf AE. 1999. Endemic families of Madagascar. III. A synoptic revision of Schizolaena (Sarcolaenaceae). Adansonia III, 21: 183–212. Lowry II PP, Haevermans T, Labat JN, Schatz GE, Leroy JF, Wolf AE. 2000. Endemic families of Madagascar. V. A synoptic revision of Eremolaena, Pentachlaena, and Perrierodendron (Sarcolaenaceae). Adansonia III, 22: 11–31. Nilsson S, Coetzee J, Grafström E. 1996. On the origin of the Sarcolaenaceae with reference to pollen morphological evidence. Grana 35: 321–334. Randrianasolo A, Miller JS. 1999. Taxonomic revision of the genus Sarcolaena (Sarcolaenaceae). Annals of the Missouri Botanical Garden 86: 702–722. Straka H. 1971. Über das System der madagassischen Sarcolaenaceae. Berichte der Deutsche Botanische Gesellschaft 84: 731–735. 289. DIPTEROCARPACEAE MARANTI FAMILY Ashton PS. 1982. Dipterocarpaceae. Flora Malesiana I, 9: 237–552. Ashton PS, Givnish TJ, Appanah S. 1988. Staggered flowering in Dipterocarpaceae: new insights into floral induction and the evolution of mast fruiting in the aseasonal tropics. American Naturalist 132: 44–66. Bate-Smith EC, Whitmore TC. 1959. Chemistry and taxonomy of the Dipterocarpaceae. Nature 184: 795–796. Beinforde C, Schäfer N, Dörfelt H, Nascimbene PC, Singh H, Heinrichs J, Reitner J, Rana RS, Schmidt AR. 2011. Ectomycorrhizas from a Lower Eocene angiosperm forest. New Phytologist 192: 988–996. Dayanandan S, Ashton PS, Williams SM, Primack RB. 1999. Phylogeny of the tropical tree family Dipterocarpaceae based on the nucleotide sequences of the chloroplast rbcL gene. American Journal of Botany 86: 1182–1190. Ducousso M, Béna D, Bourgeois C, Buyck B, Eyssartier G, Vincelette M, Rabevohitra W, Randrihasipara, Dreyfus B, Prin Y. 2003. The last common ancestor of Sarcolaenaceae and Asian dipterocarp trees was ectomycorrhizal before the India-Madagascar separation, about 88 million years ago. Molecular Ecology 13: 231–236. Dutta S, Tripathi SM, Mallick M, Mathews RP, Greenwood PF, Rao MR, Summons RE. 2011. Eocene out-of-India dispersal of Asian dipterocarps. Review of Palaeobotany and Palynology 166: 63–68. Gottwald H, Parameswaran N. 1966. Das secundäre Xylem der Familie Dipterocarpaceae. Anatomische Untersuchungen zur Taxonomie und Phylogenie. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 85: 410–508.

FURTHER READING Kamiya K, Harada K, Ogino K, Kayita T, Yamazaki T, Lee HS, Ashton PS. 1998. Molecular phylogeny of dipterocarp species using nucleotide sequences of two non-coding regions in chloroplast DNA. Tropics 7: 195–207. Kamiya K, Harada K, Tachida H, Ashton PS. 2005. Phylogeny of PgiC gene in Shorea and its closely related genera (Dipterocarpaceae), the dominant trees in Southeast Asian rain forests. American Journal of Botany 92: 775–788. Morton CM, Dayanandan S, Dissanayake D. 1999. Phylogeny and biosystematics of Pseudomonotes (Dipterocarpaceae) based on molecular and morphological data. Plant Systematics and Evolution 216: 197–205. Smits W. 1983. Dipterocarps and mycorrhiza. An ecological adaptation and a factor in forest regeneration. Flora Malesiana Bulletin 36: 3926–3927. Tsumura Y, Kado T, Yoshida K, Abe H, Ohtani M, Taguchi Y, Fukue Y, Tani N, Ueno S, Yoshimura K, Kamiya K, Harada K, Takeuchi Y, Diway B, Finkelday R, Na’iem M, Indioko S, Ng KK, Muhammad N, Lee SL. 2011. Molecular database for classifying Shorea species (Dipterocarpaceae) and techniques for checking the legitimacy of timber and wood products. Journal of Plant Research 124: 35–48. Yulita KS, Bayer RJ, West JG. 2005. Molecular phylogenetic study of Hopea and Shorea (Dipterocarpaceae): evidence from the trnL-trnF and internal transcribed spacer regions. Plant Species Biology 20: 167–182. 290. AKANIACEAE TURNIPWOOD FAMILY Boufford DE, Kjaer A, Øgaard Madsen J, Skrydstrub T. 1989. Glucosinolates in Bretscheideraceae. Biochemical Systematics and Ecology 17: 375–379. Carlquist S. 1996. Wood anatomy of Akaniaceae and Bretschneideraceae: a case of near-identity and its systematic implications. Systematic Botany 21: 607–616. Doweld AB. 1996. The systematic relevance of fruit and seed anatomy and morphology of Akania (Akaniaceae). Botanical Journal of the Linnean Society 120: 379–389. Doweld AB. 1996. The carpology and taxonomic relationships of Bretschneidera (Bretschneideraceae). Acta Botanica Malacitania 21: 79–90. Gadek PA, Quinn CJ, Rodman JE, Karol KG, Conti E, Price RA, Fernando ES. 1992. Affinities of the Australian endemic Akaniaceae: new evidence from rbcL sequences. Australian Systematic Botany 5: 717–724. Romero EJ, Hickey LJ. 1976. A fossil leaf of Akaniaceae from Paleocene beds in Argentina. Bulletin of the Torrey Botanical Club 103: 126–131. Ronse Decraene LP, Haston E. 2006. The systematic relationships of glucosinolate-producing plants and related families: a cladistic investigation based on morphological and molecular characteristics. Botanical Journal of the Linnean Society 151: 453–494. Tobe H, Peng CI. 1990. The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae). Botanical Journal of the Linnean Society 103: 139–152. Tobe H, Raven PH. 1995. Embryology and relationships of Akania (Akaniaceae). Botanical Journal of the Linnean Society 118: 261–274. 290. AKANIACEAE TURNIPWOOD FAMILY Boufford DE, Kjaer A, Øgaard Madsen J, Skrydstrub T. 1989. Glucosinolates in Bretscheideraceae.

Biochemical Systematics and Ecology 17: 375–379. Brea M, Zucol AF, Bargo MS, Fernicola JC, Vizcaíno SF. 2017. First Miocene record of Akaniaceae in Patagonia (Argentina): a fossil wood from the early Miocene Santa Cruz Formation and its palaeobiogeographical implications. Botanical Journal of the Linnean Society 183: 334–347. Carlquist S. 1996. Wood anatomy of Akaniaceae and Bretschneideraceae: a case of near-identity and its systematics implications. Systematic Botany 21: 607–616. Doweld AB. 1996. The systematic relevance of fruit and seed anatomy and morphology of Akania (Akaniaceae). Botanical Journal of the Linnean Society 120: 379–389. Doweld AB. 1996. The carpology and taxonomic relationships of Bretschneidera (Bretschneideraceae). Acta Botanica Malacitania 21: 79–90. Gadek PA, Quinn CJ, Rodman JE, Karol KG, Conti E, Price RA, Fernando ES. 1992. Affinities of the Australian endemic Akaniaceae: new evidence from rbcL sequences. Australian Systematic Botany 5: 717–724. Romero EJ, Hickey LJ. 1976. A fossil leaf of Akaniaceae from Paleocene beds in Argentina. Bulletin of the Torrey Botanical Club 103: 126–131. Ronse Decraene LP, Haston E. 2006. The systematic relationships of glucosinolate-producing plants and related families: a cladistic investigation based on morphological and molecular characteristics. Botanical Journal of the Linnean Society 151: 453–494. Tobe H, Peng CI. 1990. The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae). Botanical Journal of the Linnean Society 103: 139–152. Tobe H, Raven PH. 1995. Embryology and relationships of Akania (Akaniaceae). Botanical Journal of the Linnean Society 118: 261–274. 291. TROPAEOLACEAE NASTURTIUM FAMILY Andersson L, Andersson S. 2000. A molecular phylogeny of Tropaeolaceae and its systematic implications. Taxon 49: 721–736. Carlquist S, Donald CJ. 1996. Wood anatomy of Limnanthaceae and Tropaeolaceae in relation to habit and phylogeny. Sida 17: 333–342. Christenhusz MJM. 2012. Tropaeolum majus (Tropaeolaceae). Curtis’s Botanical Magazine 29: 331–340. Fay MF, Christenhusz MJM. 2010. Brassicales — an order of plants characterised by shared chemistry. Curtis’s Botanical Magazine 27: 165–196. Hershkovitz MA, Hernández-Pellicer CC, Arroyo MTK. 2006. Ribosomal DNA evidence for the diversification of Tropaeolum sect. Chilensia (Tropaeolaceae). Plant Systematics and Evolution 260: 1–24. King SR, Gershoff SN. 1987. Nutritional evaluation of three underexploited Andean tubers: Oxalis tuberosa (Oxalidaceae), Ullucus tuberosus (Basellaceae), and Tropaeolum tuberosum (Tropaeolaceae). Economic Botany 41: 503–511. Sparre B, Andersson L. 1991. A taxonomic revision of the Tropaeolaceae. Opera Botanica 108: 1–139. Rix M. 2012. Tropaeolum peregrinum (Tropaeolaceae). Curtis’s Botanical Magazine 29: 349–354. Watson JM, Flores AR. 2010. Tropaeolum section Chilensia: an overview (Tropaeolaceae). Curtis’s Botanical Magazine 27: 197–234.

292. MORINGACEAE HORSERADISH-TREE FAMILY Carlquist S 1998. Wood and bark anatomy of Caricaceae; correlations with systematics and habit. IAWA Journal 19: 191–206. Eilert U, Wolters B, Nahrstedt A. 1981. The antibiotic principle of seeds of Moringa oleifera and Moringa stenopetala. Planta Medica 42: 55–61. Ferguson IK. 1985. The pollen morphology of Moringaceae. Kew Bulletin 40: 25–34. Gandolfo MA, Nixon KC, Crepet WL. 1998. A new fossil f lower from the Turonian of New Jersey: Dressiantha bicapellata gen. et sp. nov. (Capparales). American Journal of Botany 85: 964–974. Olson ME. 1999. Moringa, the home page of the plant family Moringaceae. http://www.mobot.org/gradstudents/olson/moringahome.html Olson ME. 2002. Intergeneric relationships within the Caricaceae-Moringaceae clade (Brassicales), and potential morphological synapomorphies of the clade and its families. International Journal of Plant Sciences 163: 51–65. Olson ME. 2002. Combining data from DNA sequences and morphology for a phylogeny of Moringaceae. Systematic Botany 27: 55–73. Olson ME. 2003. Developmental origins of floral bilateral symmetry in Moringaceae. American Journal of Botany 90: 49–71. Verdcourt B. 1985. A synopsis of the Moringaceae. Kew Bulletin 40: 1–23. 293. CARICACEAE PAPAYA FAMILY Badillo VM. 1971. Monografía de la família Caricaceae. Asociación de profesores, Universidad Central de Venezuela, Facultad de Agronomía, Maracay. Badillo VM. 1993. Caricaceae. Segundo esquema. Revista de la Facultad de Agricultura. Universidad Central de Venezuela, Alcance (Maracay) 43: 1–111. Badillo VM. 2000. Carica L. vs. Vasconcella St.-Hil. (Caricaceae) con la rehabilitación de este último. Ernstia 10: 74–79. Carlquist S. 1998. Wood and bark anatomy of Caricaceae; correlations with systematics and habit. IAWA Journal 19: 191–206. Carvalho FA, Renner SS. 2012. A dated phylogeny of the papaya family (Caricaceae) reveals the crop’s closest relatives and the family’s biogeographic history. Molecular Phylogenetics and Evolution 65: 46–53. Christenhusz MJM, Van den Berg ERA. 1999. Caricaceae of the Guianas. Published by the authors, Culemborg. Fay MF, Christenhusz MJM. 2010. Brassicales — an order of plants characterised by shared chemistry. Curtis’s Botanical Magazine 27: 165–196. Jørgensen LB. 1995. Stomatal myrosin cells in Caricaceae. Taxonomic implications for a glucosinolate-containing family. Nordic Journal of Botany 15: 523–540. Kyndt T, Van Droogenbroeck B, Romeijn-Peeters E, Romero-Motochi JP, Scheldeman X, Goetghebeur P, Van Damme P, Gheysen G. 2005. Molecular phylogeny and evolution of Caricaceae based on rDNA internal transcribed spacers and chloroplast sequence data. Molecular Phylogenetics and Evolution 37: 442–459. Sawant AC. 1958. Crossing relationships in the genus Carica. Evolution 12: 263–266. Scheldeman X, Willemen L, Coppens d’Eeckenbrugge

Plants of the World

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FURTHER READING G, Romeijn-Peeters E, Restrepo MT, RomeroMotoche J, Jiménez D, Lobo M, Medina CI, Reyes C, Rodriguez D, Ocampo JA, Damme P, Goetghebeur P. 2007. Distribution, diversity and environmental adaptation of highland papayas (Vasconcellea spp.) in tropical and subtropical America. Biodiversity and Conservation 16: 1867–1884. Van Droogenbroeck B, Kyndt T, Romeijn-Peeters E, Van Thuyne W, Goetghebeur P, Romero.Motochi JP, Gheysen G. 2006. Evidence of natural hybridisation and introgression between Vasconcellea species (Caricaceae) from southern Ecuador revealed by chloroplast, mitochondrial and nuclear DNA markers. Annals of Botany 97: 793–805. 294. LIMNANTHACEAE MEADOWFOAM FAMILY Carlquist S, Donald CJ. 1996. Wood anatomy of Limnanthaceae and Tropaeolaceae in relation to habit and phylogeny. Sida 17: 333–342. Gentry HS, Miller RW. 1965. The search for new industrial crops IV. Prospectus of Limnanthes. Economic Botany 19: 25–32. Jain SK. 1976. Meadowfoams — mermaids of our vernal pools. Fremontia 4: 19–21. Link DA. 1992. The floral nectaries of Limnanthaceae. Plant Systematics and Evolution 179: 235–243. Mason CT. A systematic study of the genus Limnanthes. University of California Publications in Botany 25: 455–511. Ornduff R. 1971. Systematic studies of Limnanthaceae. Madroño 21: 103–111. Ornduff R, Crovello TJ. 1968. Numerical taxonomy of Limnanthaceae. American Journal of Botany 55: 173–182. Parker WH, Bohm BA. 1979. Flavonoids and taxonomy of Limnanthaceae. American Journal of Botany 66: 191–197. 295. SETCHELLANTHACEAE AZULITA FAMILY Carlquist S, Miller RB. 1999. Vegetative anatomy and relationships of Setchellanthus caeruleus (Setchellanthaceae). Taxon 48: 289–302. Iltis HH. 1999. Setchellanthaceae (Capparales), a new family for a relictual, glucosinulate-producing endemic of the Mexican deserts. Taxon 48: 257–275. Karol KG, Rodman JE, Conti E, Sytsma KJ. 1999. Nucleotide sequence of rbcL and phylogenetic relationships of Setchellanthus caeruleus (Setchellanthaceae). Taxon 48: 303–315. Tobe H, Carlquist S, Iltis HH. 1999. Reproductive anatomy and relationships of Setchellanthus caeruleus (Setchellanthaceae). Taxon 48: 277–283. Tomb S. 1999. Pollen morphology and relationships of Setchellanthus caeruleus (Setchellanthaceae). Taxon 48: 285–288. 296. KOEBERLINIACEAE ALLTHORN FAMILY Holmes WC, Yip KL, Rushing AE. 2008. Taxonomy of  Koeberlinia (Koeberliniaceae). Brittonia 60: 171–184. Mehta IJ, Moseley MF Jr. 1981. The floral anatomy of Koeberlinia Zucc.: systematic implications. American Journal of Botany 68: 482–497. Record S. 1926. The wood of Koeberlinia spinosa Zuccarini. Tropical Woods 8: 15–17. 297. BATACEAE TURTLEWEED FAMILY Behnke H-D, Turner BL. 1971. On specific sieve-tube plastids in Caryophyllales: further investigations

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with special reference to the Bataceae. Taxon 20: 731–737. Carlquist S. 1978. Wood anatomy and relationships of Bataceae, Gyrostemonaceae, and Stylobasiaceae. Allertonia 1: 297–330. Goldblatt P. 1976. Chromosome number and its significance in Batis maritima (Bataceae). Journal of the Arnold Arboretum 57: 526–530. Mabry TJ, Turner BL. 1964. Chemical investigations on the Batidaceae. Betaxanthins and their systematic implications. Taxon 13: 197–200. Rodman JE, Karol KG, Price RA, Sytsma KJ. 1996. Molecules, morphology, and Dahlgren’s expanded order Capparales. Systematic Botany 21: 289–307. Rogers GK. 1982. The Bataceae in the southeastern United States. Journal of the Arnold Arboretum 63: 375–386. Ronse De Craene LP. 2005. Floral developmental evidence for the systematic position of Batis (Bataceae). American Journal of Botany 92: 752–760. Van Heel WA. 1958. Additional investigations on Batis argillicola van Royen. Nova Guinea n.s. 9: 1–7. Van Royen P. 1956. A new Batidaceae, Batis argillicola. Nova Guinea s.n. 7: 187–195. 298. SALVADORACEAE TOOTHBRUSH-TREE FAMILY Carlquist S. 2002. Wood and bark anatomy of Salvadoraceae: ecology, relationships, history of interxylary phloem. Journal of the Torrey Botanical Society 129: 10–20. Kshetrapal S. 1970. A contribution to the vascular anatomy of the Salvadoraceae. Journal of the Indian Botanical Society 49: 92–99. Maheshwari Devi H. 1972. Salvadoraceae: a study of its embryology and systematics. Journal of the Indian Botanical Society 51: 56–62. Saxena V, Gupta S. 2011. Wood anatomy of the family Salvadoraceae from the Indian subcontinent with special reference to the ultrastructure of the vessel wall. Aliso 29: 59–63. Tobe H, Raven PH. 2012. Seed morphology and anatomy in Salvadoraceae (Brassicales): systematic and evolutionary implications. Acta Phytotaxomonica et Geobotanica 63: 1–9. 299. EMBLINGIACEAE SLIPPERCREEPER FAMILY Chandler GT, Bayer RJ. 2000. Phylogenetic placement of the enigmatic Western Australian genus Emblingia based on rbcL sequences. Plant Species Biology 15: 67–72. Erdtman G. Leins P, Melville R, Metcalfe CR. 1969. On the relationships of Emblingia. Botanical Journal of the Linnean Society 62: 169–186. Hall JC, Iltis HH, Sytsma KJ. 2004. Molecular phylogenetics of core Brassicales, placement of orphan genera Emblingia, Forchhammeria, Tirania, and character evolution. Systematic Botany 29: 654–669. Keighery GJ. 1981. The breeding system of Emblingia (Emblingiaceae). Plant Systematics and Evolution 137: 63–65. 300. TOVARIACEAE STINKBUSH FAMILY Boesewinkel FD. 1990. Ovule and seed development of Tovaria pendula Ruiz et Pavon. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 111: S389–S401. Carlquist S. 1985. Vegetative anatomy and familial placement of Tovaria. Aliso 11: 69–76.

Kjaer A. 1968. Glucosinolates in Tovariaceae. Phytochemistry 7: 131–133. 301. PENTADIPLANDRACEAE OUBLI FAMILY Berlec A, Jevnikar Z, Čanžek Majhernič A, Rogeij I, Štrukelj B. 2006. Expression of the sweet-tasting plant protein brazzein in Escherichia coli and Lactococcus lactis: a path toward sweet lactic acid bacteria. Applied Microbiology and Biotechnology 73: 158–165. Caldwell JE, Abildgaard F, Džakula Ž. Ming D, Hellekant G, Markley JL. 1998. Solution structure of the thermostable sweet-tasting protein brazzein. Nature Structural and Molecular Biology 5: 427–431. Lamphear BJ, Barker DK, Brooks CA, Delaney DE, Lane JR, Beifuss K, Love R, Thompson K, Mayor J, Clough R, Harkey R, Poage M, Drees C, Horn ME, Streatfield SJ, Nikolov Z, Woodard SL, Hood EE, Jilka JM, Howard JA. 2005. Expression of the sweet protein brazzein in maize for production of a new commercial sweetener. Journal of Plant Biotechnology 3: 103–114. Ming D, Hellekant G. 1994. Brazzein, a new highpotency thermostable sweet protein from Pentadiplandra brazzeana B. FEBS Letters 355: 106–108. Ronse De Craene LP. 2002. Floral development and anatomy of Pentadiplandra (Pentadiplandraceae): a key genus in the identification of floral morphological trends in the core Brassicales. Canadian Journal of Botany 80: 443–459. 302. GYROSTEMONACEAE BUTTONCREEPER FAMILY Carlquist S. 1978. Wood anatomy and relationships of Bataceae, Gyrostemonaceae, and Stylobasiaceae. Allertonia 1: 297–330. Goldblatt P, Nowicke JW, Mabry TJ, Behnke H-D. 1976. Gyrostemonaceae: status and affinity. Botaniska Notiser 129: 201–206. Hufford L. 1996. Developmental morphology of female flowers of Gyrostemon and Tersonia and floral evolution among Gyrostemonaceae. American Journal of Botany 83: 1471–1487. Keighery GJ. 1975. Chromosome numbers in the Gyrostemonaceae Endl. and the Phytolaccaceae Lindl.: a comparison. Australian Journal of Botany 23: 335–338. Lee DE, Lee WG, Mortimer N. 2001. Where and why have all the flowers gone? Depletion and turnover in the New Zealand Cenozoic angiosperm flora in relation to palaeogeography and climate. Australian Journal of Botany 49: 341–356. Rodman JE, Karol KG, Price RA, Conti E, Sytsma JD. 1994. Nucleotide sequences of rbcL confirm the capparalean affinity of the Australian endemic Gyrostemonaceae. Australian Systematic Botany 7: 57–69. Tobe H, Raven PH. 1991. The embryology and relationships of Gyrostemonaceae. Australian Systematic Botany 4: 407–420. 303. RESEDACEAE MIGNONETTE FAMILY Abdallah MS. 1967. The Resedaceae: a taxonomical revision of the family. Mededeelingen van de Landbouwhoogeschool te Wageningen 67(8): 1–98. Abdallah MS, De Wit HCD. 1978. The Resedaceae: a taxonomical revision of the family (final installment). Mededeelingen van de Landbouwhoogeschool te Wageningen 78(14): 99–416.

FURTHER READING Arber A. 1942. Studies in flower structure. VII. On the gynaeceum of Reseda with a consideration of paracarpy. Annals of Botany 6: 43–48. Carlquist S. 1998. Wood anatomy of Resedaceae. Aliso 16: 127–135. Carlquist S, Hansen BF, Iltis HH, Olson ME, Geiger DL. 2014. Forchhammeria and Stixis (Brassicales): stem and wood anatomical diversity, ecological and phylogenetic significance. Aliso 31: 59–75. El Naggar SM. 2002. Taxonomic significance of pollen morphology in some taxa of Resedaceae. Feddes Repertorium 113: 518–527. González Aguilera JJ, Fernández Peralta AM. 1984. Phylogenetic relationships in the family Resedaceae. Genetica 64: 185–197. Hall JC, Iltis HH, Sytsma KJ. 2004. Molecular phylogenetics of core Brassicales, placement of orphan genera Emblingia, Forchhammeria, Tirania, and character evolution. Systematic Botany 29: 654–669. Hennig L. 1929. Beiträge zur Kenntnis der Resedaceen–Blüte und Frucht. Planta 9: 507–563. Martín-Bravo S., Meimberg H, Luceño M, Märki W, Valcárcel V, Bräuchler C, Vargas P, Heubl G. 2007. Molecular systematics and biogeography of Resedaceae based on ITS and trnL-F sequences. Molecular Phylogenetics and Evolution 44: 1105–1120. Martín-Bravo S, Vargas P, Luceño M. 2009. Is Oligomeris (Resedaceae) indigenous to North America? Molecular evidence for a natural colonisation from the Old World. American Journal of Botany 96: 507–518. Mitra K, Mitra SN. 1979. Pollen morphology in relation to taxonomy and geography of Resedaceae. Bulletin of the Botanical Survey of India 18: 194–202. Schweingruber FH. 2006. Anatomical characteristics and ecological trends in the xylem and phloem of Brassicaceae and Resedaceae. IAWA Journal 27: 419–442. Sobick U. 1983. Blutenentwicklungsgeschichtliche Untersuchungen an Resedaceen unter besonderer Ber ucksichtigung von Androeceum und Gynoeceum. Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 104: 203–248. Su JX, Wang W, Zhang LB, Chen ZD. 2012. Phylogenetic placement of two enigmatic genera, Borthwickia and Stixis, based on molecular and pollen data, and the description of a new family of Brassicales, Borthwickiaceae. Taxon 61: 601–611. Weberling F. 1968. Über die Rudimentärstipeln der Resedaceae. Acta Botanica Neerlandica 17: 360–372. 304. CAPPARACEAE CAPER FAMILY Ernst WR. 1963. The genera of Capparidaceae and Moringaceae in the southeastern United States. Journal of the Arnold Arboretum 44: 81–95. Hall JC, Iltis HH, Sytsma KJ. 2004. Molecular phylogenetics of core Brassicales, placement of orphan genera Emblingia, Forchhammeria, Tirania, and character evolution. Systematic Botany. 29: 654–669. Hall JC, Sytsma KJ, Iltis HH. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. American Journal of Botany 89: 1826–1842. Hall JC. 2008. Systematics of Capparaceae and Cleomaceae: and evaluation of the generic delimitations of Capparis and Cleome using plastid DNA sequence data. Botany 86: 682–696.

Inocencio C, Cowan RS, Alcaraz F, Rivera F, Fay MF. 2005. AFLP fingerprinting in Capparis subgenus Capparis related to the commercial sources of capers. Genetic Resources and Crop Evolution 52: 137–144. Judd WS, Sanders RW, Donoghue MJ. 1994. Angiosperm family pairs: preliminary phylogenetic analyses. Harvard Papers in Botany 5: 1–51. Mithen R, Bennett R, Marquez J. 2010. Glucosinolate biochemical diversity and innovation in the Brassicales. Phytochemistry 71: 2074–2086. Rodman JE, Karol KG, Price RA, Sytsma KJ. 1996. Molecules, morphology, and Dahlgren’s expanded order Capparales. Systematic Botany 21: 289–307. 305. CLEOMACEAE SPIDERFLOWER FAMILY Bhide A, Schliesky S, Reich M, Weber APM, Becker A. 2014. Analysis of the floral transcriptome of Tarenaya hassleriana (Cleomaceae), a member of the sister group to the Brassicaceae: towards understanding the base of morphological diversity in Brassicales. BMC Genomics 15: 140 Cheng S, et al. (31 authors) 2013. The Tarenaya hassleriana genome provides insight into reproductive trait and genome evolution of crucifers. Plant Cell 8: 2813–2830. Ernst WR. 1963. The genera of Capparidaceae and Moringaceae in the southeastern United States. Journal of the Arnold Arboretum 44: 81–95. Feodorova TA, Voznesenskaya EV, Edwards GE, Roalson EH. 2010. Biogeographic patterns of diversification and the origins of C 4 in Cleome (Cleomaceae). Systematic Botany 35: 811–826. Hall JC, Sytsma KJ, Iltis HH. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. American Journal of Botany 89: 1826–1842. Hall JC. 2008. Systematics of Capparaceae and Cleomaceae: and evaluation of the generic delimitations of Capparis and Cleome using plastid DNA sequence data. Botany 86: 682–696. Hedge IC, Kjaer A, Malver O. 1980. Dipterygium — Cruciferae or Capparaceae? Notes from the Royal Botanic Gardens, Edinburgh 38: 247–250. Iltis HH. 1957. Studies in the Capparidaceae. III. Evolution and phylogeny of the western North American Cleomoideae. Annals of the Missouri Botanical Garden 44: 77–119. Marshall DM, Muhaidat R, Brown NJ, Liu Z, Stanley S, Griffith H, Sage RF, Hibbert JM. 2007. Cleome, a genus closely related to Arabidopsis, contains species spanning a developmental progression from C3 to C4 photosynthesis. The Plant Journal 51: 886–896. Patchell MJ, Roalson EH, Hall JC. 2014. Resolved phylogeny of Cleomaceae based on all three genomes. Taxon 63: 315–328. Riser JP II, Cardinal-McTeague WM, Hall JC, Hahn WJ, Sytsma KJ, Roalson EH. 2013. Phylogenetic relationships among the North American cleomoids (Cleomaceae): a test of Iltis’s reduction series. American Journal of Botany 100: 2102–2111. Sánchez-Acebo L. 2005. A phylogenetic study of the New World Cleome (Brassicaceae, Cleomoideae). Annals of the Missouri Botanical Garden 92: 179–201. Vanderpool SS, Elisens WJ, Estes JR. 1991. Pattern, tempo, and mode of evolutionary and biogeographic divergence in Oxystylis and Wislizenia (Capparaceae). American Journal of Botany 78: 925–937. Voznesenskaya EV, Koteyeva NK, Chuong DX,

Ivanova AV, Barroca J, Craven LA, Edwards GE. 2007. Physiological, anatomical and biochemical characterisation of photosynthetic types in the genus Cleome (Cleomaceae). Functional Plant Biology 34: 247–267. Woodson RE Jr. 1948.  Gynandropsis, Cleome, and´Podandrogyne. Annals of the Missouri Botanical Garden 35: 139–148. 306. BRASSICACEAE CABBAGE FAMILY Alexander PJ, Windham MD, Beck JB, Al-Shehbaz IA, Allphin L, Bailey CD. 2013. Molecular phylogenetics and taxonomy of the genus Boechera and related genera (Brassicaceae: Boechereae). Systematic Botany 38: 192–209. Al-Shehbaz IA. 1977. Protogyny in the Cruciferae. Systematic Botany 2: 327–333. Al-Shehbaz IA. 2012. A generic and tribal synopsis of the Brassicaceae (Cruciferae). Taxon 61: 931–954. Al-Shehbaz IA, Beilstein MA, Kellogg EA. 2006. Systematics and phylogeny of the Brassicaceae (Cruciferae): an overview. Plant Systematics and Evolution 259: 89–120. Arabidopsis Genome Initiative (AGI). 2000. Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408: 796–815. Arias T, Pires JC. 2012. A fully resolved phylogeny of the brassica crops and wild relatives (Brassicaceae: Brassiceae): novel clades and potential taxonomic implications. Taxon 61: 980–988. Bailey CD, Koch MA, Mayer M, Mummenhoff K, Okane SL Jr, Warwick SI, Windham MD, Al-Shehbaz IA. 2006. Toward a global phylogeny of the Brassicaceae. Molecular Biology and Evolution 23: 2142–2160. Bailey CD, Al-Shehbaz IA, Rajanikanth G. 2007. Generic limits in the tribe Halimolobeae and the description of the new genus Exhalimolobos (Brassicaceae). Systematic Botany 32: 140–156. Barber JT. 1978. Capsella bursa-pastoris seeds: are they “carnivorous”? Carnivorous Plant Newsletter 7(2): 39–42. Beilstein MA, Al-Shehbaz IA, Kellogg AE. 2006. Brassicaceae phylogeny and trichome evolution. American Journal of Botany 93: 607–619. Beilstein MA, Al-Shehbaz IA, Mathews S, Kellogg EA. 2008. Brassicaceae phylogeny inferred from phytochrome A and ndhF sequence data: tribes and trichomes revisited. American Journal of Botany 95: 1307–1327. Beilstein MA, Nagalingum NS, Clements MD, Manchester SR, Mathews S. 2010. Dated molecular phylogenies indicate a Miocene origin for Arabidopsis thaliana. Proceedings of the National Academy of Sciences of the USA 107: 18724–18728. Bennett MD, Leitch IJ, Price HJ, Johnston JS. 2003. Comparisons with Caenorabditis (~100 Mb) and Drosophila (~175 Mb) using flow cytometry show genome size in Arabidopsis to be ~157Mb and thus ~25% larger than the Arabidopsis Genome Initiative of ~125Mb. Annals of Botany 91: 1–11. Bowman JL. 2006. Molecules and morphology: comparative developmental genetics of the Brassicaceae. Plant Systematics and Evolution 259: 199–215. Clauss MJ, Koch MA. 2006. Poorly known relatives of Arabidopsis thaliana. Trends in Plant Science 11: 449–459. Fuentes-Soriano S, Al-Shehbaz IA. 2013. Phylogenetic relationships of mustards with multiaperturate pollen (Physarieae, Brassicaceae) based on the plastud ndhF gene: implications for morphological diversification. Systematic Botany 38: 178–191.

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FURTHER READING Hall JC, Sytsma KJ, Iltis HH. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. American Journal of Botany 89: 1826–1842. Hall JC, Tisdale TE, Donohue K, Wheeler A, Al-Yahya MA, Kramer EM. 2011. Convergent evolution of a complex fruit structure in the tribe Brassiceae (Brassicaceae). American Journal of Botany 98: 1989–2003. Johnston JS, Pepper AE, Hall AE, Chen ZJ, Hodnett G, Drabek J, Lopez R, Price HJ. 2005. Evolution of genome size in Brassicaceae. Annals of Botany 95: 229–235. Jordon-Thaden I, Hase I, Al-Shehbaz IA, Koch MA. 2010. Molecular phylogeny and systematics of the genus Draba (Brassicaceae) and identification of its most closely related genera. Molecular Phylogenetics and Evolution 55: 524–540. Kiefer M, Schmicki R, German DA, Lysák M, Al-Shehbaz IA, Franzke A, Mummenhoff K, Stamatakis A, Koch MA. 2014. BrassiBase: introduction to a novel database on Brassicaceae evolution. Plant and Cell Physiology 55: e3. Koch M, Al-Shehbaz IA, Mummenhoff K. 2003. Molecular systematics, evolution, and population biology in the mustard family (Brassicaceae). Annals of the Missouri Botanical Garden 90: 151–171. Koch M, Haubold B, Mitchell-Olds T. 2000. Comparative analysis of chalcone synthase and alcohol dehydrogenase loci in Arabidopsis, Arabis and related genera (Brassicaceae). Molecular Biology and Evolution 17: 1483–1498. Koch MA, Karl R, Kiefer M, Al-Shehbaz IA. 2010. Colonizing the American continent: systematics of the genus Arabis in North America (Brassicaceae). American Journal of Botany 97: 1040–1057. Koch MA, Marhold K. 2012. Phylogeny and systematics of Brassicaceae — Introduction. Taxon 61: 929–930. (see also other paper in this special issue of Taxon). Krause EHL. 1902. Brassicaceae in Sturm J (ed.) Deutschlands Flora, Abt. II, Cryptogamie 6. Kosten der Verfassers, Nürnberg. Lysák MA, Lexer C. 2006. Towards the era of comparative evolutionary genomics in Brassicaceae. Plant Systematics and Evolution 259: 175–198. Lysák MA, Koch MA, Beaulieu JM, Meister A, Leitch IJ. 2009. The dynamic ups and downs of chromosome size evolution in Brassicaceae. Molecular Biology and Evolution 26: 85–98. Meyer FK. 2003. Kritische Revision der ‘Thlaspi’ — Arten Europas, Afrikas und Voderasienes. Hausknechtia 9: 115–124. Meyerowitz EM. 2001. Prehistory and history of Arabidopsis research. Plant Physiology 125: 15–19. O’Kane SL Jr, Al-Shehbaz IA. 2003. Phylogenetic position and generic limits of Arabidopsis (Brassicaceae) based on sequences of nuclear ribosomal DNA. Annals of the Missouri Botanical Garden 90: 603–612. Resetnik I, Satovic Z, Schneeweiss GM, Liber Z. 2013. Phylogenetic relationships in Brassicaceae tribe Alysseae inferred from nuclear ribosomal and chloroplast DNA sequence data. Molecular Phylogenetics and Evolution 69: 772–786. Rollins RC, Banerjee UC. 1979. Pollen of the Cruciferae. Publications of the Bussey Institution, Harvard University 1979: 33–64. Warwick SI, Sauder CA. 2005. Phylogeny of tribe Brassiceae (Brassicaceae) based on chloroplast restriction site polymorphisms and nuclear

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ribosomal internal transcribed spacer and chloroplast trnL intron sequences. Canadian Journal of Botany 83: 467–483. War wick SI, Al-Shehbaz IA, Sauder CA. 2006. Phylogenetic position of Arabis arenicola and generic limits of Eutrema and Aphragmus (Brassicaceae) based on sequences of nuclear ribosomal DNA. Canadian Journal of Botany 84: 269–281. Warwick SI, Francis A, Al-Shehbaz IA. 2006b. Brassicaceae: species checklist and database on CD-ROM. Plant Systematics and Evolution 259: 249–258. Warwick SI, Sauder CA, Al-Shehbaz IA, Jacquemoud F. 2007. Phylogenetic relationships in the tribes Anchonieae, Chorisporeae, Euclidieae, and Hesperideae (Brassicaceae) based on nuclear ribosomal ITS DNA sequences. Annals of the Missouri Botanical Garden 94: 56–78. Warwick SI, Mummenhoff K, Saunder CA, Koch MA, Al-Shehbaz IA. 2010. Closing the gaps: phylogenetic relationships in the Brassicaceae based on DNA sequence data of nuclear ribosomal ITS region. Plant Systematics and Evolution 285: 209–232. 307. AEXTOXICACEAE OLIVILLO FAMILY Carlquist S. 2003. Wood anatomy of Aextoxicaceae and Berberidopsidaceae is compatible with their inclusion in Berberidopsidales. Systematic Botany 28: 317–325. Ronse de Craene LP, Stuppy W. 2010. Floral development and anatomy of Aextoxicon punctatum (Aextoxicaceae-Berberidopsidales): an enigmatic tree at the base of the core eudicots. International Journal of Plant Sciences 171: 244–257. 308. BERBERIDOPSIDACEAE CORAL-VINE FAMILY Baas P. 1984. Vegetative anatomy and taxonomy of Berberidopsis and Streptothamnus (Flacourtiaceae). Blumea 30: 39–44. Carlquist S. 2003. Wood anatomy of Aextoxicaceae and Berberidopsidaceae is compatible with their inclusion in Berberidopsidales. Systematic Botany 28: 317–325. Ronse de Craene LP. 2004. Floral development of Berberidopsis corallina: a crucial link in the evolution of flowers in the core eudicots. Annals of Botany 94: 741–751. Van Heel WA. 1984. Flowers and fruits in Flacourtiaceae. V. The seed anatomy and pollen morphology of Berberidopsis and Streptothamnus. Blumea 30: 31–37. Veldkamp JF. 1984. Berberidopsis (Flacourtiaceae) in Australia. Blumea 30: 21–29. 309. OLACACEAE TALLOW-WOOD FAMILY Baas P, Van Oosterhout E, Scholtes CJL. 1982. Leaf anatomy and classification of the Olacaceae, Octoknema and Erythropalum. Allertonia 3: 155–210. Kuo J, Pate JS, Davidson NJ. 1989. Ultrastructure of the haustorial interface and apoplastic continuum between host and the root hemiparasite Olax phyllanthi (Labill.) R. Br. (Olacaceae). Protoplasma 150: 27–39. Maas PJM, Baas P, Boesewinkel FD, Hiepko P, Lobreau-Callen D, Van den Oever L, Ter Welle BJH. 1992. The identity of “unknown Z”: Maburea Maas, a new genus of Olacaceae in Guyana. Botanische Jahrbücher für Systematik, Pflanzengeschichte und

Pflanzengeographie 114: 275–291. Malécot V, Lobreau-Callen D. 2005. A survey of the species assigned to the fossil pollen genus Anacolosidites. Grana 44: 314–336. Malécot V, Nickrent DL. 2008. Molecular phylogenetic relationships of Olacaceae and related Santalales. Systematic Botany 33: 97–106. Malécot V, Nickrent DL, Baas P, Van den Oever L, Lobreau-Callen D. 2004. A morphological cladistic analysis of Olacaceae. Systematic Botany 29: 569–586. Nickrent DL, Malécot V, Vidal-Russell R, Der JP. 2010. A revised classification of Santalales. Taxon 59: 538–558. Pate JS, Davidson NJ, Kuo J, Milburn JA. 1990. Water relations of the root hemiparasite Olax phyllanthi (Labill.) R.Br. (Olacaceae) and its multiple hosts. Oecologia 84: 186–193. Pate JS, Pate SR, Kuo J, Davidson NJ. 1990. Growth, resource allocation and haustorial biology of the root hemiparasite Olax phyllanthi (Olacaceae). Annals of Botany 65: 437–449. 310. OPILIACEAE BALLY-COMA FAMILY Der JP, Nickrent DL. 2008. A molecular phylogeny of Santalaceae. Systematic Botany 33: 107–116. Hiepko P. 1984. Opiliaceae. Flora Malesiana ser. 1, vol. 10. Hiepko P. 2000. Opiliaceae. Flora Neotropica Monograph 82. Williams LO. 1966. The agonandras of Mexico and Central America. Ciencia (Mexico) 24: 227–228. 311. BALANOPHORACEAE SNAKEMUSHROOM FAMILY Barkman TJ, McNeal JR, Lim S-H, Coat G, Croom HB, Young ND, dePamphilis CW. 2007. Mitochondrial DNA suggests at least 11 origins of parasitism in angiosperms and reveals genomic chimerisms in parasitic plants. BMC Evolutionary Biology 7: 248. Borchsenius F, Olesen JM. 1990. The Amazonian root holoparasite Lophophytum mirabile (Balanophoraceae) and its pollinators and herbivores. Journal of Tropical Ecology 6: 501–505. Eberwein R, Nickrent DL, Weber A. 2009. Development and morphology of flowers and inflorescences in Balanophora papuana and B. elongata (Balanophoraceae). American Journal of Botany 96: 1055–1067. González AM, Mauseth JD. 2010. Morphogenesis is highly aberrant in the vegetative body of the holoparasite Lophophytum leandri (Balanophoraceae): all typical vegetative organs are absent and many tissues are highly modified. International Journal of Plant Sciences 171: 499–508. Holzapfel S. 2001. Studies of the New Zealand rootparasite Dactylanthus taylorii (Balanophoraceae). Englera 22: 1–107. Hooker JD. 1856. On the structure and affinities of Balanophoraceae. Transactions from the Linnean Society of London 22: 1–68. Kuijt J, Dong W-X. 1990. Surface features of the leaves of Balanophoraceae — a family without stomata? Plant Systematics and Evolution 170: 29–35. Su HJ, Hu JM. 2012. Rate heterogeneity in six proteincoding genes from the holoparasite Balanophora (Balanophoraceae) and other taxa of Santalales. Annals of Botany 110: 1137–1147. 312. SANTALACEAE SANDALWOOD FAMILY Der JP, Nickrent DL. 2008. A molecular phylogeny of Santalaceae. Systematic Botany 33: 107–116.

FURTHER READING Harbaugh DT, Baldwin BG. 2007. Phylogeny and biogeography of sandalwoods (Santalum, Santalaceae): repeated dispersals throughout the Pacific. American Journal of Botany 94: 1028–1040. Kuijt J. 2003. Monograph of Phoradendron (Viscaceae). Systematic Botany Monographs 66. Moore TE, Verboom GA, Forest F. 2010. Phylogenetics and biogeography of the parasitic genus Thesium (Santalaceae), with an emphasis on the Cape of South Africa. Botanical Journal of the Linnean Society 162: 435–452. Mwang’ingo PL, Teklehaimanot Z, Hall JB, Zilihona JE. 2007. Sex distribution, reproductive biology and regeneration in the dioecious species Osyris lanceolata (African sandalwood) in Tanzania. Tanzania Journal of Forestry and Nature Conservation 76: 118–133. Nickrent DL, García MP. 2009. On the brink of holoparasitism: plastome evolution in dwarf mistletoes (Arceuthobium, Viscaceae). Journal of Molecular Evolution 68: 603–615. Nickrent DL, García MP, Martin RL, Mathiasen RL. 2004. A phylogeny of all species of Arceuthobium (Viscaceae) using nuclear and chloroplast DNA sequences. American Journal of Botany 91: 125–138. Norveto CA. 2011. Study of the comparative wood anatomy of the species of Amphorogynaceae, Cervantesiaceae, Nanodeaceae, Santalaceae and Thesiaceae. Journal of the Botanical Research Institute of Texas 5: 643–659. Pate JS. 2001. Haustoria in action: case studies of nitrogen acquisition by woody xylem-tapping hemiparasites from their hosts. Protoplasma 215: 204–217. Press MC, Graves JD (eds). 1995. Parasitic plants. Chapman, Hall, London. Tennakoon KU, Pate JS, Arthur D. 1997. Ecophysiological aspects of the woody root hemiparasite Santalum acuminatum (R. Br.) A. DC. and its common hosts in southwestern Australia. Annals of Botany 80: 245–256. Vidal-Russell R, Nickrent DL. 2008. The first mistletoes: origin of aerial parasitism in Santalales. Molecular Phylogenetics and Evolution 47: 523–537. Wilson CA, Calvin CL. 2003. Development, taxonomic significance and ecological role of the cuticular epithelium in the Santalales. IAWA Journal 24: 129–138.

V. 2013. Host use and resource sharing by fruit/ seed-infesting insects on Schoepfia schreberi (Olacaceae). Environmental Entomology 42: 231–239. Malécot V, Nickrent DL. 2008. Molecular phylogenetic relationships of Olacaceae and related Santalales. Systematic Botany 33: 97–106. Werth CR, Baird WV, Musselman LJ. 1979. Root parasitism in Schoepfia Schreb. (Olacaceae). Biotropica 11: 140–143.

313. MISODENDRACEAE FEATHERYMISTLETOE FAMILY Carlquist S. 1985. Wood and stem anatomy of Misodendraceae: systematic and ecological conclusions. Brittonia 37: 58–75. Kuijt J. 1969. The biology of parasitic flowering plants. University of California, Berkeley. Vidal-Russell R, Nickrent DL. 2007. A molecular phylogeny of the feathery mistletoe Misodendrum. Systematic Botany 32: 560–568.

315. LORANTHACEAE SHOWY-MISTLETOE FAMILY Barlow BA, Wiens D. 1977. Host-parasite resemblance in Australian mistletoes: the case for cryptic mimicry. Evolution 31: 69–84. Fay MF, Bennett JR, Dixon KW, Christenhusz MJM. 2010. Parasites, their relationships and the disintegration of Scrophulariaceae sensu lato. Curtis’s Botanical Magazine 26: 286–313. Feuer S, Kuijt J. 1985. Fine structure of mistletoe pollen. VI. Small-f lowered Neotropical Loranthaceae. Annals of the Missouri Botanical Garden 72: 187–212. Hopper SD. 2010. 660. Nuytsia floribunda. Curtis’s Botanical Magazine 26: 333–368. Johri BM, Bhatnagar SP. 1972. Loranthaceae. Council of Scientific and Industrial Research, New Delhi. Kuijt J. 2009. Monograph of Psittacanthus (Loranthaceae). Systematic Botany Monographs 86. Kuijt J. 2010. A note on stamen position and petal number in Loranthaceae. Blumea 55: 224–225. Kuijt J. 2011. Pulling the skeleton out of the closet: resurrection of Phthirusa Martius and the consequent revival of Passovia (Loranthaceae). Plant Diversity and Evolution 129: 159–211. Mathiasen RL, Nickrent DL, Shaw DC, Watson DM. 2008. Mistletoes: pathology, systematics, ecology, and management. Plant Disease 92: 988–1006. Nickrent DL, Malécot V, Vidal-Russell R, Der JP. 2010. A revised classification of Santalales. Taxon 59: 538–558. Polhill RM, Weins D. 1998. Mistletoes of Africa. Royal Botanic Gardens, Kew. Press MC, Phoenix GK. 2005. Impacts of parasitic plants on natural communities. New Phytologist 166: 737–751. Vidal-Russell R, Nickrent DL. 2008. Evolutionary relationships in the showy mistletoe family (Loranthaceae). American Journal of Botany 95: 1015–1029. Visser J. 1981. South African parasitic flowering plants. Juta, Cape Town. Wilson CA, Calvin CL. 2006. Character divergences and convergences in canopy-dwelling Loranthaceae. Botanical Journal of the Linnean Society 150: 101–113. Wilson CA, Calvin CL. 2006. An origin of aerial branch parasitism in the mistletoe family, Loranthaceae. American Journal of Botany 93: 787–796.

314. SCHOEPFIACEAE WHITEWOOD FAMILY Der JP, Nickrent DL. 2008. A molecular phylogeny of Santalaceae. Systematic Botany 33: 107–116. Huang CF, Gan XW, Bai HY, Hu LH. 2008. Schoepfin A, B, C: three new chalcone C-glycosides from Schoepfia chinensis. Natural Product Research 22: 623–627. López-Ortega M, Pérez-Rodríguez P, Rojas JC, Hernández RMS, López-Mata L, Rico-Gray

316. FRANKENIACEAE SEA-HEATH FAMILY Brochmann C, Lobin W, Sunding P, Stabbetorp OE. 1995. Parallel ecoclinal evolution and taxonomy of Frankenia (Frankeniaceae) in the Cape Verde Islands, W Africa. Nordic Journal of Botany 15: 603–623. Campbell N, Thomson WW. 1976. The ultrastructure of Frankenia salt glands. Annals of Botany II, 40: 681–686.

Gaskin JF, Ghahremaninejad F, Zhang D-Y, Londo JP. 2004. A systematic overview of Frankeniaceae and Tamaricaceae from nuclear rDNA and plastid sequence data. Annals of the Missouri Botanica Garden 91: 402–410. Olson ME, Gaskin JF, Ghahremaninejad F. 2003. Stem anatomy is congruent with molecular phylogenies placing Hypericopsis persica in Frankenia (Frankeniaceae): comments on vasicentric tracheids. Taxon 52: 525–532. Summerhayes VS. 1930. A revision of the Australian species of Frankenia. Journal of the Linnean Society, Botany 48: 337–387. Walia K, Kapil RN. 1965. Embryology of Frankenia Lin. with some comments on the systematic position of the Frankeniaceae. Botaniska Notiser 118: 412–429. Whalen MA. 1987. Wood anatomy of the American frankenias (Frankeniaceae): systematic and evolutionary implications. American Journal of Botany 74: 1211–1223. Whalen MA. 1989. Systematics of Frankenia (Frankeniaceae) in North and South America. Systematic Botany Monographs 17. 317. TAMARICACEAE SALTCEDAR FAMILY Baum BR. 1978. The genus Tamarix. Israel Academy of Sciences and Humanities, Jerusalem. Gaskin JF, Ghahremaninejad F, Zhang DY, Londo JP. 2004. A systematic overview of Frankeniaceae and Tamaricaceae from nuclear rDNA and plastid sequence data. Annals of the Missouri Botanical Garden 91: 402–410. Gupta AK, Murty YS. 1987. Floral anatomy in Tamaricaceae. Journal of the Indian Botanical Society 66: 275–282. Qaiser M. 1987. Studies in the seed morphology of the family Tamaricaceae from Pakistan. Botanical Journal of the Linnean Society 94: 469–484. Ronse Decraene LP. 1990. Morphological studies in Tamaricales. I. Floral ontogeny and anatomy of Reaumuria vermiculata L. Beiträge zur Biologie der Pflanzen 65: 181–203. Waly NM. 1999. Wood anatomical characters of the Egyptian Tamarix L. species and its taxonomic significance. Taeckholmia 19: 115–125. Wang Y, Liu Y, Liu S, Huang H. 2009. Molecular phylogeny of Myricaria (Tamaricaceae): implications for taxonomy and conservation in China. Botanical Studies 50: 343–352. Zhang D, Chen Z, Sun H, Yin L, Pan B. 2000. Systematic studies on some questions of Tamaricaceae based on ITS sequence. Acta Botanica Boreali-Occidentalia Sinica 20: 421–431. 318. PLUMBAGINACEAE THRIFT FAMILY Carlquist S, Boggs CJ. 1996. Wood anatomy of Plumbaginaceae. Bulletin of the Torrey Botanical Club 123: 135–147. De Laet J, Clinckemaille J, Jansen S, Smets E. 1995. Floral ontogeny of the Plumbaginaceae. Journal of Plant Research 108: 289–304. Flowers TJ, Colmer TD. 2008. Salinity tolerance in halophytes. New Phytologist 179: 945–963. Hanson AD, Rathinasabapathi B, Rivoal J, Burnet M, Dillon MO, Gage DA. 1994. Osmoprotective compounds in the Plumbaginaceae: a natural experiment in metabolic engineering of stress tolerance. Proceedings of the National Academy of Sciences of the U.S.A. 91: 206–310. Lledó MD, Crespo MB, Cameron KM, Fay MF, Chase MW. 1998. Systematics of Plumbaginaceae based

Plants of the World

727

FURTHER READING upon cladistic analysis of rbcL sequence data. Systematic Botany 23: 21–29. Lledó MD, Karis PO, Crespo MB, Fay MF, Chase MW. 2001. Phylogenetic position and taxonomic status of the genus Aegialitis and subfamilies Staticoideae and Plumbaginoideae (Plumbaginaceae): evidence from plastid DNA sequences and morphology. Plant Systematics and Evolution 229: 107–124. Lledó MD, Crespo MB, Fay MF, Chase MW. 2005. Molecular phylogenetics of Limonium and related genera (Plumbaginaceae): biogeographical and systematic implications. American Journal of Botany 92: 1189–1198. Nowicke J, Skvarla JJ. 1977. Pollen morphology and the relationship of the Plumbaginaceae, Polygonaceae, and Primulaceae to the order Centrospermae. Smithsonian Contributions to Botany 37: 1–64. 319. POLYGONACEAE KNOTWEED FAMILY Burke JM, Sanchez A, Kron KA, Luckow M. 2010. Placing the woody genera of Polygonaceae: a hypothesis of character evolution and phylogeny. American Journal of Botany 97: 1377–1390. Carlquist S. 2003. Wood anatomy of Polygonaceae: analysis of a family with exceptional wood diversity. Botanical Journal of the Linnean Society 141: 25–51. Cuñado N, Navajas-Pérez R, De la Herrán R, Ruiz Rejón C, Ruiz Rejón M, Santos JL, Garrido-Ramos MA. 2007. The evolution of sex chromosomes in the genus Rumex (Polygonaceae): identification of a new species with heteromorphic sex chromosomes. Chromosome Research 15: 825–832. Fan DM, Chen JM, Meng Y, Wen J, Huang JL, Yang YP. 2013. Molecular phylogeny of Koenigia L. (Polygonaceae: Persicarieae): implications for classification, character evolution and biogeography. Molecular Phylogenetics and Evolution 69: 1093–1100. Galasso G, Banfi E, De Mattia F, Grassi F, Sgorbati S, Labra M. 2009. Molecular phylogeny of Polygonum L. s.l. (Polygonoideae, Polygonaceae), focusing on European taxa: preliminary results and systematic considerations based on rbcL plastidial sequence data. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 15: 113–148. Kempton EA. 2013. Systematics of Eriogonoideae s.s. (Polygonaceae). Systematic Botany 37: 723–737. Kostikova A, Litsois G, Burgy S, Milani L, Pearman PB, Salamin N. 2014. Scale-dependent adaptive evolution and morphological convergence to climatic niche in Californian eriogonoids (Polygonaceae). Journal of Biogeography 41: 1326–1337. Lamb Frye AS, Kron KA. 2003. rbcL phylogeny and character evolution in Polygonaceae. Systematic Botany 28: 326–332. Sanchez A, Kron KA. 2009. Phylogenetic relationships of Afrobrunnichia Hutch., Dalziel (Polygonaceae) based on three chloroplast genes and ITS. Taxon 58: 781–792. Sanchez A, Schuster TM, Burke JM, Kron KA. 2011. Taxonomy of Polygonoideae (Polygonaceae): a new tribal classification. Taxon 60: 151–160. Sanchez A, Schuster TM, Kron KA. 2009. A large scale phylogeny of Polygonaceae based on molecular data. International Journal of Plant Sciences 170: 1044–1055. Schuster TM, Reveal JL, Kron KA. 2011. Phylogeny of Polygoneae (Polygonaceae: Polygonoideae). Taxon 60: 1653–1666.

728

Christenhusz, Fay & Chase

Schuster TM, Setaro SD, Kron KA. 2013. Age estimates for the buckwheat family Polygonaceae based on sequence data calibrated by fossils and with a focus on the amphi-Pacific Muehlenbeckia. PLoS ONE 8: e61261. Sun Y, Wang A, Wan D, Wang Q, Liu J. 2012. Rapid radiation of Rheum (Polygonaceae) and parallel evolution of morphological traits. Molecular Phylogenetics and Evolution 63: 150–158. Tian X, Luo J, Wang A, Mao K, Liu J. 2011. On the origin of the woody buckwheat Fagopyrum tibeticum (=Parapteropyrum tibeticum) in the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution 61: 515–520. Wang A, Yang M, Liu J. 2005. Molecular phylogeny, recent radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequences. Annals of Botany 96: 489–498. 320. DROSERACEAE SUNDEW FAMILY Albert VA, Williams SE, Chase MW. 1992. Carnivorous plants: phylogeny and structural evolution. Science 257: 1491–1495. Cameron KM, Wurdack KJ, Jobson RW. 2002. Molecular evidence for the common origin of snaptraps among carnivorous plants. American Journal of Botany 89: 1503–1509. Chase MW, Christenhusz MJM, Sanders D, Fay MF. 2010. Murderous plants: Victorian Gothic, Darwin and modern insights into vegetable carnivory. Botanical Journal of the Linnean Society 161: 329–356. Conran JG, Jaudzems VG, Hallam ND. 1997. Droseraceae germination patterns and their taxonomic significance. Botanical Journal of the Linnean Society 123: 211–223. Darwin C. 1875. Insectivorous plants. John Murray, London. Darwin C. 1880. The power of movement in plants. John Murray, London. Dixon KW, Pate SJ. 1978. Phenology, morphology and reproductive biology of the tuberous sundews, Drosera erythrorhiza Lindl. Australian Journal of Botany 26: 441–454. Escalante-Pérez M, Krol E, Stange A, Geiger D, Al-Rasheid KAS, Hause B, Neher E, Hedrich R. 2011. A special pair of phytohormones controls excitability, slow closure, and external stomach formation in the Venus flytrap. Proceedings of the National Academy of Science of the USA 108: 15492–15497. Forterre Y, Skotheim JM, Dumais J, Mahadevan L. 2005. How the Venus flytrap snaps. Nature 433: 421–425. Gibson TC, Waller DM. 2009. Evolving Darwin’s ‘most wonderful’ plant: ecological steps to a snaptrap. New Phytologist 183: 575–587. Hartmeyer I, Hartmeyer SRH. 2010. Snap-tentacles and runway lights: summary of comparative examination of Drosera tentacles. Carnivorous Plants Newsletter 39: 101–113. Heslop-Harrison Y. 1976. Carnivorous plants, a century after Darwin. Endeavour 126: 114–122. Juniper BE, Robins RJ, Joel DM. 1989. The carnivorous plants. Academic Press, London. McPherson S. 2008. Glistening carnivores. The stickyleaved insect-eating plants. Redfern Natural History Productions, Poole. Płachno BJ, Adamec L, Lichtscheidl IK, Peroutka M, Adlassnig W, Vrba J. 2006. Fluorescence labelling of phosphatase activity in digestive glands of

carnivorous plants. Plant Biology 8: 813–820. Poppinga S, Joyeux M. 2011. Different mechanisms of snap-trapping in two closely related carnivorous plants Dionaea muscipula and Aldrovanda vesiculosa. Physics Review E 84: 041926. Poppinga S, Hartmeyer SRH, Seidel R, Masselter T, Hartmeyer I, Speck T. 2012. Catapulting tentacles in a sticky carnivorous plant. PLoS ONE 7: e45735. Rivadavia F, Kondo K, Kato M, Hasebe M. 2003. Phylogeny of the sundews, Drosera (Droseraceae), based on chloroplast rbcL and nuclear 18S ribosomal DNA sequences. American Journal of Botany 90: 123–130. Rivadavia F, De Miranda VFO, Hoogenstrijd G, Pinheiro F, Heubl G, Fleischmann A. 2012. Is Drosera meristocaulis a pygmy sundew? Evidence of long-distance dispersal between Western Australia and northern South America. Annals of Botany 110: 11–21. Seine R, Barthlott W. 1993. On the morphology of trichomes and tentacles of Droseraceae Salisb. Beiträge zur Biologie der Pflanzen 67: 345–366. Williams SE, Albert VA, Chase MW. 1994. Relationships of Droseraceae: a cladistic analysis of rbcL sequence and morphological data. American Journal of Botany 1027–1037. 321. NEPENTHACEAE ASIAN-PITCHERPLANT FAMILY Adlassnig W, Peroutka M, Lendl T. 2011. Traps of carnivorous plants as a habitat: composition of the fluid, biodiversity and mutualistic activities. Annals of Botany 107: 181–194. Alamsyah F, Ito M. 2014. Phylogenetic analysis of Nepenthaceae based on internal transcribed spacer (ITS) nrDNA sequences. Acta Phytotaxonomica et Geobotanica 64: 113–126. Carlquist S. 1981. Wood anatomy of Nepenthaceae. Bulletin of the Torrey Botanical Club 108: 324–330. Cheek M, Jebb M. 2001. Nepenthaceae. Flora Malesiana I, 15: 1–157. Chin L, Moran JA, Clarke C. 2010. Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size. New Phytologist 186: 461–470. Clarke CM, Bauer U, Lee CC, Tuen AA, Rembold K, Moran JA. 2009. Tree shrew lavatories: a novel nitrogen sequestration strategy in a tropical pitcher plant. Biology Letters 5: 632–635. Heslop-Harrison Y. 1976. Carnivorous plants, a century after Darwin. Endeavour 126: 114–122. Heubl GR, Wistuba A, 1997. A cytological study of the genus Nepenthes L. (Nepenthaceae). Sendtnera 4: 169–174. Jebb M, Cheek M. 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42: 1–106. Juniper BE, Robins RJ, Joel DM. 1989. The carnivorous plants. Academic Press, London. Krutzsch W. 1989. Paleogeography and historical geography (paleochorology) in the Neophyticum. Plant Systematics and Evolution 162: 5–61. McPherson S. 2009. Pitcher plants of the Old World (2 vols). Redfern Natural History Productions, Poole. Meimberg H, Heubl G. 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology 8: 831–840. Moran JA, Merbach MA, Livingston NJ, Clarke CM, Booth WE. 2001. Termite prey specialisation in the pitcher plant Nepenthes albomarginata —evidence from stable isotope analysis. Annals of Botany 88: 307–311.

FURTHER READING Pavlovic A, Masarovicová E, Hudák J. 2007. Carnivorous syndrome in Asian pitcher plants of the genus Nepenthes. Annals of Botany 100: 527–536. 322. DROSOPHYLLACEAE DEW-PINE FAMILY Adamec L. 2009. Ecophysiological investigation on Drosophyllum lusitanicum: why doesn’t the plant dry out? Carnivorous Plant Newsletter 38: 71–74. Carlquist S, Wilson EJ. 1995. Wood anatomy of Drosophyllum (Droseraceae): ecological and phylogenetic considerations. Bulletin of the Torrey Botanical Club 122: 185–189, Cuénoud P, Savolainen V. Chatrou LW, Powell M, Grayer RJ, Chase 2002. Molecular phylogeny of Caryophyllales based on 18S rDNA, rbcL, atpB, and matK. American Journal of Botany 89: 132–144. Hoshi Y, Hotta K. 1998. A chromosome phylogeny of the Droseraceae by using CMA-DAPI fluorescent banding. Cytologia 63: 329–339. McPherson S. 2008. Glistening carnivores. The stickyleaved insect-eating plants. Redfern Natural History Productions, Poole. Ortega Olivencia A, Carrasco Claver JP, Devesa Alcarez JA. 1995. Floral and reproductive biology of Drosophyllum lusitanicum (L.) Link (Droseraceae). Botanical Journal of the Linnean Society 118: 331–351. Płachno BJ, Adamec L, Huet H. 2009. Mineral nutrient uptake from prey and glandular phosphatase activity as a dual test of carnivory in semidesert plants with glandular leaves suspected of carnivory. Annals of Botany 104: 649–654. Williams SE, Albert VA, Chase MW. 1994. Relationships of Droseraceae: a cladistic analysis of rbcL sequence and morphological data. American Journal of Botany 1027–1037. 323. DIONCOPHYLLACEAE HOOKLEAF-VINE FAMILY Airy Shaw HK. 1952. On the Dioncophyllaceae, a remarkable new family of flowering plants. Kew Bulletin 1951: 327–347. Bringman G, Wenzel M, Bringmann HP, Schlauer J. 2001. Uptake of the amino acid alanine by digestive leaves: proof of carnivory in the tropical liana Triphyophyllum peltatum (Dioncophyllaceae). Carnivorous Plant Newsletter 30: 15–21. Cameron KM, Chase MW, Swensen SM. 1995. Molecular evidence for the relationship of Triphyophyllum (Dioncophyllaceae) and Ancistrocladus (Ancistrocladaceae). American Journal of Botany 82: 117. François G, Bringmann G, Phillipson JD, Aké Assi L, Dochez C, Rübenacker M, Schneider C, Warhurst DC, Kirby GC. 1994. Activity of extracts and naphthylisoquinoline alkaloids from Triphyophyllum peltatum, Ancistrocladus abbreviatus and A. barteri against Plasmodium falciparum in vitro. Phytochemistry 35: 1461–1464. Green S, Green TL, Heslop-Harrison Y. 1979. Seasonal heterophylly and leaf gland features in Triphyophyllum (Dioncophyllaceae), a new carnivorous plant genus. Journal of the Linnean Society, Botany 78: 99–116. Heubl G, Bringman G, Meimberg H. 2006. Molecular phylogeny and character evolution of carnivorous plant families in Caryophyllales — revisited. Plant Biology 8: 821–830. Juniper BE, Robins RJ, Joel DM. 1989. The carnivorous plants. Academic Press, London. McPherson S. Carnivorous plants and their habitats (2

vols.). Redfern Natural History Productions, Poole. McPherson S. 2008. Glistening carnivores. The stickyleaved insect-eating plants. Redfern Natural History Productions, Poole. Meimberg H, Dittrich P, Bringmann C, Schlauer J, Heubl G. 2000. Molecular phylogeny of Caryophyllales s.l. based on matK sequences with special emphasis on carnivorous taxa. Plant Biology 2: 218–228. Metcalfe CR. 1952. The anatomical structure of the Dioncophyllaceae in relation to the taxonomic affinities of the family. Kew Bulletin 1951: 351–368. 324. ANCISTROCLADACEAE KARDAL FAMILY Airy Shaw HK. 1952. On the Dioncophyllaceae, a remarkable new family of flowering plants. Kew Bulletin 1951: 327–347. Bringman G, Pokorny F. 1995. The naphtylisoquinoline alkaloids. Alkaloids 46: 127–271. Heubl GR, Turini F, Mudogo V, Kajahn I, Bringmann G. 2010. Ancistrocladus iliboensis (D.R. Congo), a new liana with unique alkaloids. Plant Ecology and Evolution 143: 63–69. Keng H. 1967. Observations on Ancistrocladus. Gardens’ Bulletin, Singapore 22: 113–121. Meimberg H, Dittrich P, Bringmann C, Schlauer J, Heubl G. 2000. Molecular phylogeny of Caryophyllales s.l. based on matK sequences with special emphasis on carnivorous taxa. Plant Biology 2: 218–228. Van Tieghem P. 1903. Sur les Ancistrocladacées. Journal de Botanique (L. Morot) 17: 151–168. 325. RHABDODENDRACEAE CLUBFRUITTREE FAMILY Fay MF, Cameron KM, Prance GT, Lledó MD, Chase MW. 1997. Familial relationships of Rhabdodendron (Rhabdodendraceae): plastid rbcL sequences indicate a caryophyllid placement. Kew Bulletin 52: 923–932. Nelson B, Prance GT. 1984. Observations on the pollination of Rhabdodendron macrophyllum (Spr. ex Benth.) Huber. Acta Amazonica 14: 411–426. Prance GT. 1968. The systematic position of Rhabdodendron Gilg, Pilg. Bulletin du Jardin botanique de l’État à Bruxelles 38: 127–146. Prance GT. 1972. Rhabdodendraceae. Flora Neotropica Monographs 11: 1–22. Puff C, Weber A. 1976. Contribution to the morphology, anatomy, and caryology of Rhabdodendron, and a reconsideration of the systematic position of the Rhabdodendraceae. Plant Systematics and Evolution 125: 195–222. Record JS. 1934. The woods of Rhabdodendron and Duckeodendron. Tropical Woods 33: 6–10. 326. SIMMONDSIACEAE JOJOBA FAMILY Bailey DC. 1980. Anomalous growth and vegetative anatomy of Simmondsia chinensis. American Journal of Botany 67: 147–161. Carlquist S. 2002. Wood anatomy and successive cambia in Simmondsia (Simmondsiaceae): evidence for inclusion in Caryophyllales s.l. Madroño 49: 158–164. Gentry HS. 1958. The natural history of jojoba (Simmondsia chinensis) and its cultural aspects. Economic Botany 12: 261–295. Rost TL, Schimper D, Schell P, Allen S. 1977. Anatomy of jojoba (Simmondsia chinensis) seed and the utilisation of liquid wax during germination. Economic Botany 31: 140–147. Sherbrooke WC, Haase FW. 1974. Jojoba: a wax-producing shrub of the Sonoran Desert. Arid Lands

Resource Information Paper 5. Office of Arid Lands Studies, University of Sonora, Tuscon. Van Tieghem P. 1898. Sur le genre Simmondsia, considéré comme type d’une famille distincte, les Simmondsiacées. Journal de Botanique (L. Morot) 12: 103–11 327. PHYSENACEAE BALLOONFRUIT FAMILY Capuron R. 1968. Sur le genre Physena Noronh. ex Thouars. Adansonia II, 8: 355–357. Dickison WC, Miller RB. 1993. Morphology and anatomy of the Malagasy genus Physena (Physenaceae), with a discussion of the relationships of the genus. Adansonia séries 3, 15: 85–106. Inoue M, Ohtani K, Kasai R, Okukubo M, Andriansiferana M, Yamasaki K, Koike T. 2009. Cytotoxic 16-ß[(D-xylopyranosyl)oxy]oxohexadecanyl triterpene glycosides from a Malagasy plant, Physena sessiliflora. Phytochemistry 70: 1195–1202. Miller RB, Dickison WC. 1992. Wood anatomy of Asteropeia (Asteropeiaceae) and Physena (Physenaceae): two endemics from Madagascar. American Journal of Botany 79: 41. Morton MC, Karol KG, Chase MW. 1997. Taxonomic affinities of Physena (Physenaceae) and Asteropeia (Theaceae). Botanical Review 63: 231–239. 328. ASTEROPEIACEAE MANOKO FAMILY Miller RB, Dickison WC. 1992. Wood anatomy of Asteropeia (Asteropeiaceae) and Physena (Physenaceae): two endemics from Madagascar. American Journal of Botany 79(6): 41. Morton MC, Karol KG, Chase MW. 1997. Taxonomic affinities of Physena (Physenaceae) and Asteropeia (Theaceae). Botanical Review 63: 231–239. Schatz GE, Lowry II PP, Wolf AE. 1999. Endemic families of Madagascar. IV. A synoptic revision of Asteropeia (Asteropeiaceae). Adansonia séries 3, 21: 255–268. 329. MACARTHURIACEAE MACARTHURIA FAMILY Behnke HD, Mabry TJ, Neumann P, Bathlott W. 1983. Ultrastructural, micromorphological and phytochemical evidence for a “central position” of Macarthuria (Molluginaceae) within the Caryophyllales. Plant Systematics and Evolution 143: 151–161. Brockington SF, Walker RH, Glover B, Soltis PS, Soltis DE. 2011. Complex pigment evolution in the Caryophyllales. New Phytologist 190: 854–864. Christenhusz MJM, Christin P-A, Sage RF. 2014. On the disintegration of Molluginaceae: a new genus and family (Kewa, Kewaceae) segregated from Hypertelis, and the placement of Macarthuria in Macarthuriaceae. Phytotaxa 181: 238–243. Hofmann U. 1973. Centrospermen-Sudien 6. Morphologische Untersuchungen zur Umgrenzung und Gliederung der Aizoaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 93: 247–324. Lepschi BJ. 1996. A taxonomic revision of Macarthuria (Molluginaceae) in Western Australia. Nuytsia 11: 37–54. 330. MICROTEACEAE JUMBY-PEPPER FAMILY Schäferhoff B, Müller KF, Borsch T. 2009. Caryophyllales phylogenetics: disentangling Phytolaccaceae and Molluginaceae and description of Microteaceae as a new isolated family. Willdenowia 39: 209–228.

Plants of the World

729

FURTHER READING 331. CARYOPHYLLACEAE CARNATION FAMILY Brantjes NBM, Leemans JAAM. 1976. Silene otites (Caryophyllaceae) pollinated by nocturnal Lepidoptera and mosquitoes. Acta Botanica Neerlandica 25: 281–295. Charlesworth D. 2008. Plant sex chromosomes. Genome Dynamics 4: 83–94. Desfeux C, Lejeune B. 1996. Systematics of Euromediterranean Silene (Caryophyllaceae): evidence from a phylogenetic analysis using ITS sequences. Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la Vie 319: 351–358. Dillenberger MS, Kadereit JW. 2014. Maximum polyphyly: multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae). Taxon 63: 64–88. Erixon P, Oxelman B. 2008. Reticulate or treelike chloroplast DNA evolution in Silene (Caryophyllaceae)? Molecular Phylogenetics and Evolution 48: 313–325, Fior S, Karis PO, Casazza G, Minuto L, Sala F. 2006. Molecular phylogeny of the Caryophyllaceae (Caryophyllales) inferred from chloroplast matK and nuclear rDNA ITS sequences. American Journal of Botany 93: 399–411. Fior S, Karis PO. 2007. Phylogeny, evolution and systematics of Moehringia (Caryophyllaceae) as inferred from molecular and morphological data: a case for homology reassessment. Cladistics 23: 362–372. Frajman B, Heidari N, Oxelman B. 2009. Phylogenetic relationships of Atocion and Viscaria (Sileneae, Caryophyllaceae) inferred from chloroplast, nuclear ribosomal, and low-copy gene DNA sequences. Taxon 58: 811–824. Greenberg AK, Donoghue MJ. 2011. Molecular systematics and character evolution in Caryophyllaceae. Taxon 60: 1637–1652. Harbaugh DT, Nepokroeff M, Rabeler RK, McNeill J, Zimmer EA, Wagner WL. 2010. A new lineage based tribal classification of the family Caryophyllaceae. International Journal of Plant Sciences 171: 185–198. Jordan GJ, Macphail MK. 2003. A Middle-Late Eocene inflorescence of Caryophyllaceae from Tasmania, Australia. American Journal of Botany 90: 761–768. Mameli E, Aschieri E. 1920. Richerche anatomische e biomiche sul Lychnis viscaria. Atti dell’Istituto Botanico dell’Universita di Pavia, series II 17: 119–129. McNeill J, Bassett IJ. 1974. Pollen morphology and the infrageneric classification of Minuartia (Caryophyllaceae). Canadian Journal of Botany 52: 1225–1231. Valente LM, Savolainen V, Vargas P. 2010. Unparalleled rates of species diversification in Europe. Proceedings of the Royal Society B 277: 1489–1496. Weller SG, Sakai AK, Wagner WL, Herbst DR. 1990. Evolution of dioecy in Schiedea (Caryophyllaceae: Alsinoideae) in the Hawaiian Islands: biogeographical and ecological factors. Systematic Botany 6: 126–135. Yashina S, Gubin S, Maksimovich S, Yashina A, Gakhova E, Gilichikinsky D. 2012. Regeneration of whole fertile plants from 30,000-y-old fruit tissue buried in Siberian permafrost. Proceedings of the National Academy of Aciences (USA) 109: 4008–4013.

730

Christenhusz, Fay & Chase

332. ACHATOCARPACEAE SNAKE-EYES FAMILY Brown GK, Varadarajan GS. 1985. Studies in Caryophyllales I: re-evaluation of classification of Phytolaccaceae s.l. Systematic Botany 10: 49–63. Carlquist S. 2000. Wood and bark anatomy of Achatocarpaceae. Sida 19: 71–78. Skvarla JJ, Nowicke JW. 1982. Pollen fine structure and relationships of Achatocarpus Triana and Phaulothamnus A. Gray. Taxon 31: 244–249. 333. AMARANTHACEAE SPINACH FAMILY Fuentes-Bazan S, Mansion G, Borsch T. 2012. Towards a species level tree of the globally diverse genus Chenopodium (Chenopodiaceae). Molecular Phylogenetics and Evolution 62: 359–374. Fuentes-Bazan S, Uotila P, Borsch T. 2012. A novel phylogeny-based generic classification for Chenopodium sensu lato and a tribal rearrangement of Chenopodioideae (Chenopodiaceae). Willdenowia 42: 5–24. Kadereit G, Borsch T, Weising K, Freitag H. 2003. Phylogeny of Amaranthaceae and Chenopodiaceae and the evolution of C 4 photosynthesis. International Journal of Plant Sciences 164: 959–986. Kadereit G, Freitag H. 2011. Molecular phylogeny of Camphorosmeae (Camphorosmoideae, Chenopodiaceae): implications for biogeography, evolution of C 4 photosynthesis and taxonomy. Taxon 60: 51–78. Kadereit G, Gotzek D, Jacobs S, Freitag H. 2005. Origin and age of Australian Chenopodiaceae. Organisms, Diversity and Evolution 5: 59–80. Kadereit G, Ball P, Beer S, Mucina L, Sokoloff D, Teege P, Yaprack AE, Freitag H. 2007. A taxonomic nightmare comes true: phylogeny and biogeography of glassworts (Salicornia L., Chenopodiaceae). Taxon 56: 1143–1170. Kadereit G, Ackerly D, Pirie MD. 2012. A broader model for C 4 photosynthesis evolution in plants inferred from the goosefoot family (Chenopodiaceae s. str.). Proceedings of the Royal Society B 279: 3304–3311. Masson R, Kadereit G. 2013. Phylogeny of Polycnemoideae (Amaranthaceae): implications for biogeography, character evolution and taxonomy. Taxon 62: 100–111. Müller KF, Borsch T. 2005. Phylogenetics of Amaranthaceae based on matK/trnK sequence data: evidence from parsimony, likelihood and Bayesian analyses. Annals of the Missouri Botanical Garden 92: 66–102. Prideaux GJ, Ayliffe IK, DeSantis LRG, Schubert BW, Murray PF, Gagan MK, Cerling TE. 2009. Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo. Proceedings of the National Academy of Sciences of the U.S.A. 106: 11646–11650. 334. STEGNOSPERMATACEAE CUBANTANGLE FAMILY Bedell HG. 1980. A taxonomic and morphological reevaluation of Stegnospermaceae (Caryophyllales). Systematic Botany 5: 419–431. Carlquist S. 2012. How wood evolves: a new synthesis. Botany 90: 901–940. Hof m a n n U. 1977. Die St el lu ng von Stegnosperma innerhalb der Centrospermen. Centrospermenstudien 9. Berichte der Deutschen Botanichen Gesellschaft 90: 39–52.

335. LIMEACEAE LIZARD-FOOT FAMILY Hassan NMS, Meve U, Liede-Schumann S. 2005. Seed coat morphology of Aizoaceae-Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance. Botanical Journal of the Linnean Society 148: 189–206. Hofmann U. 1973. Centrospermen-Studien 6. Morphologische Untersuchungen zur Umgrenzung und Gliederung der Aizoaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 93: 247–324. Sharma HP. 1963. Studies in the order Centrospermales II. Vascular anatomy of the flower of certain species of Molluginaceae. Journal of the Indian Botanical Society 42: 19–32. 336. LOPHIOCARPACEAE SANDAARBOSSIE FAMILY Adamson RS. 1958. The South African species of Aizoaceae. IV. Mollugo, Pharnaceum, Coelanthum, and Hypertelis. Journal of South African Botany 24: 11–66. Behnke HD. 1974. Elektronenmikroskopische Untersuchungen and Siebröhren-Plastiden und ihre Aussage über die systematische Stellung von Lophiocarpus. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 94: 114–119. Christin PA, Sage TL, Edwards EJ, Ogburn RM, Khoshravesh R, Sage RF. 2011. Complex evolutionary transitions and the significance of C3-C 4 intermediate forms of photosynthesis in Molluginaceae. Evolution 65: 643–660. Hofmann U. 1973. Centrospermen-Studien 6. Morphologische Untersuchungen zur Umgrenzung und Gliederung der Aizoaceen. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 93: 247–324. Narayana HS, Lodha BC. 1972. Embryology of Orygia decumbens Forssk. Phytomorphology 13: 54–59. Thulin M, Moore A, El-Seedi H, Larsson A, Christin PA, Edwards EJ. 2016. Phylogeny and generic delimitation in Molluginaceae, new pigment data in Caryophyllales and the new family Corbichorniaceae. Taxon 65: 775–793. 337. KEWACEAE SURING FAMILY Adamson RS. 1958. The South African species of Aizoaceae. IV. Mollugo, Pharnaceum, Coelanthum, and Hypertelis. Journal of South African Botany 24: 11–66. Brockington SF, Walker RH, Glover B, Soltis PS, Soltis DE. 2011. Complex pigment evolution in the Caryophyllales. New Phytologist 190: 854–864. Christenhusz MJM, Christin P-A, Sage RF. 2014. On the disintegration of Molluginaceae: a new genus and family (Kewa, Kewaceae) segregated from Hypertelis, and the placement of Macarthuria in Macarthuriaceae. Phytotaxa 181: 238–243. Christin PA, Sage TL, Edwards EJ, Ogburn RM, Khoshravesh R, Sage RF. 2011. Complex evolutionary transitions and the significance of C3-C 4 intermediate forms of photosynthesis in Molluginaceae. Evolution 65: 643–660. Ronse Decraene LP. 2013. Reevaluation of the perianth and androecium in Caryophyllales: implications for flower evolution. Plant Systematics and Evolution 299: 1599–1636. 338. BARBEUIACEAE LIANE-BARBEU FAMILY Behnke HD. 1993. Further studies of the sieve-element plastids of the Caryophyllales including Barbeuia, Corrigiola, Lyallia, Microtea, Sarcobatus and

FURTHER READING Telephium. Plant Systematics and Evolution 186: 231–243. Carlquist S. 2000. Wood anatomy, stem anatomy, and cambial activity of Barbeuia (Caryophyllales). IAWA Journal 20: 31–41. 339. GISEKIACEAE STORK’S-HENNA FAMILY Bissinger K, Khoshravesh R, Kotrade JP, Oakley J, Sage TL, Sage RF, Hartmann HE, Kadereit G. 2014. Gisekia (Gisekiaceae): phylogenetic relationships, biogeography, and ecophysiology of a poorly known C4 lineage in the Caryophyllales. American Journal of Botany 101: 499–509. Gilbert MG. 1993. A review of Gisekia (Gisekiaceae). Kew Bulletin 48: 343–356. Hassan NMS, Meve U, Liede-Schumann S. 2005. Seed coat morphology of Aizoaceae-Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance. Botanical Journal of the Linnean Society 148: 189–206. 340. AIZOACEAE DEWPLANT FAMILY Bittrich V. 1990. Systematic studies in Aizoaceae. Mitteilungen aus dem Institut für allgemeine Botanik in Hamburg 23b: 491–507. Carlquist S. 2007. Successive cambia in Aizoaceae: products and process. Botanical Journal of the Linnean Society 153: 141–155. Chesselet P, Mössmer M, Smith GF. 1995. Research priorities in the succulent plant family Mesembryanthemaceae Fenzl. South African Journal of Science 91: 192–209. Hartmann HEK. 1988. Fruit types in Mesembryanthema. Beiträge zur Biologie der Pflanzen 63: 313–349. Hartmann HEK, Niesler IM. 2007. On the evolution of nectaries in Aizoaceae. Bradleya 27: 69–120. Ihlenfeldt HD. 1994. Diversification in an arid world: the Mesembryanthemaceae. Annual Review of Ecology and Systematics 25: 521–546. Kellner A, Ritz CM, Schlittenhardt P, Hellwig FH. 2011. Genetic differentiation of the genus Lithops L. (Ruschioideae, Aizoaceae) reveals a high level of convergent evolution and reflects geographic distribution. Plant Biology 13: 368–380. Klak C. Bruyns PV, Hanácek P. 2013. A phylogenetic hypothesis of the recently diversified Ruschieae (Aizoaceae) in southern Africa. Molecular Phylogenetics and Evolution 69: 1005–1020. Klak C, Bruyns PV, Hedderson TAJ. 2007. A phylogeny and new classification for Mesembryanthemoideae (Aizoaceae). Taxon 56: 737–756. Klak C, Khunou A, Reeves G, Hedderson TAJ. 2003. A phylogenetic hypothesis for the Aizoaceae (Caryophyllales) based on four plastid DNA regions. American Journal of Botany 90: 1433–1445. Klak C, Reeves G, Hedderson TAJ. 2004. Unmatched tempo of evolution in southern African semi-desert iceplants. Nature 427: 63–65. 341. PHYTOLACCACEAE POKEWEED FAMILY Behnke HD, Chang C, Eifert IJ, Mabry TJ. 1974. Betalains and P-type sieve-tube plastids in Petiveria and Agdestis (Phytolaccaceae). Taxon 23: 541–542. Brown GK, Varadarajan GS. 1985. Studies in Caryophyllales I: Re-evaluation of classification of Phytolaccaceae s.l. Systematic Botany 10: 49–63. Carlquist S. 2000. Wood and stem anatomy of phytolaccoid and rivinoid Phytolaccaceae (Caryophyllales): ecology, systematics, nature of successive cambia. Aliso 19: 13–29.

Cevallos-Ferriz SRS, Estrada-Ruiz E, PérezHer ná ndez BR. 20 08. Phy tolaccaceae infructescence from Cerro del Pueblo formation, Upper Cretaceous (late Campanian), Coahuila, Mexico. American Journal of Botany 95: 77–83. Lee J, Kim SY, Park SH, Ali MA. 2013. Molecular phylogenetic relationships among members of the family Phytolaccaceae sensu lato inferred from internal transcribed spacer sequences of nuclear ribosomal DNA. Genetics and Molecular Research 12: 4515–4525. Nowicke JW. 1968. Palynotaxonomic study of the Phytolaccaceae. Annals of the Missouri Botanical Garden. 55: 294–364. Rogers GK. 1985. The genera of Phytolaccaceae in the southeastern United States. Journal of the Arnold Arboretum 66: 1–37. Ronse Decraene LP. 2013. Reevaluation of the perianth and androecium in Caryophyllales: implications for flower evolution. Plant Systematics and Evolution 299: 1599–1636. Zheng HC, Ma SW, Chai TY. 2010. Ovular development and perisperm formation in Phytolacca americana (Phytolaccaceae) and their systematic significance in Caryophyllales. Journal of Systematics and Evolution 48: 318–325. 342. PETIVERIACEAE HENWOOD FAMILY Bissinger K, Khoshravesh R, Kotrade JP, Oakley J, Sage TL, Sage RF, Hartmann HE, Kadereit G. 2014. Gisekia (Gisekiaceae): phylogenetic relationships, biogeography, and ecophysiology of a poorly known C4 lineage in the Caryophyllales. American Journal of Botany 101: 499–509. Brockington SF, Alexandre R, Ramdial J, Moore MJ, Crawley S, Dhingra A, Hilu K, Soltis DE, Soltis PS. 2009. Phylogeny of the Caryophyllales sensu lato: revisiting hypotheses on pollination biology and perianth differentiation in the core Caryophyllales. International Journal of Plant Sciences 170: 627–643. Carlquist S. 2000. Wood and stem anatomy of phytolaccoid and rivinoid Phytolaccaceae (Caryophyllales): ecology, systematics, nature of successive cambia. Aliso 19: 13–29. Jansen S, Ronse Decraene LP, Smets E. 2000. On the wood and stem anatomy of Monococcus echinophorus (Phytolaccaceae s.l.). Systematic Geography of Plants 70: 171–179. Rohwer J. 1982. A taxonomic revision of the genera Seguieria Loefl. and Gallesia Cesar. Mitteilungen der Botanisches Staatssammlung München 18: 231–288. 343. SARCOBATACEAE GREASEWOOD FAMILY Cuénoud P, Savolainen V, Chatrou LW, Powell M, Grayer RJ, Chase MW. 2001. Molecular phylogenetics of Caryophyllales based on nuclear 18S rDNA and plastid rbcL, atpB, and matK DNA sequences. American Journal of Botany 89: 132–144. Drenovsky RE, Richards JH. 2005. Nitrogen addition increases fecundity in the desert shrub Sarcobatus vermiculatus. Oecologia 143: 349–356. Khan MA, Gul B, Weber DJ. 2001. Seed germination in relation to salinity and temperature in Sarcobatus vermiculatus. Biologia Plantarum 45: 133–135. 344. NYCTAGINACEAE FOUR-O’CLOCKS FAMILY Baker HG. 1964. Variation in style length in relation to outbreeding in Mirabilis. Evolution 18: 507–509.

Bogle AL. 1974. The genera of Nyctaginaceae in the southeastern United States. Journal of the Arnold Arboretum 55: 1–37. Burger AE. 2005. Dispersal and germination of seeds of Pisonia grandis, and Indo-Pacific tropical tree associated with insular seabird colonies. Journal of Tropical Ecology 21: 263—271. Carlquist S. 2004. Lateral meristems, successive cambia and their products: a reinterpretation based on roots and stems of Nyctaginaceae. Botanical Journal of the Linnean Society 146: 129–143. Cruden RW. 1973. Reproductive biology of weedy and cultivated Mirabilis. American Journal of Botany 60: 802–809. Douglas NA, Manos PS. 2007. Molecular phylogeny of Nyctaginaceae: taxonomy, biogeography, and characters associated with radiation of xerophytic genera in North America. American Journal of Botany 95: 856–872. Douglas NA, Spellenberg R. 2010. A new tribal classification of Nyctaginaceae. Taxon 59: 905–910. Levin RA. 2000. Phylogenetic relationships within Nyctaginaceae: evidence from nuclear and chloroplast genomes. Systematic Botany 25: 738–750. MacDaniels LH. 1981. A study of cultivars of Bougainvillea. Baileya 21: 77–100. Sattler R, Perlin L. 1982. Floral development of Bougainvillea spectabilis Willd., Boerhavia diffusa L. and Mirabilis jalapa L. (Nyctaginaceae). Botanical Journal of the Linnean Society 84: 161–182. Vanvinckenroye P, Cresens E, Ronse de Craene LP, Smets EF. 1993. A comparative floral development study in Pisonia, Bougainvillea and Mirabilis (Nyctaginaceae) with special emphasis on the gynoecium and floral nectaries. Bulletin du Jardin Botanique National de la Belgique 62: 69–96. 345. MOLLUGINACEAE CARPETWEED FAMILY Adamson RS. 1958. The South African species of Aizoaceae. IV. Mollugo, Pharnaceum, Coelanthum, and Hypertelis. Journal of South African Botany 24: 11–66. Arakaki M, Christin PA, Nyffeler R, Lendel A, Eggli U, Ogburn RM, Spriggs E, Moore MJ, Edwards EJ. 2011. Contemporaneous and recent radiations of the world’s major succulent lineages. Proceedings of the National Academy of Sciences of the USA 108: 8379–8384. Boetsch JR. 2002. The Aizoaceae and Molluginaceae of the southeastern United States. Castanea 67: 42–53. Chase MW, et al. (40 additional authors). 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Annals of the Missouri Botanical Garden 80: 528–580. Christin PA, Sage TL, Edwards EJ, Ogburn RM, Khoshravesh R, Sage RF. 2011. Complex evolutionary transitions and the significance of C3-C 4 intermediate forms of photosynthesis in Molluginaceae. Evolution 65: 643–660. Shar ma HP. 1963. St udies in the order Centrospermales. II. Vascular anatomy of the flower of certain species of the Molluginaceae. Journal of the Indian Botanical Society 42: 19–32. Thulin M, Moore A, El-Seedi H, Larsson A, Christin PA, Edwards EJ. 2016. Phylogeny and generic delimitation in Molluginaceae, new pigment data in Caryophyllales and the new family Corbichorniaceae. Taxon 65: 775–793.

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FURTHER READING 346. MONTIACEAE BLINKS FAMILY Applequist WL, Wagner WL, Zimmer EA, Nepokroeff M. 2006. Molecular evidence resolving the systematic position of Hectorella (Portulacaceae). Systematic Botany 31: 310–319. Carlquist S. 1998. Wood anatomy of Portulacaceae and Hectorellaceae: ecological, habital and systematic implications. Aliso 16: 137–153. Cranwell LM. 1963. The Hectorellaceae: pollen type and taxonomic speculation. Grana Palynologica 4: 195–202. Hershkovitz MA. 1993. Revised circumscriptions and subgeneric taxonomies of Calandrinia and Montiopsis (Portulacaceae) with notes on phylogeny of the portulacaceous alliance. Annals of the Missouri Botanical Garden 80: 333–365. Hershkovitz MA. 2006. Ribosomal and chloroplast DNA evidence for diversification of western American Portulacaceae in the Andean region. Gayana Botanica 63: 13–74. Hershkovitz MA, Zimmer EA. 2000. Ribosomal DNA evidence and disjunctions of western American Portulacaceae. Molecular Phylogenetics and Evolution 15: 419–439. Mathew B. 1989. The genus Lewisia, a Kew Magazine monograph. Royal Botanic Gardens, Kew and Timber Press, Portland. Nilsson Ö. 1967. Studies in Montia L. and Claytonia L. and allied genera. III. Pollen morphology. Grana Palynologica 7: 279–363. Nyffeler R, Eggli U. 2010. Disintegrating Portulacaceae: a new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. O’Quinn R, Hufford L. 2005. Molecular systematics of Montieae (Portulacaceae): implications for taxonomy, biogeography and ecology. Systematic Botany 30: 314–331. 347. DIDIEREACEAE OCTOPUS-TREE FAMILY Appelquist WL, Wallace RS. 2000. Phylogeny of the Madagascan endemic family Didiereaceae. Plant Systematics and Evolution 221: 157–166. Appelquist WL, Wallace RS. 2003. Expanded circumscription of Didiereaceae and its division into three subfamilies. Adansonia Séries 3, 25: 13–16. Erbar C, Leins P. 2006. Floral ontogeny and systematic position of the Didiereaceae. Plant Systematics and Evolution 261: 165–185. Rauh W. 1956. Mor phologische, entwicklungsgeschichtliche, histogenetische und anatomische Untersuchungen and den Sprossen der Didiereaceen. Abhandlungen der Akademie der Wissenschaften und Literatur, MathematischNaturwissenschaftliche Klasse, Jahrgang 1956, 6: 1–104. 348. BASELLACEAE MALABAR-SPINACH FAMILY Bogle AL. 1969. The genera of Portulacaceae and Basellaceae in the southeastern United States. Journal of the Arnold Arboretum 50: 566–598. Carlquist S. 1999. Wood, stem and root anatomy of Basellaceae with relation to habit, systematics, and cambial variants. Flora 194: 1–12. Eriksson R. 2007. A synopsis of Basellaceae. Kew Bulletin 62: 297–320. King SR, Gershoff SN. 1987. Nutritional evaluation of three underexploited Andean tubers: Oxalis tuberosa (Oxalidaceae), Ullucus tuberosus (Basellaceae), and Tropaeolum tuberosum (Tropaeolaceae). Economic Botany 41: 503–511.

732

Christenhusz, Fay & Chase

Nowicke JW. 1996. Pollen morphology, exine structure and the relationships of Basellaceae and Didiereaceae to Portulacaceae. Systematic Botany 21: 187–208. Pietilä L. 1995. Pollination requirements for seed set in ulluco (Ullucus tuberosus). Euphytica 84: 127–131. 349. HALOPHYTACEAE VERDOLAGA FAMILY Hunziker JH, Behnke HD, Eifert IJ, Mabry TJ. 1974. Halophytum ameghinoi: a betalain-containing and P-type sieve-tube plastid species. Taxon 23: 537–539. Pozner R, Cocucci A. 2006. Floral structure, anther development, and pollen dispersal of Halophytum ameghinoi (Halophytaceae). International Journal of Plant Sciences 167: 1091–1098. Soriano A. 1946. Halophytaceae. Nueva familia del orden Centrospermae. Notas del Museo de La Plata 11: 161–175. 350. TALINACEAE FAMEFLOWER FAMILY Nyffeler R, Eggli U. 2009. Disintegrating Portulacaceae: a new familial classification of the suborder Portulacinae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. Von Poellnitz K. 1934. Monographie der Gattung Talinum Adans. Feddes Repertorium 35: 161–173. 351. PORTULACACEAE PURSLANE FAMILY Christin PA, Osborne CP, Sage RF, Arakaki M, Edwards EJ. 2011. C4 eudicots are not younger than C4 monocots. Journal of Experimental Botany 62: 3171–3181. Geesink R. 1969. An account of the genus Portulaca in Indo-Australia and the Pacific (Portulacaceae). Blumea 17: 275–307. Legrand CD. 1962. Las especies americanas de Portulaca. Anales del Museo de Historia Natural de Montevideo ser. 2, 7: 1–147. Matthews JF, Levins PA. 1985. The genus Portulaca in the southeastern United States. Castanea 50: 96–104. Nyffeler R, Eggli U. 2009. Disintegrating Portulacaceae: a new familial classification of the suborder Portulacinae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. Ocampo G. 2013. Morphological characterisation of seeds in Portulacaceae. Phytotaxa 141: 1–24. Ocampo G, Columbus JT. 2012. Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae). Molecular Phylogenetics and Evolution 63: 97–112. Ocampo G, Koteyeva NK, Voznesenskaya EV, Edwards GE, Sage TL, Sage RF, Columbus JT. 2013. Evolution of leaf anatomy and photosynthetic pathways in Portulacaceae. American Journal of Botany 100: 2388–2402. 352. ANACAMPSEROTACEAE LOVE-PLANT FAMILY Gerbaulet M. 1992–1993. Die Gattung Anacampseros L. (Portulacaceae). Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 113: 477–576; 114: 1–28. Nyffeler R. 2007. The closest relatives of cacti: insights from phylogenetic analyses of chloroplast and mitochondrial sequences with special emphasis on relationships in the tribe Anacampseroteae. American Journal of Botany 94: 89–101.

353. CACTACEAE CACTUS FAMILY Anderson EF. 2001. The cactus family. Timber Press, Portland. Arakaki M, Christin PA, Nyffeler R, Lendel A, Eggli U, Ogburn RM, Spriggs E, Moore MJ, Edwards EJ. 2011. Contemporaneous and recent radiations of the world’s major succulent lineages. Proceedings of the National Academy of Sciences of the U.S.A. 108: 8379–8384. Bárcenas RT, Yesson C, Hawkins JA. 2011. Molecular systematics of the Cactaceae. Cladistics 27: 470–489. Benson LD. 1982. The cacti of the United States and Canada. Stanford University Press, Stanford. Calvente A, Zappi DC, Forest F, Lohmann LG. 2011. Molecular phylogeny, evolution, and biogeography of South American epiphytic cacti. International Journal of Plant Sciences 172: 902–914. Edwards EJ. 2006. Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae). New Phytologist 172: 479–489. Edwards EJ, Nyffeler R, Donoghue MJ. 2005. Basal cactus phylogeny: implications of Pereskia paraphyly for the transition to the cactus life form. American Journal of Botany 92: 1177–1188. Gibson AC, Nobel PS. 1986. The cactus primer. Harvard University Press, Cambridge. Hernández-Hernández T, Hernández HM, De-Nova JA, Puente R, Eguiarte LE, Magallón S. 2011. Phylogenetic relationships and evolution of growth form in Cactaceae (Caryophyllales, Eudicotyledoneae). American Journal of Botany 98: 44–61. Hunt DR (comp.). 1999. CITES Cactaceae checklist, ed. 2. Royal Botanic Gardens, Kew. Hunt DR, Taylor NP. 1990. The genera of Cactaceae: progress towards consensus. Bradleya 8: 85–107. Majure LC, Puente R, Griffith MP, Judd WS, Soltis PS, Soltis DE. 2012. Phylogeny of Opuntia s.s. (Cactaceae): clade delineation, geographic origins, and reticulate evolution. American Journal of Botany 99: 847–864. Metzing D, Kiesling R. 2008. The study of cactus evolution: the pre-DNA era. Haseltonia 14: 6–25. Nobel PS. 1988. Environmental biology of agaves and cacti. Cambridge University Press, Cambridge. Nobel PS (ed.). 2002. Cacti: biology and uses. University of California Press, Berkeley. Nyffeler R. 2002. Phylogenetic relationships in the cactus family (Cactaceae) based on evidence from trnK/matK and trnL-trnF sequences. American Journal of Botany 89: 312–326. Nyffeler R, Eggli U. 2010. A farewell to dated ideas and concepts — molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6: 109–149. Ogburn RM, Edwards EJ. 2010. Anatomical variation in Cactaceae and relatives: trait lability and evolutionary innovation. American Journal of Botany 96: 391–408. Schlumpberger BO, Renner SS. 2012. Molecular phylogenetics of Echinopsis (Cactaceae): polyphyly at all levels and convergent evolution of pollination modes and growth forms. American Journal of Botany 99: 1335–1349. Schwager H, Masselter T, Speck T, Neinhuis C. 2013. Functional morphology and biomechanics of branch-stem junctions in columnar cacti. Proceedings of the Royal Society B 280: 1471–2954. Vásquez-Sánchez M, Terrazas T, Arias S, Ochoterena H. 2013. Molecular phylogeny, origin and taxonomic implications of the tribe Cacteae

FURTHER READING (Cactaceae). Systematics and Biodiversity 11: 103–116. Wallace RS. 1995. Molecular systematic study of the Cactaceae: using chloroplast DNA variation to elucidate cactus phylogeny. Bradleya 13: 1–12. 354. NYSSACEAE TUPELO-TREE FAMILY Eyde RH. 1968. The peculiar gynoecial vasculature of Cornaceae and its systematic significance. Phytomorphology 17: 172–182. Eyde RH. 1997. Fossil record and ecology of Nyssa (Cornaceae). Botanical Review 63: 97–123. Eyde RH, Xiang QY. 1990. Fossil mastixioid (Cornaceae) alive in eastern Asia. American Journal of Botany 77: 689–692. Hook DD, Brown CL, Kormanik PP. 1970. Lenticel and water root development of swam tupelo under various flooding conditions. Botanical Gazette 131: 217–224. Horne AS. 1909. The structure and affinities of Davidia involucrata Baill. Transactions of the Linnean Society of London II, 7: 303–326. Li S, Zhang Z, Cain A, Wang B, Long M, Taylor J. 2005. Antifungal activity of camptothecin, trifolin, and hyperoside isolated from Camptotheca acuminata. Journal of Agricultural and Food Chemistry 53: 32–37. Lorence A, Nessler CL. 2004. Camptothecin, over four decades of surprising findings. Phytochemistry 65: 2735–2749. Mai D. 1993. On the extinct Mastixiaceae (Cornales) in Europe. Geophytology 23: 53–63. Manchester SR. 2002. Leaves and fruits of Davidia (Cornales) from the Paleocene of North America. Systematic Botany 27: 368–382. Manchester SR, Crane PR, Golovnea LB. 1999. An extinct genus with affinities to extant Davidia and Camptotheca (Cornales) from the Paleocene of North America and eastern Asia. International Journal of Plant Sciences 160: 188–207. Manchester SR, Chen ZD, Lu AM, Uemura K. 2009. Eastern Asian endemic seed plant genera and their paleogeographic history throughout the Northern Hemisphere. Journal of Systematics and Evolution 47: 1–41. Matthew KM. 1976. A revision of the genus Mastixia (Cornaceae). Blumea 23: 51–93. Sirikantaramas S, Yamazaki M, Saito K. 2009. A survival strategy: the coevolution of the camptothecin biosynthethic pathway and self-resistance mechanism. Phytochemistry 70: 1894–1898. Vekemans D, Viane T, Caris P, Geuten K. 2011. Transference of function shapes f loral organ identity in the dove tree inf lorescence. New Phytologist 193: 216–228. Wen J, Stuessy TF. 1993. The phylogeny and biogeography of Nyssa (Cornaceae). Systematic Botany 18: 68–79. Xiang QY, Thomas DT, Xiang QP. 2011. Resolving and dating the phylogeny of Cornales — effects of sampling, data partitions and fossil calibrations. Molecular Phylogenetics and Evolution 50: 123–138. 355. HYDROSTACHYACEAE WATERSPIKE FAMILY Albach DC, Soltis DE, Chase MW, Soltis PS. 2001. Phylogenetic placement of the enigmatic angiosperm Hydrostachys. Taxon 50: 781–805. Cusset C. 1973. Révision des Hydrostachyaceae. Adansonia II, 13: 75–119. Hempel AL, Reeves PA, Olmstead RG, Jansen

RK. 1995. Implications of rbcL sequence data for higher order relationships of the Loasaceae and the anomalous aquatic plant Hydrostachys (Hydrostachyaceae). Plant Systematics and Evolution 194: 25–37. Xiang QY. 1999. Systematic affinities of Grubbiaceae and Hydrostachyaceae within Cornales — insights from rbcL sequences. Harvard Papers in Botany 4: 527–542. 356. HYDRANGEACEAE HORTENSIA FAMILY Fairbairn JW, Lou TC. 1950. A pharmacognostical study of Dichroa febrifuga Lour., a Chinese antimalarial plant. Journal of Pharmacy and Pharmacology 2: 162–177. Hao G, Hu C. 1996. A study of pollen morphology of Hydrangeoideae (Hydrangeaceae). Journal of Tropical and Subtropical Botany 4: 26–31. He P. 1990. Taxonomy of Deutzia (Hydrangeaceae) from Sichuan, China. Phytologia 69: 332–339. Hu SY. 1954, 1955, 1956. A monograph of the genus Philadelphus. Journal of the Arnold Arboretum 35: 275–333; 36: 52–109; 37: 15–90. Hufford L. 1995. Seed morphology of Hydrangeaceae and its phylogenetic implications. International Journal of Plant Sciences 156: 555–580. Hufford L. 1997. A phylogenetic analysis of Hydrangeaceae using morphological data. International Journal of Plant Sciences 158: 652–672. Hufford L, Moody ML, Soltis DE. 2001. A phylogenetic analysis of Hydrangeaceae based on sequences of the plastid gene matK and their combination with rbcL and morphological data. International Journal of Plant Sciences 162: 835–846. McClintock E. 1956. The cultivated hydrangeas. Baileya 4: 165–175. Mendoza CG, Wanke S, Salomo K, Goetghebeur P, Samain MS. 2013. Application of the phylogenetic informativeness method to chloroplast markers: a test case of closely related species in tribe Hydrangeeae (Hydrangeaceae). Molecular Phylogenetics and Evolution 66: 233–242. Nevling LI. 1964. Climbing hydrangeas and their relatives. Arnoldia 24: 17–39. Rinehart TA, Scheffler BE, Reed SM. 2006. Genetic diversity estimates for the genus Hydrangea and development of a molecular key based on SSR. Journal of the American Society of Horticultural Science 131: 787–797. Roels P, Ronse Decraene LP, Smets EF. 1997. A floral ontogenetic investigation of the Hydrangeaceae. Nordic Journal of Botany 17: 235–254. Samain MS, Wanke S, Goetghebeur P. 2010. Unraveling extensive paraphyly in the genus Hydrangea s.l. with implications for the systematics of tribe Hydrangeeae. Systematic Botany 35: 593–600. Stern WL. 1978. Comparative anatomy and systematics of woody Saxifragaceae. Hydrangea. Botanical Journal of the Linnean Society 76: 83–113. Turner BL. 2001. Taxonomic revision of the genus Fendlera (Hydrangeaceae). Lundellia 4: 1–11. Wyman D. 1965. The mock-oranges. Arnoldia 25: 29–36. Zaïkonnikova TI. 1966. Deutzias — ornamental shrubs — a monograph of the genus Deutzia Thunberg (Saxifragaceae). Nauka, Moscow. 357. LOASACEAE BLAZINGSTAR FAMILY Ackermann M, Weigend M. 2006. Nectar, floral morphology and pollination syndrome in Loasaceae subfamily Loasoideae (Cornales). Annals of Botany 98: 503–514.

Carlquist S. 1984. Wood anatomy of Loasaceae with relation to systematics, habit and ecology. Aliso 10: 538–602. Darlington J. 1934. A monograph of Mentzelia. Annals of the Missouri Botanical Garden 21: 103–226. Hempel AL, Reeves PA, Olmstead RG, Jansen RK. 1995. Implications of rbcL sequence data for higher order relationships of the Loasaceae and the anomalous aquatic plant Hydrostachys (Hydrostachyaceae). Plant Systematics and Evolution 194: 25–37. Hufford L, McMahon MM, Sherwood AM, Reeves G, Chase MW. 2003. The major clades of Loasaceae: phylogenetic analysis using the plastid matK and trnL-trnF regions. American Journal of Botany 90: 1215–1228. Hufford L, McMahon MM, O’Quinn R, Poston ME. 2005. A phylogenetic analysis of Loasaceae subfamily Loasoideae based on plastid DNA sequences. International Journal of Plant Sciences 166: 289–300. Moody ML, Hufford L. 2000. Floral ontogeny and morphology of Cevallia, Fuentesia, and Gronovia (Loasaceae subfamily Gronovoideae). International Journal of Plant Sciences 161: 869–883. Moody ML, Hufford L, Soltis DE, Soltis PS. 2001. Phylogenetic relationships of Loasaceae subfamily Gronovioideae inferred from matK and ITS sequence data. American Journal of Botany 88: 326–336. Müller AA. Kufer JK, Dietl KG, Weigend M. 1999. Iridoid glucosides — chemotaxonomic markers in Loasoideae. Phytochemistry 52: 67–78. Weigend M. 1997. Nasa and the conquest of South America. Published by the author, Munich. Weigend M, Aitzetmüller K, Bruehl L. 2004. The seeds of Loasaceae subfam. Loasoideae (Cornales) I: seed release, seed numbers and fatty acid composition. Flora 199: 424–436. Weigend M, Gottschling M. 2006. Evolution of funnel-revolver flowers and ornithophily in Nasa (Loasaceae). Plant Biology 8: 120–142. Weigend M, Gottschling M, Hoot S, Ackermann M. 2004. A preliminary phylogeny of Loasaceae subfam. Loasoideae (Angiospermae: Cornales) based on trn(UAA) sequence data, with consequences for systematics and historical biogeography. Organisms, Diversity and Evolution 4: 73–90. Xiang QY. 1999. Systematic affinities of Grubbiaceae and Hydrostachyaceae within Cornales — insights from rbcL sequences. Harvard Papers in Botany 4: 527–542. 358. CURTISIACEAE ASSEGAI FAMILY Eyde RH. 1968. The peculiar gynoecial vasculature of Cornaceae and its systematic significance. Phytomorphology 17: 172–182. Manchester SR, Xiang QY, Ziang QP. 2007. Curtisia (Cornales) from the Eocene of Europe and its phytogeographical significance. Botanical Journal of the Linnean Society 155: 127–134. Marais W. 1985. Curtisia. Kew Magazine 2: 368. Yembaturova EY, Van Wyk BE, Tilney PM. 2009. A review of the genus Curtisia (Curtisiaceae). Bothalia 39: 87–96. Xiang QY, Moody ML, Soltis DE, Fan CZ, Soltis PS. 2002. Relationships within Cornales and circumscription of Cornaceae — matK and rbcL sequence data and effects of outgroups and long branches. Molecular Phylogenetics and Evolution 24: 35–57.

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FURTHER READING 359. GRUBBIACEAE KOOLHOUT FAMILY Carlquist S. 1977. A revision of Grubbiaceae. Journal of South African Botany 43: 115–128. Carlquist S. 1977. Wood anatomy of Grubbiaceae. Journal of South African Botany 43: 129–144. Carlquist S. 1978. Vegetative anatomy and systematics of Grubbiaceae. Botaniska Notiser 131: 117–126. Dahlgren R, Van Wijk AE. 1988. Structures and relationships of families endemic to or centered in southern Africa. Annals of the Missouri Botanical Garden 25: 1–94. Xiang QY. 1999. Systematic affinities of Grubbiaceae and Hydrostachyaceae within Cornales — insights from rbcL sequences. Harvard Papers in Botany 4: 527–542. Xiang QY, Moody ML, Soltis DE, Fan CZ, Soltis PS. 2002. Relationships within Cornales and circumscription of Cornaceae — matK and rbcL sequence data and effects of outgroups and long branches. Molecular Phylogenetics and Evolution 24: 35–57. 360. CORNACEAE DOGWOOD FAMILY Bloembergen S. 1939. A revision of the genus Alangium. Bulletin du Jardin Botanique de Buitenzorg III, 6: 139–235. Cappiello P, Shadow D. 2005. Dogwoods, the genus Cornus. Timber Press, Portland. Eyde RH. 1968. The peculiar gynoecial vasculature of Cornaceae and its systematic significance. Phytomorphology 17: 172–182. Eyde RH. 1968. Flowers, fruits, and phylogeny of Alangiaceae. Journal of the Arnold Arboretum 49: 167–192. Eyde RH, Bartlett A, Barghoorn ES. 1969. Fossil record of Alangium. Bulletin of the Torrey Botanical Club 96: 288–314. Fan C, Xiang QY. 2001. Phylogenetic relationships within Cornus (Cornaceae) based on 26S rDNA sequences. American Journal of Botany 88: 1131–1138. Feng CM, Manchester SR, Xiang QY. 2009. Phylogeny and biogeography of Alangiaceae (Cornales) inferred from DNA sequences, morphology, and fossils. Molecular Phylogenetics and Evolution 51: 201–214. Manchester SR, Ziang XP, Ziang QY. 2010. Fruits of cornelian cherries (Cornaceae: Cornus subg. Cornus) in the Paleocene and Eocene of the Northern Hemisphere. International Journal of Plant Sciences 171: 882–891. Murrell ZE. 1993. Phylogenetic relationships in Cornus (Cornaceae). Systematic Botany 18: 469–495. Noshiro S, Baas P. 1998. Systematic wood anatomy of Cornaceae and allies. IAWA Journal 19: 43–97. Xiang QY, Moody ML, Soltis DE, Fan CZ, Soltis PS. 2002. Relationships within Cornales and circumscription of Cornaceae — matK and rbcL sequence data and effects of outgroups and long branches. Molecular Phylogenetics and Evolution 24: 35–57. Xiang QY, Thomas DT. 2008. Tracking character evolution and biogeographic history through time in Cornaceae — does choice of methods matter? Journal of Systematics and Evolution 46: 349–374. Xiang QY, Thomas DT, Xiang QP. 2011. Resolving and dating the phylogeny of Cornales — effects of sampling, data partitions and fossil calibrations. Molecular Phylogenetics and Evolution 50: 123–138. 361. BALSAMINACEAE BALSAM FAMILY Akiyama S, Wakabayashi M, Ohba H. 1992. Chromosome evolution in Himalayan Impatiens

734

Christenhusz, Fay & Chase

(Balsaminaceae). Botanical Journal of the Linnean Society 109: 247–257. Grey-Wilson C. 1980. Impatiens of Africa. Balkema, Rotterdam. Grey-Wilson C. 1980. Hydrocera triflora, its floral morphology and relationship with Impatiens. Kew Bulletin 35: 213–219. Janssens SB, Geuten K, Yuan YM, Song Y, Küpfer P, Smets EF. 2006. Phylogenetics of Impatiens and Hydrocera (Balsaminaceae) using chloroplast atpB-rbcL spacer sequences. Systematic Botany 31: 171–180. Janssens SB, Knox EB, Huysmans D, Smets EF, Merckx VSFT. 2009. Rapid radiation of Impatiens (Balsaminaceae) during Pliocene and Pleistocene: results of a global climate change. Molecular Phylogenetics and Evolution 52: 806–824. Janssens SB, Smets EF, Vrijdaghs A. 2012. Floral development of Hydrocera and Impatiens reveal evolutionary trends in the most early diverged lineages of Balsaminaceae. Annals of Botany 109: 1285–1296. Ku r t to A. 1996. Impatiens glandulifera (Balsaminaceae) as an ornamental and escape in Finland, with notes on the other Nordic countries. Symbolae Botanicae Upsalienses 31: 221–228. Lens F, Eeckhout S, Zwartjes R, Smets E, Janssens SB. 2012. The multiple fuzzy origins of woodiness within Balsaminaceae using an intergrated approach. Where do we draw the line? Annals of Botany 109: 783–799. Yuan YM, Song Y, Geuten K, Rahelivololona E, Wohlhauser S, Fischer E, Smets E, Küpfer P. 2004. Phylogeny and biogeography of Balsaminaceae inferred from ITS sequences. Taxon 53: 391–403. 362. MARCGRAVIACEAE SHINGLE-VINE FAMILY De Roon AC, Dressler S. 1997. New taxa of Norantea Aubl. s.l. (Marcgraviaceae) from Central America and adjacent South America. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 119: 327–335. Dressler S. 1997. Marcgravia umbellata (Marcgraviaceae). Curtis’s Botanical Magazine ser. 6, 14: 130–136. Dressler S, Tschapka M. 2002. Bird versus bat pollination in the genus Marcgravia and the description of a new species. Curtis’s Botanical Magazine ser. 6, 19: 104–114. Lens F, Dressler S, Vinckler S, Janssens S, Dessein S, Van Evelghem L, Smets EF. 2005. Palynological variation in balsaminoid Ericales: I. Marcgraviaceae. Annals of Botany 1047–1060. Punt W. 1971. Pollen morphology of the genera Norantea, Soubourea, and Marcgravia (Marcgraviaceae). Pollen and Spores 13: 199–232. Simon R, Holderied MW, Koch CU, Von Helversen O. 2011. Floral acoustics: conspicuous echoes of a disc-shaped leaf attract bat pollinators. Science 333: 631–633. Tschapka M, Dressler S, Von Helversen O. 2006. Bat visits to Marcgravia pittieri and notes on the inflorescence diversity within the genus Marcgravia (Marcgraviaceae). Flora 201: 383–388. Ward NM, Price RA. 2002. Phylogenetic relationships of Marcgraviaceae: insights from three chloroplast genes. Systematic Botany 27: 149–160. Zotz G. 2013. The systematic distribution of vascular epiphytes — a critical update. Botanical Journal of the Linnean Society 171: 453–481.

363. TETRAMERISTACEAE PUNAH FAMILY Collins JP, Berkelhamer RC, Melser M. 1977. Notes on the natural history of the mangrove Pelliciera rhizophorae Tr., Pl. (Theaceae). Brenesia 10/11: 17–29. Graham A. 1977. New records of Pelliciera (Theaceae/ Pellicieraceae) in the Tertiary of the Caribbean. Biotropica 9: 48–52. Janssens SB, Lens F, Dressler S, Geuten K, Smets EF, Vinckler S. 2005. Palynological variation in balsaminoid Ericales. II. Balsaminaceae, Tetrameristaceae, Pellicieraceae and general conclusions. Annals of Botany 96: 1061–1073. Lens F, Dressler S, Janssens S, Van Evelghem L, Smets EF. 2005. Relationships with balsaminoid Ericales: a wood anatomical approach. American Journal of Botany 92: 941–953. Martínez-Millán M. 2010. Fossil record and age of the Asteridae. Botanical Review 76: 83–135. Tomlinson PB. 1986. The botany of mangroves. Cambridge University Press, Cambridge. 364. FOUQUIERIACEAE OCOTILLO FAMILY Dahlgren R, Jensen SR, Nielsen BJ. 1976. Iridoid compounds in Fouquieriaceae and notes on its possible affinities. Botaniska Notiser 129: 207–212. Henrickson J. 1969. Anatomy of periderm and cortex of Fouquieriaceae. Aliso 7: 97–26. Henrickson J. 1972. A taxonomic revision of the Fouquieriaceae. Aliso 7: 439–537. Humphrey RR. 1935. A study of Idria columnaris and Fouquieria splendens. American Journal of Botany 22: 95–99. Johansen DA. 1936. Morphology and embryology of Fouquieria. American Journal of Botany 23: 95–99. Schönenberger J. 2009. Comparative floral structure and systematics of Fouquieriaceae and Polemoniaceae (Ericales). International Journal of Plant Sciences 170: 1132–1167. Schultheis L, Baldwin BG. 1999. Molecular phylogenetics of Fouquieriaceae: evidence from nuclear rDNA ITS studies. American Journal of Botany 86: 578–589. 365. POLEMONIACEAE JACOB’S-LADDER FAMILY Arisawa M, Pezzuto J, Kinghorn A, Douglas A, Cordell G, Farnsworth N. 1984. Plant anticancer agents. 30. Cucurbitacins from Ipomopsis aggregata (Polemoniaceae). Journal of Pharmaceutical Science 73: 411–413. Carlquist S, Eckhart V, Michener D. 1984. Wood anatomy of Polemoniaceae. Aliso 10: 547–572. Day A, Moran R. 1986. Acanthogilia, a new genus of Polemoniaceae from Baja California, Mexico. Proceedings of the Californian Academy of Sciences 44: 111–126. Grant V. 1959. Natural history of the phlox family. Nijhoff, The Hague. Grant V. 1998. Primary classification and phylogeny of the Polemoniaceae, with comments on molecular cladistics. American Journal of Botany 85: 741–752. Grant V. 2001. A guide to understanding recent classifications of the family Polemoniaceae. Lundellia 4: 12–24. Grant V, Grant K. 1965. Flower pollination in the phlox family. Columbia University Press, New York. Johnson LA, Schutz JL, Soltis DE, Soltis PS. 1996. Monophyly and generic relationships of Polemoniaceae based on matK sequences.

FURTHER READING American Journal of Botany 83: 1207–1224. Johnson LA, Chan LM, Weese TL, Busby LD, McMurry S. 2008. Nuclear and cpDNA sequences combined privide string inference of higher phylogenetic relationships in the phlox family (Polemoniaceae). Molecular Phylogenetics and Evolution 48: 997–1012. Lott TA, Manchester SR, Dilcher DL. 1998. A unique and complete polemoniaceous plant from the middle Eocene of Utah, USA. Review of Palaeobotany and Palynology 104: 39–49. Monfils AK, Prather LA. 2004. The conserved nature and taxonomic utility of pollen morphology in Cantua (Polemoniaceae). Grana 43: 249–256. Plitmann U, Levin D. 1990. Breeding systems in the Polemoniaceae. Plant Systematics and Evolution 170: 205–214. Porter JM. 1997. Phylogeny of Polemoniaceae based on nuclear ribosomal internal transcribed spacer DNA sequences. Aliso 15: 57–77. Porter JM, Johnson LA. 1998. Phylogenetic relationships of Polemoniaceae: inferences from mitochondrial nad1b intron sequences. Aliso 17: 157–188. Porter JM, Johnson LA. 2000. A phylogenetic classification of Polemoniaceae. Aliso 19: 55–91. Porter JM, Johnson LA, Wilken D. 2010. Phylogenetic systematics of Ipomopsis (Polemoniaceae): relationships and divergence times estimated from chloroplast and nuclear DNA sequences. Systematic Botany 35: 181–200. Schönenberger J. 2009. Comparative floral structure and systematics of Fouquieriaceae and Polemoniaceae (Ericales). International Journal of Plant Sciences 170: 1132–1167. 366. LECYTHIDACEAE CANNONBALL-TREE FAMILY Appel O. 1996. Morphology and systematics of the Scytopetalaceae. Botanical Journal of the Linnean Society 121: 207–227. Berry EW. 1924. A fossil flower from the Miocene of Trinidad. American Journal of Science 7: 103–108. Carlquist S. 1988. Wood anatomy of Scytopetalaceae. Aliso 12: 63–76. Huang YY, Mori SA, Kelly LM. 2011. A morphological cladistic analysis of Lecythidoideae with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis. Brittonia 63: 396–417. Mori SA, Prance GT. 1987. A guide to collecting Lecythidaceae. Annals of the Missouri Botanical Garden 74: 321–330. Mori SA, Prance GT. 1990. Taxonomy, ecology and economic botany of the Brazil nut (Bertholletia excelsa Humb., Bonpl.: Lecythidaceae). Advances in Economic Botany 8: 130–150. Mori SA, Tsou CH, Wu CC, Cronholm B, Anderberg AA. 2007. Evolution of Lecythidaceae with an emphasis on the circumscription of Neotropical genera: information from combined ndhF and trnL-F sequence data. American Journal of Botany 94: 289–301. Mori SA, Smith NP, Cornejo X, Prance GT. 2010 onwards. The Lecythidaceae pages. http:// sweetgum.nybg.org/lp/index.php. New York Botanical Garden, New York. Morton CM, Mori SA, Prance GT, Karol KG, Chase MW. 1997. Phylogenetic relationships of Lecythidaceae: a cladistic analysis using rbcL sequence and morphological data. American Journal of Botany 84: 530–540. Morton CM, Prance GT, Mori SA, Thorburn LG.

1998. Recircumscription of Lecythidaceae. Taxon 47: 817–827. Prance GT. 2008. A revision of Foetidia (Lecythidaceae subfamily Foetidioideae). Brittonia 60: 336–348. Tsou CH. 1994. The embryology, reproductive morphology and systematics of Lecythidaceae. Memoires of the New York Botanical Garden 71: 1–112. 367. SLADENIACEAE RIBFRUIT FAMILY Anderberg AA, Rydin C, Källersjö M. 2002. Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. American Journal of Botany 89: 677–687. Giraud B, Bussert R, Schrank E. 1992. A new theacean wood from the Cretaceous of northern Sudan. Review of Palaeobotany and Palynology 75: 289–299. Liang D, Baas P. 1991. The wood anatomy of the Theaceae. IAWA Bulletin II, 12: 333–353. Shui YM, Zhang GJ, Zhou ZK, Mo MZ. 2002. Sladenia integrifolia (Sladeniaceae), a new species from China. Novon 12: 539–542. 368. PENTAPHYLACACEAE SAINTEDWOOD FAMILY Carlquist S. 1984. Wood anatomy and relationships of Pentaphylacaceae: significance of vessel features. Phytomorphology 34: 84–90. Liang D, Baas P. 1991. The wood anatomy of the Theaceae. IAWA Bulletin II, 12: 333–353. Mai DH. 1971. Über fossile Lauraceae und Theaceae in Mitteleuropa. Feddes Repertorium 82: 313–341. Prince LM, Parks CM. 2001. Phylogenetic relationships of Theaceae inferred from chloroplast DNA data. American Journal of Botany 88: 2309–2320. Su Y, Liao W, Wang T, Sun Y, Wei Q, Chang H. 2011. Phylogeny and evolutionary divergence times in Apterosperma and Eurydendron: evidence of a Tertiary origin in southern China. Biochemical Systematics and Ecology 39: 769–777. Zhang RJ, Ma HY, Wang YH. 2007. Early floral development of endangered Eurydendron excelsum (Termstroemioideae: Theaceae). Acta Botanica Yunnanica 29: 648–654. 369. SAPOTACEAE SAPODILLA FAMILY Govarts R, Frodin DG, Pennington TD. 2001. World check-list and bibliography of Sapotaceae. Royal Botanic Gardens, Kew. Hall JB. 1996. Vitellaria paradoxa: a monograph. University of Wales, Bangor. Harley MM. 1990. The pollen morphology of Sapotaceae. Kew Bulletin 46: 379–491. Herhey DR. 2004. The widespread misconception that the tambalacoque or calvaria tree absolutely required the dodo bird for its seeds to germinate. Plant Science Bulletin 50: 105–108. Morse K. 2004. Ardent for argan. Saudi Aramco World 55(5): 12–15. Pennington TD. 1991. The genera of Sapotaceae. Royal Botanic Gardens, Kew. Swenson U, Anderberg AA. 2005. Phylogeny, character evolution, and classification of Sapotaceae (Ericales). Cladistics 21: 101–130. Swenson U, Bartisch IV, Munzinger J. 2007. Phylogeny, diagnostic characters and generic limitation of Australasian Chrysophylloideae (Sapotaceae, Ericales): evidence from ITS sequence data and morphology. Cladistics 23: 201–228.

Swenson U, Richardson JE, Bartisch IV. 2008. Multigene phylogeny of the pantropical subfamily Chrysophylloideae (Sapotaceae): evidence of generic polyphyly and extensive morphological homoplasy. Cladistics 24: 1006–1031. Swenson U, Nylinder S, Munzinger J. 2013. Towards a natural classification of Sapotaceae subfamily Chrysophylloideae in Oceania and Southeast Asia based on nuclear sequence data. Taxon 62: 746–770. 370. EBENACEAE PERSIMMON FAMILY Anderberg AA, Rydin C, Källersjö M. 2002. Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. American Journal of Botany 89: 677–687. Berry PE. 1999. A synopsis of the family Lissocarpaceae. Brittonia 51: 214–216. Berry PE, Savolainen V, Sytsma KJ, Hall JC, Chase MW. 2001. Lissocarpa is sister to Diospyros (Ebenaceae). Kew Bulletin 56: 725–729. Contreras LS, Lersten NR. 1984. Extrafloral nectaries in Ebenaceae: anatomy, morphology, and distribution. American Journal of Botany 71: 865–872. Duangjai S, Wallnöfer B, Samuel R, Munzinger J, Chase MW. 2006. Generic delimitation and relationships in Ebenaceae sensu lato: evidence from six plastid DNA regions. American Journal of Botany 93: 1808–1827. Duangjai S, Samuel R, Munzinger J, Forest F, Wallnöfer B, Barfuss MHJ, Fischer G, Chase MW. 2009. A multi-locus plastid phylogenetic analysis of the pantropical genus Diospyrus (Ebenaceae), with an emphasis on the radiation and biogeographic origins on the New Caledonian endemic species. Molecular Phylogenetics and Evolution 52: 602–620. Hiern WP. 1873. A monograph of the Ebenaceae. Transactions of the Cambridge Philosophical Society 12: 27–300. Hillis WE Soenardi P. 1994. Formation of ebony and streaked woods. IAWA Journal 15: 425–437. Hunter JR. 1981. Tendu (Diospyros melanoxylon) leaves, bidi cigarettes, and resource management. Economic Botany 35: 450–459. Morton CM, Chase MW, Kron KA, Swensen SM. 1997. A molecular evaluation of the monophyly of the order Ebenales based upon rbcL sequence data. Systematic Botany 21: 567–586. Wallnöfer B. 2001. The biology and systematics of Ebenaceae: a review. Annalen des Naturhistorischen Museums in Wien B, 103: 485–512. 371. PRIMULACEAE PRIMROSE FAMILY Anderberg AA, Ståhl B. 1995. Phylogenetic interrelationships in the order Primulales, with special emphasis on the family circumscriptions. Canadian Journal of Botany 73: 1699–1730. Anderberg AA, Ståhl B, Källersjö M. 1998. Phylogenetic relationships in the Primulales inferred from rbcL sequence data. Plant Systematics and Evolution 211: 93–102. Anderberg AA, Zhang X, Källersjö M. 2000. Maesaceae, a new primuloid family in the order Ericales s.l. Taxon 49: 183–187. Anderberg AA, Manns U, Källersjö M. 2007. Phylogeny and f loral evolution of the Lysimachieae (Ericales, Myrsinaceae): evidence from ndhF sequence data. Willdenowia 37: 407–421.

Plants of the World

735

FURTHER READING Bone RE, Strijk JS, Fritsch PW, Buerki S, Strasberg D, Thébaud C, Hodkinson TR. 2012. Phylogenetic inference of Badula Juss. (Primulaceae), a rare and threatened genus endemic to the Mascarene Archipelago. Botanical Journal of the Linnean Society 169: 284–296. Carlquist S. 1992. Wood anatomy of sympetalous dicotyledon families: a summary with comments on systematic relationships and evolution of the woody habit. Annals of the Missouri Botanical Garden 79: 303–332. Grey-Wilson C. 1989. The genus Dionysia. Alpine Garden Society, Woking. Hao G, Yuan YM, Hu CM, Ge XJ, Zhao NX. 2004. Molecular phylogeny of Lysimachia (Myrsinaceae) based on chloroplast trnL-F and nuclear ribosomal ITS sequences. Molecular Phylogenetics and Evolution 31: 323–339. Källersjö M, Bergqvist G, Anderberg AA. 2000. Generic realignment in primuloid families of the Ericales s.l.: a phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology. American Journal of Botany 87: 1325–1341. Knudsen JT, Ståhl B. 1994. Floral odours in the Theophrastaceae. Biochemical Systematics and Ecology 22: 259–268. Larsen K, Hu CM. 1995. Reduction of Tetradisia to Ardisia. Nordic Journal of Botany 15: 161–162. Lens F, Jansen S, Caris P, Serlet L, Smets E. 2005. Comparative anatomy of the primuloid clade (Ericales s.l.). Systematic Botany 30: 163–183. Martins L, Oberprieler C, Hellwig FH. 2003. A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data. Plant Systematics and Evolution 237: 75–85. Mast AR, Kelso S, Richards AJ, Land DJ, Feller DMS, Conti E. 2001. Phylogenetic relationships in Primula L. and related genera (Primulaceae) based on noncoding chloroplast DNA. International Journal of Plant Sciences 162: 1381–1400. Mathew B. 2012. Genus Cyclamen: science, cultivation, art and culture. Royal Botanic Gardens, Kew. Pipoly JJ, Ricketson JM. 1999. Discovery of the IndoMalesian genus Hymenandra (Myrsinaceae) in the Neotropics, and its boreotropical implications. Sida 18: 701–746. Punt W, De Leeuw van Weenen JS, Van Oostrum AP. 1974. Primulaceae. The Northwest European pollen flora, 3. Review of Palaeobotany and Palynology 17: 31–70. Richards AJ. 2002. Primula 2nd ed. Timber Press, Portland. Ståhl B. 1995. A revision of Clavija (Theophrastaceae). Opera Botanica 107: 1–77. Strijk JS, Bone RE, Thébaud C, Buerki S, Fritsch PW, Hodkinson TR, Strasberg D. 2014. Timing and tempo of evolutionary diversification in a biodiversity hotspot: Primulaceae on Indian Ocean islands. Journal of Biogeography 41: 810–822. Thrift I, Källersjö M, Anderberg AA. 2002. The monophyly of Primula (Primulaceae) evaluated by analysis of sequences from the chloroplast genes rbcL. Systematic Botany 27: 396–407. Wanntorp L, Ronse Decraene LP, Peng C-I, Anderberg AA. 2011. Floral ontogeny and morphology of Stimpsonia and Ardisiandra, two aberrant genera of the primuloid clade of Ericales. International Journal of Plant Science. 173: 1023–1035. Yesson C, Toomey NH, Culham A. 2009. Cyclamen: time, sea and speciation biogeography using

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a temporally calibrated phylogeny. Journal of Biogeography 36: 1234–1252. 372. THEACEAE TEA FAMILY Bloembergen S. 1952. A critical study in the complex-polymorphous genus Schima (Theaceae). Reinwardtia 2: 133–183. Bozeman JR, Bozeman GA. 1986. This very curious tree. Tipularia 1: 9–15. Cabrera C, Artacho R, Giménez R. 2006. Beneficial effects of green tea — a review. Journal of the American College of Nutrition 25: 79–99. Chang HT, Bartholomew B. 1984. Camellias. Timber Press, Portland. Grote PJ, Dilcher DL. 1989. Investigations of angiosperms from the Eocene of North America: a new genus of Theaceae based on fruit and seed remains. Botanical Gazette 150: 190–206. Grote PJ, Dilcher DL. 1992. Fruits and seeds of tribe Gordonieae (Theaceae) from the Eocene of North America. American Journal of Botany 79: 744–753. Heiss ML, Heiss RJ. 2011. The story of tea: a cultural history and drinking guide. Ten Speed Press, New York. Keng H. 1962. Comparative morphological studies in Theaceae. University of California Publications in Botany 33: 269–384. Li J, Del Tredici P, Yang S, Donoghue MJ. 2002. Phylogenetic relationships and biogeography of Stewartia (Camellioideae, Theaceae) inferred from nuclear ribosomal DNA ITS sequences. Rhodora 04: 117–133. Li R, Yang JB, Yang SX, Li DZ. 2011. Phylogeny and taxonomy of the Pyrenaria complex (Theaceae) based on nuclear ribosomal ITS sequences. Nordic Journal of Botany 29: 780–787. Liang D, Baas P. 1991. The wood anatomy of the Theaceae. IAWA Bulletin II, 12: 333–353. Mai DH. 1971. Über fossile Lauraceae und Theaceae in Mitteleuropa. Feddes Repertorium 82: 313–341. Prince LM, Parks CM. 2001. Phylogenetic relationships of Theaceae inferred from chloroplast DNA data. American Journal of Botany 88: 2309–2320. Prince LM. 2002. Circumscription and biogeographic patterns in the eastern North American-East Asian genus Stewartia (Theaceae: Stewartieae): insight from chloroplast and nuclear DNA sequence data. Castanea 67: 290–301. Prince LM. 2007. A brief nomenclatural review of genera and tribes in Theaceae. Aliso 24: 105–121. Sealy JR. 1958. A revision of the genus Camellia. Royal Horticultural Society, London. Su Y, Liao W, Wang T, Sun Y, Wei Q, Chang H. 2011. Phylogeny and evolutionary divergence times in Apterosperma and Eurydendron: evidence of a Tertiary origin in southern China. Biochemical Systematics and Ecology 39: 769–777. Vijayan K, Zhang WJ, Tsou CH. 2009. Molecular taxonomy of Camellia (Theaceae) inferred from nrITS sequences. American Journal of Botany 96: 1348–1360. Yang SX, Yang JB, Lei LG, Li DZ, Yoshino H, Ikeda T. 2004. Reassessing the relationships between Gordonia and Polyspora (Theaceae) based on the combined analyses of molecular data from the nuclear, plastid and mitochondrial genomes. Plant Systematics and Evolution 248: 45–55. Yang JB, Yang SX, Li DZ, Lei LG, Ikeda T, Yoshino H. 2006. Phylogenetic relationships of Theaceae inferred from mitochondrial matR gene sequence data. Acta Botanica Yunnanica 29: 29–36.

373. SYMPLOCACEAE SWEETLEAF FAMILY Baas P, Van den Oever L, Zandee M. 1981. Comparative wood anatomy of Symplocos and latitude and altitude of provenance. IAWA Bulletin n.s. 2: 3–24. Fritsch PW, Cruz BC, Almeda F, Wang Y, Shi S. 2006. Phylogeny of Symplocos based on DNA sequences of the chloroplast trnC-trnD intergenic region. Systematic Botany 31: 181–192. Fritsch PW, Kelly LM, Wang Y, Almeda F, Kriebel R. 2008. Revised infrafamilial classification of Symplocaceae based on phylogenetic data from DNA sequences and morphology. Taxon 57: 823–852. Kircheimer F. 1949. Die Symplocaceae des erdgeschichtlichen Vergangenheit. Palaeontographica 90B b: 1–52. Nooteboom HP. 1991. Revision of the Symplocaceae of the Old World, New Caledonia excepted. Leiden Botanical Series 1. Universitaire Pers, Leiden. Wang YH, Fritsch PW, Shi S, Almeda F, Cruz BC, Kelly LM. 2004. Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data. American Journal of Botany 91: 1901–1914. Wood CE Jr, Channell RB. 1960. The genera of the Ebenales in the southeastern United States. Journal of the Arnold Arboretum 41: 1–35. 374. DIAPENSIACEAE PINCUSHION-PLANT FAMILY Doonan S. 1993. The Diapensia family. Bulletin of the American Rock Garden Society 51: 97–106. Murphy HT, Hardin JW. 1976. A new and unique venation pattern in the Diapensiaceae. Bulletin of the Torrey Botanical Club 103: 177–179. Nesom GL. 1983. Galax (Diapensiaceae): geographic variation in chromosome studies. Systematic Botany 8: 1–4. Rönblom K, Anderberg AA. 2002. Phylogeny of Diapensiaceae based on molecular data and morphology. Systematic Botany 27: 383–395. Scott PJ, Day RT. 1983. Diapensiaceae: a review of the taxonomy. Taxon 32: 417–423. Wood CE Jr, Channell RB. 1959. The Empetraceae and Diapensiaceae of the southeastern United States. Journal of the Arnold Arboretum 40: 161–171. Xi YZ, Tang YC. 1990. Pollen morphology and phylogenetic relationships in the Diapensiaceae. Cathaya 2: 89–112. 375. STYRACACEAE STORAX FAMILY Copeland HF. 1938. The Styrax of northern California and the relationships of the Styracaceae. American Journal of Botany 25: 771–780. Dickison WC. 1993. Floral anatomy of the Styracaceae, including observations on intra-ovarian trichomes. Botanical Journal of the Linnean Society 112: 223–255. Dickison WC, Phend KD. 1985. Wood anatomy of the Styracaceae: evolutionary and ecological considerations. IAWA Bulletin, n.s. 6: 3–22. Fritsch PW. 1996. Isozyme analysis of intercontinental disjuncts within Styrax (Styracaceae): implications for the Madrean-Tethyan hypothesis. American Journal of Botany 83: 342–355. Fritsch PW. 1999. Phylogeny of Styrax based on morphological characters, with implications for biogeography and infrageneric classification. Systematic Botany 24: 356–378. Fritsch PW, Morton CM, Chen T, Meldrum C. 2001. Phylogeny and biogeography of the Styracaceae.

FURTHER READING International Journal of Plant Sciences 162: S95–S116. Langenheim J. 2003. Plant resins: chemistry, evolution, ecology, and ethnobotany. Timber Press, Portland. Miers J. 1859. On the natural order Styraceae, as distinguished from the Symplocaceae. Annual Magazine of Natural History, ser. 3, 3: 394–404. Morton CM, Dickison WC. 1992. Comparative pollen morphology of the Styracaceae. Grana 31: 1–15. Morton CM, Chase MW, Kron KA, Swensen SM. 1997. A molecular evaluation of the monophyly of the order Ebenales based upon rbcL sequence data. Systematic Botany 21: 567–586. Schadel WE, Dickison WC. 1979. Leaf anatomy and venation patterns of the Styracaceae. Journal of the Arnold Arboretum 60: 8–37. Van Steenis CGGJ. 1932. The Styracaceae of Netherlands India. Bulletin du Jardin Botanique de Buitenzorg III, 12: 212–272. Wood CE Jr, Channell RB. 1960. The genera of the Ebenales in the southeastern United States. Journal of the Arnold Arboretum 41: 1–35. 376. SARRACENIACEAE AMERICANPITCHERPLANT FAMILY Bayer RJ, Hufford L, Soltis DE. 1996. Phylogenetic relationships in Sarraceniaceae based on rbcL and ITS sequences. Systematic Botany 21: 121–134. Bell CR. 1949. A cytotaxonomic study of the Sarraceniaceae of North America. Journal of the Elisha Mitchell Scientific Society 65: 137–166. Bennett KF, Ellison AM. 2009. Nectar not colour may lure insects to their death. Biology Letters 5: 469–472. Chase MW, Christenhusz MJM, Sanders D, Fay MF. 2010. Murderous plants: Victorian Gothic, Darwin and modern insights into vegetable carnivory. Botanical Journal of the Linnean Society 161: 329–356. Darwin C. 1875. Insectivorous plants. Murray, London. De Buhr LE. 1975. Phylogenetic relationships of the Sarraceniaceae. Taxon 24: 297–306. Jaffe K, Michelangeli F, Gonzaléz JM, Miras B, Ruiz MC. 1992. Carnivory in pitcher plants of the genus Heliamphora (Sarraceniaceae). New Phytologist 122: 733–744. McPherson S. 2006. Pitcher plants of the Americas. McDonald, Woodward Publishing Company, Granville. McPherson S, Schnell D. 2011. Sarraceniaceae of North America. Redfern Natural History Publications, Poole. McPherson S, Wistuba A, Fleischmann A, Nerz J. 2011. Sarraceniaceae of South America. Redfern Natural History Publications, Poole. Neyland R, Merchant M. 2006. Systematic relationships of Sarraceniaceae inferred from nuclear ribosomal DNA sequences. Madroño 53: 223–232. Schaefer HM, Ruxton GD. 2008. Fatal attraction: carnivorous plants roll out the red carpet to lure insects. Biology Letters 4: 153–155. Schneider EL, Carlquist S. 2004. Perforation plate pit remnants in vessels of Sarraceniaceae: possible indicators of relationship and ecology. Journal of the Torrey Botanical Club 131: 1–7. 377. RORIDULACEAE FLYCATCHER-BUSH FAMILY Anderson B. 2005. Adaptations to foliar absorption of faeces: a pathway in plant carnivory. Annals of Botany 95: 757–761. Anderson B, Midgley JJ. 2002. It takes two to tango

but three is a tangle: mutualists and cheaters on the carnivorous plant Roridula. Oecologia 132: 369–373. Carlquist S. 1976. Wood anatomy of Roridulaceae: ecological and phylogenetic implications. American Journal of Botany 63: 1003–1008. Chase MW, Christenhusz MJM, Sanders D, Fay MF. 2010. Murderous plants: Victorian Gothic, Darwin and modern insights into vegetable carnivory. Botanical Journal of the Linnean Society 161: 329–356. Conran JG, Dowd JW. 1994. The phylogenetic relationships of the Byblis-Roridula (ByblidaceaeRoridulaceae) complex inferred from 18S rRNA partial sequences. Plant Systematics and Evolution 188: 73–86. Dolling WR, Palmer JM. 1991. Pameridea (Hemiptera: Miridae): predaceous bugs specific to highly viscid plant genus Roridula. Systematic Entomology 16: 319–328. Ellis AG, Midgley JJ. 1996. A new plant-animal mutualism involving a plant with sticky leaves and a resident hemipteran. Oecologia 106: 478–481. McPherson S. 2008. Glistening carnivores. The stickyleaved insect-eating plants. Redfern Natural History Productions, Poole. McPherson S. 2010. Carnivorous plants and their habitats, 2 vols. Redfern Natural History Productions, Poole. Midgley JJ, Stock WD. 1998. Natural abundance of delta N-15 confirms insectivorous habit of Roridula gorgonias, despite it having no proteolytic enzymes. Annals of Botany 82: 387–388. Warren BH, Hawkins JA. 2006. The distribution of species diversity across a flora’s component lineages: dating the Cape’s ‘relicts’. Proceedings of the Royal Society B, 273: 2149–2158. 378. ACTINIDIACEAE KIWIFRUIT FAMILY Anderberg AA, Rydin C, Källersjö M. 2002. Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. American Journal of Botany 89: 677–687. Atkinson RG, Cipriani G, Whittaker DJ, Gardner RC. 1997. The allopolyploid origin of kiwifruit, Actinidia deliciosa (Actinidiaceae). Plant Systematics and Evolution 205: 111–124. Chat J, Jáuregui B, Petit RJ, Nadot S. 2004. Reticulate evolution in kiwifruit (Actinidia, Actinidiaceae) identified by comparing their maternal and paternal phylogenies. American Journal of Botany 91: 736–747. Cui ZX. 1993. Actinidia in China. Shangdon Scientific Press, Jinan. Dickison WC. 1972. Observations on the floral morphology of some species of Saurauia, Actinidia and Clematoclethra. Journal of the Elisha Mitchell Scientific Society 88: 43–54. He ZC, Li JQ, Cai Q, Wang Q. 2005. The cytology of Actinidia, Saurauia and Clematoclethra (Actinidiaceae). Botanical Journal of the Linnean Society 147: 369–374. Huang H, Li Z, Li J. Kubisiak TL, Layne DR. 2002. Phylogenetic relationships in Actinidia as revealed by RAPD analysis. Journal of the American Society of Horticultural Science 127: 759–766. Hunter GE. 1966. Revision of Mexican and Central American Saurauia (Dilleniaceae). Annals of the Missouri Botanical Garden 53: 47–89. Keller JA, Herendeen PS, Crane PF. 1996. Fossil f lowers and fruits of the Actinidiaceae from

the Campanian (Late Cretaceous) of Georgia. American Journal of Botany 83: 528–541. Li XW, Li JQ, Soejarto DD. 2007. Actinidiaceae. Flora of China 12: 334–360. Liang CF, Ferguson AR. 1986. The botanical nomenclature of the kiwifruit and related taxa. New Zealand Journal of Botany 24: 183–184. Record M. 1945. A collection of woody plants from Melanesia. Tropical Woods 81: 9–30 [p. 27]. Soejarto DD. 1980. Revision of South American Saurauia (Actinidiaceae). Fieldiana Botany new series 2: 1–141. Warrington IJ, Weston GC (eds). 1990. Kiwifruit: science and management. Richards, Auckland. 379. CLETHRACEAE LILY-OF-THE-VALLEYTREE FAMILY Anderberg AA, Zhang X. 2002. Phylogenetic relationships of Cyrillaceae and Clethraceae (Ericales) with special emphasis on the genus Purdiaea Planch. Organisms, Diversity and Evolution 2: 127–136. Berazaín R, Rodríguez S. 1992. Novedades taxonómicas en el género Purdiaea Planchon (Cyrillaceae) en Cuba. Revista del Jardín Botánico Nacional, Universidad de la Habana 13: 21–25. Fior S, Karis PO, Anderberg AA. 2003. Phylogeny, taxonomy, and systematic position of Clethra (Clethraceae, Ericales) with notes on bibliography: evidence from plastid and nuclear DNA sequences. International Journal of Plant Sciences 164: 997–1006. Giebel KP, Dickison WC. 1975. Wood anatomy of Clethraceae. Journal of the Elisha Mitchell Scientific Society 91: 17–26. Kron KA, Chase MW. 1993. Systematics of Ericaceae, Empetraceae, Epacridaceae, and related taxa based upon rbcL sequence data. Annals of the Missouri Botanical Garden 80: 735–741. Sleumer H. 1967. Monographia clethracearum. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 87: 36–175. 380. CYRILLACEAE LEATHERWOOD FAMILY Anderberg AA, Zhang X. 2002. Phylogenetic relationships of Cyrillaceae and Clethraceae (Ericales) with special emphasis on the genus Purdiaea Planch. Organisms, Diversity and Evolution 2: 127–136. Barghoorn ES, Spackman W. 1949. A preliminary study of the flora of the Brandon lignite. American Journal of Science 247: 33–39. Copeland HF. 1953. Observations on the Cyrillaceae, particularly on the reproductive structures of the North American species. Phytomorphology 3: 405–411. Schneider EL, Carlquist S. 2003. Unusual pit membrane remnants in perforation plates of Cyrillaceae. Journal of the Torrey Botanical Society 130: 225–230. Thomas JL. 1960. A monographic study of the Cyrillaceae. Contributions from the Gray Herbarium 186: 1–114. Thomas JL. 1961. The genera of the Cyrillaceae and Clethraceae of the southeastern United States. Journal of the Arnold Arboretum 42: 96–106. Vijayaraghavan MR, Dhar U. 1978. Embryology of Cyrilla and Cliftonia (Cyrillaceae). Botaniska Notiser 131: 127–138. Zhang XP, Anderberg AA. 2002. Pollen morphology in the ericoid clade of the order Ericales, with special emphasis on Cyrillaceae. Grana 41: 201–215.

Plants of the World

737

FURTHER READING 381. ERICACEAE HEATHER FAMILY Anderberg AA, 1994. Phylogeny of Empetraceae, with special emphasis on character evolution in the genus Empetrum. Systematic Botany 19: 35–46. Anderberg AA. 1994. Cladistic analysis of Enkianthus with notes on the early diversification of the Ericaceae. Nordic Journal of Botany 14: 385–401. Bell TL, Pate JS, Dixon KW. 1996. Relationships between fire response, morphology, root anatomy and starch distribution in south-west Australian Epacridaceae. Annals of Botany II, 77: 357–364. Bidartondo MI, Bruns RD. 2001. Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographic structure. Molecular Ecology 10: 2285–2295. Braukmann TWA, Stefanovic S. 2012. Plastid genome evolution in mycoheterotrophic Ericaceae. Plant Molecular Biology 79: 5–20. Carlquist S. 1989. Wood and bark anatomy of Empetraceae; comments on paedomorphosis in woods of certain small shrubs. Aliso 12: 497–515. Collinson ME, Crane PR. 1978. Rhododendron seeds from the Palaeocene of England. Botanical Journal of the Linnean Society 76: 195–205. Crayn DM, Kron KA, Gadek PA, Quinn CJ. 1998. Phylogenetics and evolution of epacrids: a molecular analysis using the plastid gene rbcL with a reappraisal of the position of Lebetanthus. Australian Journal of Botany 46: 187–200. Cullings KW. 1996. Single phylogenetic origin of ericoid mycorrhizae within the Ericaceae. Canadian Journal of Botany 74: 1896–1909. Cullings KW, Szaro TM, Bruns TD. 1996. Evolution of extreme specialisation within a lineage of ectomycorrhizal epiparasites. Nature 379: 63–66. Eck P. 1990. The American cranberry. Rutgers University Press, New Brunswick. Freudenstein JV. 1999. Relationships and character transformation in Pyroloideae (Ericaceae) based on ITS sequences, morphology and development. Systematic Botany 24: 398–408. Fritsch PW, Lu L, Bush CM, Cruz BC, Kron KA, Li DZ. 2011. Phylogenetic analysis of the wintergreen group (Ericaceae) based on six genic regions. Systematic Botany 36: 990–1003. Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. American Naturalist 180: 438–449. Hileman LC, Vasey MC, Parker VT. 2001. Phylogeny and biogeography of the Arbutoideae (Ericaceae): implications for the Madean-Tethyan hypothesis. Systematic Botany 26: 131–143. Jordan GJ, Hill RS. 1996. The fossil record of the Epacridaceae. Annals of Botany II, 77: 341–346. Judd WS, Kron KA. 1993. Circumscription of Ericaceae (Ericales) as determined by preliminary cladistic analysis based on morphological, anatomical and embryological features. Brittonia 45: 99–114. Kron KA, Chase MW. 1993. Systematics of Ericaceae, Empetraceae, Epacridaceae, and related taxa based upon rbcL sequence data. Annals of the Missouri Botanical Garden 80: 735–741 Kron KA, Judd WS, Stevens PF, Crayn DM, Anderberg AA, Gadek PA, Quinn CJ, Luteyn JL. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68: 335–423. Luteyn JL. 1983. Ericaceae part I, Cavendishia. Flora Neotropica Monograph 35. Luteyn JL (ed.) 1995. Ericaceae part II, the superiorovaried genera. Flora Neotropica Monograph 66.

738

Christenhusz, Fay & Chase

Luteyn JL. 2002. Diversity, adaptation, and endemism in Neotropical Ericaceae: biogeographical patterns in the Vaccinieae. Botanical Review 68: 55–87. McGuire AF, Kron KA. 2005. Phylogenetic relationships of European and African ericas. International Journal of Plant Sciences 166: 311–318. Nixon KC, Crepet WL. 1993. Late Cretaceous fossil flowers of ericalean affinity. American Journal of Botany 80: 616–623. Oliver EGH. 2000. Systematics of Ericeae (Ericaceae: Ericoideae) species with indehiscent and partially dehiscent fruits. Contributions from the Bolus Herbarium 19: 1–483. Quinn CJ, Crayn DM, Heslewood MM, Brown EA, Gadek PA. 2003. A molecular estimate of the phylogeny of Styphelieae (Ericaceae). Australian Systematic Botany 16: 581–594. Quinn CJ, Brown EA, Heslewood MM, Crayn DM. 2005. Generic concepts in Styphelieae (Ericaceae): the Cyathodes group. Australian Systematic Botany 18: 493–454. Szkudlarz P. 2009. Variation in seed morphology in the genus Erica L. (Ericaceae). Biodiversity Research and Conservation 16: 1–105. Vander Kloet SP, Avery TS. 2007. The taxonomic utility of staminal features in Vaccinieae (Ericaceae). Taxon 56: 897–904. Wahlert GA, Parker VT, Vasey MC. 2009. The phylogeny of Arctostaphylos (Ericaceae) inferred from nuclear ribosomal ITS sequences. Journal of the Botanical Research Institute of Texas 3: 673–682. 382. MITRASTEMONACEAE NIPPLEDAISY FAMILY Barkman TJ, McNeal JR, Lim SH, Coat G, Croom HB, Young ND, de Pamphilis CW. 2007. Mitochondrial DNA suggests at least 11 origins of parasitism in angiosperms and reveals genomic chimaerism in parasitic plants. BMC Evolutionary Biology 7: 248. Beehler BM. 1994. Canopy-dwelling honeyeater aggressively defends terrestrial nectar source. Biotropica 26: 459–461. Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. American Naturalist 180: 438–449. Meijer W, Veldkamp JF. 1993. A revision of Mitrastema (Rafflesiaceae). Blumea 38: 221–229. Nickrent DL, Blarer A, Qiu YL, Vidal-Russell R, Anderson FE. 2004. Phylogenetic inference in Rafflesiales: the influence of rate heterogeneity and horizontal gene transfer. BMC Evolutionary Biology 4: 40. Watanabe H. 1936–1937. Morphologisch-biologische Studien über die Gattung Mitrastemon I, II, III, IV, V, VI, VII. Journal of Japanese Botany 12: 603–618, 698–711, 759–773, 847–858; 13: 14–24, 75–86, 154–162. 383. ONCOTHECACEAE KANAK-LAUREL FAMILY Baas P. 1975. Vegetative anatomy and the affinities of Aquifoliaceae, Sphenostemon, Phelline, and Oncotheca. Blumea 22: 311–407. Cameron KM. 2003. On the phylogenetic position of the New Caledonian endemic families Paracryphiaceae, Oncothecaceae, and Strasburgeriaceae: A comparison of molecules and morphology. Botanical Review 68: 428–443. Carpenter CS, Dickison WC. 1976. The morphology and relationships of Oncotheca balansae. Botanical Gazette 137: 141–153.

384. ICACINACEAE FALSE-YAM FAMILY Angulo DF, De Stefano RD, Stull GW. 2013. Systematics of Mappia (Icacinaceae) and endemic genus of tropical America. Phytotaxa 116: 1–18. Byng JW, Bernardini B, Joseph JA, Chase MW, Utteridge TMA. 2014. Phylogenetic relationships of Icacinaceae, focussing on the vining genera. Botanical Journal of the Linnean Society 176: 277–294. Howard RA. 1992. A revision of Casimirella, including Humirianthera (Icacinaceae). Brittonia 44: 166–172. Kårehed J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88: 2259–2274. Lens F, Kårehed J, Baas P, Jansen S, Rabaey D, Huysmans S, Hamann T, Smets E. 2008. The wood anatomy of the polyphyletic Icacinaceae s.l., and their relationships within asterids. Taxon 57: 525–552. Pigg KB, Manchester SR, DeVore ML. 2008. Fruits of Icacinaceae (tribe Iodeae) from the Late Paleocene of western North America. American Journal of Botany 95: 824–832. Potgeiter MJ, Van Wyk AE. 1994. Fruit structure of the genus Cassinopsis Sond. (Icacinaceae) in Africa. South African Journal of Botany 60: 117–122. Rankin BD, Stockey RA, Beard G. 2008. Fruits of Icacinaceae from the Eocene Appian Way locality of Vancouver Island, British Columbia. International Journal of Plant Sciences 169: 305–314. Sleumer H. 1971. Icacinaceae. Flora Malesiana ser. 1, 7: 1–87. Stull GW, Moore RW, Manchester SR. 2011. Fruits of Icacinaceae from the Eocene of southeastern North America and their biogeographic implications. International Journal of Plant Sciences 172: 935–947. Stull GW. Herrera F, Manchester SR, Jaramillo C, Tiffney BH. 2012. Fruits of an “Old World” tribe (Phytocreneae; Icacinaceae) from the Paleogene of North and South America. Systematic Botany 37: 784–794. Stull GW, Duno de Stefano R, Soltis DE, Soltis PS. 2015. Resolving basal lamiid phylogeny and the circumscription of Icacinaceae with a plastomescale data set. American Journal of Botany 102: 1794–1813. Utteridge TMA, Nagamasu H, Teo SP, White LC, Gasson P. 2005. Sleumeria (Icacinaceae): a new genus from northern Borneo. Systematic Botany 30: 635–643. 385. METTENIUSACEAE URUPAGUA FAMILY Byng JW, Bernardini B, Joseph JA, Chase MW, Utteridge TMA. 2014. Phylogenetic relationships of Icacinaceae, focussing on the vining genera. Botanical Journal of the Linnean Society 176: 277–294. De Stefano RD, Carnevali F-C G. 2011. Morphologyinferred phylogeny and a revision of the genus Emmotum (Icacinaceae). Annals of the Missouri Botanical Garden 98: 1–27. González F, Betancur J, Maurin O. Freudenstein JV, Chase MW. 2007. Metteniusaceae, an earlydiverging family in the lamiid clade. Taxon 56: 795–800. González F, Rudall PJ. 2010. Flower and fruit characters in the early-divergent lamiid family Metteniusaceae, with particular reference to the evolution of pseudomonomery. American Journal of Botany 97: 191–206.

FURTHER READING Kårehed J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88: 2259–2274. Lozano-Contreras G, De Lozano NB. 1988. Metteniusaceae. Flora de Colombia, monograph 11: 1–53. Potgeiter MJ, Van Wyk AE. 1994. Fruit structure of the southern African species of Apodytes E. Meyer ex Arn. (Icacinaceae). Botanical Journal of the Linnean Society 115: 221–233. Stull GW, Duno de Stefano R, Soltis DE, Soltis PS. 2015. Resolving basal lamiid phylogeny and the circumscription of Icacinaceae with a plastomescale data set. American Journal of Botany 102: 1794–1813. 386. EUCOMMIACEAE CHINESE-RUBBERTREE FAMILY Call VB, Dilcher DL. 1997. The fossil record of Eucommia (Eucommiaceae) in North America. American Journal of Botany 84: 798–814. Eckhardt T. 1963. Some observations on the morphology and embryology of Eucommia ulmoides Oliv. Journal of the Indian Botanical Society 42A: 27–34. Forrest T. 1995. Two thousand years of eating bark: Magnolia officinalis var. biloba and Eucommia ulmoides in traditional Chinese medicine. Arnoldia 55: 13–18. Manchester SR, Chen ZD, Lu AM, Uemura K. 2009. Eastern Asian endemic seed plant genera and their paleogeographic history throughout the Northern Hemisphere. Journal of Systematics and Evolution 47: 1–41. Tippo O. 1940. The comparative anatomy of the secondary xylem and the phylogeny of the Eucommiaceae. American Journal of Botany 27: 832–838. Vincent MA. Eucommia ulmoides (hardy rubber-tree; Eucommiaceae) as an escape in North America. The Michigan Botanist 41: 141–145. Wang YF, Li CS, Collinson ME, Lin J, Sun QJ. 2003. Eucommia (Eucommiaceae), a potential biothermometer for the reconstruction of paleoenvironments. American Journal of Botany 90: 1–7. Zhang K, Dong J, Ma B, Gao J, Han X. 2002. Studies on the distribution differences of the secondary metabolites in Eucommia ulmoides. Scientia Silvae Sinicae 38: 12–16. Zhang YL, Wang EH, Chien NF. 1988. A study on pollen morphology of Eucommia ulmoides Oliver. Acta Phytotaxonomica Sinica 26: 367–370. Zhang ZY, Lu AM, Pan KY. 1990. The anatomy, embryology and systematic relationships of Eucommiaceae. Acta Phytotaxonomica Sinica 28: 430–441. 387. GARRYACEAE TASSELBUSH FAMILY Burge DO. 2011. Molecular phylogenetics of Garrya (Garryaceae). Madroño 58: 249–255. Dahling GV. 1978. Systematics and evolution of Garrya. Contributions to the Gray Herbarium of Harvard University 209: 1–104. Hallock FA. 1930. The relationship of Garrya. The development of the flowers and seeds of Garrya and its bearing on the phylogenetic position of the genus. Annals of Botany 176: 771–812. Iwashina T, Kamenosono K, Hatta H. 1997. Flavonoid glycosides of Aucuba japonica and Helwingia japonica (Cornaceae): phytochemical relationship with the genus Cornus. Journal of Japanese Botany 72: 337–346.

Liston A. 2003. A new interpretation of floral morphology in Garrya (Garryaceae). Taxon 52: 271–276. Muller O, Hirose T, Werger MJA, Hikosaka K. 2011. Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understory evergreen shrub. Annals of Botany 108: 529–536. Noshiro S, Baas P. 1998. Systematic wood anatomy of Cornaceae and allies. IAWA Journal 19: 43–97. 388. RUBIACEAE COFFEE FAMILY Andersson L. 2002. Relationships and generic circumscriptions in the Psychotria complex. Systematic Geography of Plants 72: 167–202. Barrabé L, Maggia L, Pillon Y, Rigault F, Mouly A, Davis AP, Buerki S. 2014. New Caledonian lineages of Psychotria (Rubiaceae) reveal different evolutionary histories and the largest documented plant radiation for the Archipelago. Molecular Phylogenetics and Evolution 71: 15–35. Bremer B. 1996. Phylogenetic studies within Rubiaceae and relationships to other families based on molecular data. Opera Botanica Belgica 7: 33–50. Bremer B. 2009. A review of molecular phylogenetic studies of Rubiaceae. Annals of the Missouri Botanical Garden 96: 4–26. Bremer B, Eriksson O. 1992. Evolution of fruit characters and dispersal modes in the tropical family Rubiaceae. Biological Journal of the Linnean Society 47: 79–95. Bremer B, Eriksson O. 2009. Time tree of Rubiaceae: phylogeny and dating the family, subfamilies and tribes. International Journal of Plant Sciences 170: 766–793. Cantley JT, Swenson NG, Markey A, Keeley SC. 2014. Biogeographic insights on Pacific Coprosma (Rubiaceae) indicate two colonizations to the Hawaiian Islands. Botanical Journal of the Linnean Society 174: 412–424. Davis AP, Govaerts R, Bridson DM, Ruhsam M, Moat J, Brummit NA. 2009. A global assessment of distribution, diversity, endemism, and taxonomic effort in the Rubiaceae. Annals of the Missouri Botanical Garden 96: 68–78. Fay MF, Bremer B, Prance GT, Van der Bank M, Bridson D, Chase MW. 2000. Plastid rbcL sequence data show Dialypetalanthus to be a member of Rubiaceae. Kew Bulletin 55: 853–864. Graham A. 2009. Fossil record of the Rubiaceae. Annals of the Missouri Botanical Garden 96: 90–108. Kainulainen K, Razafimandimbison SG, Bremer B. 2013. Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae (Rubiaceae). Botanical Journal of the Linnean Society 173: 387–406. Koek-Noorman J. 1977. Systematische Holzanatomie einiger Rubiaceen. Berichte der Deutchen Botanischen Gesellschaft 90: 183–190. Lens F, Groeninckx I, Smets E, Dessein S. 2009. Woodiness within the Spermacoceae-Knoxieae alliance (Rubiaceae): retention of the basal woody condition in Rubiaceae or recent innovation? Annals of Botany 103: 1049–1064. Manns U, Wikström N, Taylor CM, Bremer B. 2012. Historical biogeography of the predominantly Neotropical subfamily Cinchonoideae (Rubiaceae): into or out of America? International Journal of Plant Sciences 173: 261–289. Maurin O, Davis AP, Chester M, Mvungi EF, Jaufeerally-Fakim Y, Fay MF. 2007. Towards a phylogeny for Coffea (Rubiaceae): identifying

well-supported lineages based on nuclear and plastid DNA sequences. Annals of Botany 100: 1565–1583. Mouly A, Razafimandimbison SG, Florence J, Jérémie J, Bremer B. 2009. Paraphyly of Ixora and new tribal delimitation of Ixoreae (Rubiaceae): inference from combined chloroplast (rps16, rbcL, and trnT-F) sequence data. Annals of the Missouri Botanical Garden 96: 146–160. Riba J, Valle M, Urbano G, Yritia M, Morte A, Barbanoj MJ. 2003. Human pharmacology of ayahuasca: subjective and cardiovascular effects, monoamine metabolite excretion, and pharmacokinetics. Journal of Pharmacology and Experimental Therapeutics 306: 73–83. Robbrecht E, Manen JF. 2006. The major evolutionary lineages of the coffee family (Rubiaceae, angiosperms). Combined analysis (nDNA and cpDNA) to infer the position of Coptosapelta and Luculia, and supertree construction based on rbcL, rps16, trnL-trnF and atpB-rbcL data. A new classification in two subfamilies, Cinchonoideae and Rubioideae. Systematics and Geography of Plants 76: 85–145. Rydin C, Kainulainen K, Razafimandimbison SG, Smedmark JEE, Bremer B. 2009. Deep divergences in the coffee family and the systematic position of Acranthera. Plant Systematics and Evolution 278: 101–123. Smedmark JEE, Razafimandimbison SG, Wikstöm N, Bremer B. 2014. Inferring geographic range evolution of a pantropical tribe in the coffee family (Lasiantheae, Rubiaceae) in the face of topological uncertainty. Molecular Phylogenetics and Evolution 70: 182–194. Soza VL, Olmstead RG. 2010. Molecular systematics of tribe Rubieae (Rubiaceae): evolution of major clades, development of leaf-like whorls, and biogeography. Taxon 59: 755–771. Ukers WH. 1922. All about coffee. The Tea and Coffee Trade Journal Company, New York. 389. GENTIANACEAE GENTIAN FAMILY Favre A, Yuan YM, Küpfer P, Alvarez N. 2010. Phylogeny of subtribe Gentianinae (Gentianaceae): biogeographic inferences despite limitations in temporal calibration points. Taxon 59: 1701–1711. Halda JJ. 1996. The genus Gentiana. SEN, Dobré. Ho TN, Liu SW. 2001. A worldwide monograph of Gentiana. Science Press, Beijing. Kadereit JW, Von Hagen KB. 2003. The evolution of flower morphology in Gentianaceae-Swertiinae and the roles of key innovations and niche width for the diversification of Gentianella and Halenia in South America. International Journal of Plant Sciences 164: S441–S452. Kissling J. 2010. Taxonomy of Exochaenium and Lagenias: two resurrected genera of tribe Exaceae (Gentianaceae). Systematic Botany 37: 238–253. Maas PJM, Ruyters P. 1986. Voyria and Voyriella (saprophytic Gentianaceae). Flora Neotropica Monograph 41. Merckx VSFT, Kissling J, Hentrich H, Janssens SB, Mennes CB, Specht CD, Smets EF. 2013. Phylogenetic relationships of the mycoheterotrophic genus Voyria and the implications for the biogeographic history of Gentianaceae. American Journal of Botany 100: 712–721. Molina J, Struwe L. 2009. Utility of secondary structure in phylogenetic reconstructions using nrDNA ITS sequences — an example from Potalieae (Gentianaceae: Asteridae). Systematic Botany 34: 414–428.

Plants of the World

739

FURTHER READING Oehler E. 1927. Entwicklungsgeschichtlichezytologische Untersuchungen an einigen saprophytischen Gentianaceen. Planta 3: 641–733. Refulio-Rodriguez NF, Olmstead R. 2014. Phylogeny of Lamiidae. American Journal of Botany 101: 287–299. Struwe L, Albert VA. 2002. Gentianaceae: systematics and natural history. Cambridge University Press, Cambridge. Von Hagen KB, Kadereit JW. 2001. The phylogeny of Gentianella (Gentianaceae) and its colonisation of the Southern Hemisphere as revealed by nuclear and chloroplast sequence variation. Organisms, Diversity and Environment 1: 61–79. Von Hagen KB, Kadereit JW. 2003. The diversification of Halenia (Gentianaceae): ecological opportunity versus key innovation. Evolution 57: 2507–2518. 390. LOGANIACEAE INDIAN-PINK FAMILY Aniszewski T. 2007. Alkaloids — secrets of life. Elsevier, Amsterdam. Backlund M, Oxelman B, Bremer B. 2000. Phylogenetic relationships within the Gentianales based in ndhF and rbcL sequences, with particular reference to the Loganiaceae. American Journal of Botany 87: 1029–1043. Bremer B, Struwe L. 1992. Phylogeny of the Rubiaceae and Loganiaceae: congruence or conflict between morphological and molecular data. American Journal of Botany 79: 1171–1194. Gibbons KL, Henwood MJ, Conn BJ. 2012. Phylogenetic relations in Loganieae (Loganiaceae) inferred from nuclear ribosomal and chloroplast DNA sequence data. Australian Systematic Botany 25: 331–340. Leenhouts PW. 1962. Loganiaceae. Flora Malesiana, ser. 1, vol. 6. Leeuwenberg AJM. 1969. The Loganiaceae of Africa VIII. Strychnos III. Revision of the species with notes on the extra-African sections. Mededeelingen van de Landbouwhogeschool te Wageningen 69: 1–316. 391. GELSEMIACEAE YELLOW-JESSAMINE FAMILY Jiao Z, Li J. 2007. Phylogeny of intercontinental disjunct Gelsemiaceae inferred from chloroplast and nuclear DNA sequences. Systematic Botany 32: 617–627. Leeuwenberg AJM. 1961. The Loganiaceae of Africa II. A revision of Mostuea Didr. Mededeelingen van de Landbouwhogeschool te Wageningen 61: 1–31. Refulio-Rodriguez NF, Olmstead R. 2014. Phylogeny of Lamiidae. American Journal of Botany 101: 287–299. Rueangsawang K, Chantaranothai P. 2014. Studies on Thai Pteleocarpaceae. Tropical Natural History 14: 1–6. Struwe L, Soza VL, Manickam S, Olmstead RG. 2014. Gelsemiaceae (Gentianales) expanded to include the enigmatic Asian genus Pteleocarpa. Botanical Journal of the Linnean Society 175: 482–496. 392. APOCYNACEAE PERIWINKLE FAMILY Aniszewski T. 2007. Alkaloids — secrets of life. Elsevier, Amsterdam. Bruyns PV. 2000. Phylogeny and biogeography of the stapeliads. Plant Systematics and Evolution 221: 199–226. Bruyns PV. 2005. Stapeliads of southern Africa and Madagascar, 2 volumes. Umdaus Press, Hatfield. Bruyns PV, Al Farsi A, Hedderson T. 2010. Phylogenetic

740

Christenhusz, Fay & Chase

relationships in Caralluma R. Br. (Apocynaceae). Taxon 59: 1031–1043. Burge DO, Mugford K, Hastings AP, Agrawal AA. 2013. Phylogeny of the plant genus Pachypodium (Apocynaceae). PeerJ 1: e70. Chuba D, Goyder D, Chase MW, Fishbein M. 2017. Phylogenetics of the African Asclepias complex (Apocynaceae) based on three plastid DNA regions. Systematic Botany 42: 148–159. Civeyrel L, Le Thomas A, Ferguson K, Chase MW. 1998. Critical examination of palynological characters used to delimit Asclepiadaceae in comparison to the molecular phylogeny obtained from plastid matK sequences. Molecular Phylogenetics and Evolution 9: 517–527. Endress ME, Liede-Schumann S, Mevre U. 2007. Advances in Apocynaceae: the enlightenment, an introduction. Annals of the Missouri Botanical Garden 94: 259–267. Endress ME, Liede-Schumann S, Meve U. 2014. An updated classification for Apocynaceae. Phytotaxa 159: 175–194. Fishbein M, Chuba D, Ellison C, Mason-Gamer RJ, Lynch SP. 2011. Phylogenetic relationships of Asclepias (Apocynaceae) inferred from non-coding cloroplast DNA sequences. Systematic Botany 36: 1008–1023. Goyder DJ. 2009. Nomenclatural changes resulting from the transfer of tropical African Sarcostemma to Cynanchum (Apocynaceae: Asclepiadoideae). Kew Bulletin 63: 471. Goyder DJ, Nicholas A, Liede-Schumann S. 2007. Phylogenetic relationships in subtribe Asclepiadinae (Apocynaceae: Asclepiadoideae). Annals of the Missouri Botanical Garden 94: 423–434. Ionta GM, Judd WS. 2007. Phylogenetic relationships in Periplocoideae (Apocynaceae s.l.) and insights into the origin of pollinia. Annals of the Missouri Botanical Garden 94: 360–375. Lahaye R, Klackenberg J, Källersjö M, Van Campo E, Civeyrel L. 2007. Phylogenetic relationships between derived Apocynaceae s.l. and within Secamonoideae based on four chloroplast sequences. Annals of the Missouri Botanical Garden 94: 376–391. Leeuwenberg AJM. 1994. Taxa of Apocynaceae above the genus level. Series of revisions of Apocynaceae. XXXVIII. Wageningen Agricultural University Papers 94: 45–60. Lens F, Endress ME, Baas P, Jansen S, Smets E. 2008. Wood anatomy of Rauvolfioideae (Apocynaceae): a search for meaningful non-DNA characters at the tribal level. American Journal of Botany 95: 1199–1215. Lens F, Endress ME, Baas P, Jansen S, Smets E. 2009. Vessel grouping patterns in subfamilies Apocynoideae and Periplocoideae confirm phylogenetic value of wood structure within Apocynaceae. American Journal of Botany 96: 2168–2182. Liede S, Kunze H. 2002. Cynanchum and Cynanchinae (Apocynaceae — Asclepiadoideae): a molecular, anatomical and latex triterpenoid study. Organisms, Diversity and Evolution 2: 239–269. Liede S, Täuber A. 2000. Sarcostemma R. Br. (Apocynaceae–Asclepiadoideae) — a controversial generic circumscription reconsidered: evidence from trnL-F spacers. Plant Systematics and Evolution 225: 133–140. Liede-Schumann S, Kong H, Meve U, Thiv M. 2012. Vincetoxicum and Tylophora (Apocynaceae:

Asclepiadoideae: Asclepiadeae) — two sides of the same medal: independent shifts from tropical to temperate habitats. Taxon 61: 803–825. Livschultz T. 2010. The phylogenetic position of milkweeds (Apocynaceae subfamilies Secamonoideae and Asclepiadoideae): evidence from the nucleus and chloroplast. Taxon 59: 1016–1030. Livschultz T, Middleton DJ, Endress ME, Williams J. 2007. Phylogeny of Apocynoideae and the APSA clade. Annals of the Missouri Botanical Garden 94: 323–361. Meve U, Liede-Schumann S. 2007. Ceropegia (Apocynaceae, Ceropegieae, Stapeliinae): paraphyletic, but still taxonomically sound. Annals of the Missouri Botanical Garden 94: 392–406. Nazar N, Goyder DJ, Clarkson JJ, Mahmood T, Chase MW. 2013. The taxonomy and systematics of Apocynaceae: where we stand in 2012. Botanical Journal of the Linnean Society 171: 482–490. Plowes DCH. 1995. A reclassification of Caralluma R.Brown (Stapelieae: Asclepiadaceae). Haseltonia 3: 49–70. Rapini A, Chase MW, Konno TUP. 2006. Phylogenetics of South American Asclepiadeae (Apocynaceae). Taxon 55: 119–124. Sennblad B, Bremer B. 1996. The familial and subfamilial relationships of Apocynaceae and Asclepiadaceae evaluated with rbcL data. Plant Systematics and Evolution 202: 153–175. Simões AO, Livschultz T, Conti E, Endress ME. 2007. Phylogeny and systematics of the Rauvolfioideae (Apocynaceae) based on molecular and morphological evidence. Annals of the Missouri Botanical Garden 94: 268–297. Simões AO, Endress ME, Conti E. 2010. Systematics and character evolution of Tabernaemontaneae (Apocynaceae, Rauvolfioideae) based on molecular and morphological evidence. Taxon 59: 772–790. Surveswaran S, Sun M, Grimm GW, Liede-Schumann S. 2014. On the systematic position of some Asian enigmatic genera of Asclepiadoideae (Apocynaceae). Botanical Journal of the Linnean Society 174: 601–619. Venter HJT, Verhoeven RL. 2001. Diversity and relationships within the Periplocoideae (Apocynaceae). Annals of the Missouri Botanical Garden 88: 550–568. 393. BORAGINACEAE FORGET-ME-NOT FAMILY Cohen JI. 2014. A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of character evolution. Cladistics 30: 139–169. Ferguson DM. 1999. Phylogenetic analysis and relationships in Hydrophyllaceae based on ndhF sequence data. Systematic Botany 23: 253–268. Gottschling M, Diane N, Hilger HH, Weigend M. 2004. Testing hypotheses on disjunctions present in the primarily woody Boraginales: Ehretiaceae, Cordiaceae, and Heliotropaceae, inferred from ITS1 sequence data. International Journal of Plant Sciences 165: S123–S135. Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. American Naturalist 180: 438–449. Luebert F, Brokamp G, Wen J, Weigend M, Hilger HH. 2011. Phylogenetic relationships and morphological diversity in Neotropical Heliotropium (Heliotropaceae). Taxon 60: 663–680. Luebert F et al. (19 others). 2016. Familial classification of the Boraginales. Taxon 65: 502–522.

FURTHER READING Moore MJ, Jansen RK. 2006. Molecular evidence for the age, origin, and evolutionary history of the American desert plant Tiquilia (Boraginaceae). Molecular Phylogenetics and Evolution 39: 668–687. Nazaire M, Hufford L. 2014. A broad phylogenetic analysis of Boraginaceae: implications for the relationships of Mertensia. Systematic Botany 37: 758–783. Nazaire M, Wang XQ, Hufford L. 2014. Geographic origins and patterns of radiation of Mertensia (Boraginaceae). American Journal of Botany 101: 104–118. Weigend M, Gottschling M, Selvi F, Hilger HH. 2010. Fossil and extant Western Hemisphere Boraginaceae, and the polyphyly of “Trigonotidae” Riedl (Boraginaceae: Boraginoideae). Systematic Botany 35: 409–419. Weigend M, Luebert F, Gottschling M, Couvreur TLP, Hilger HH, Miller JS. 2014. From capsules to nutlets — phylogenetic relationships in the Boraginales. Cladistics 30: 508–518. 394. VAHLIACEAE FLINDERSBUSH FAMILY Bridson DM. 1974. A revision of the family Vahliaceae. Kew Bulletin 30: 163–182. Cutler DF, Gregory M. 1998. Anatomy of the dicotyledons, volume 4, Saxifragales (sensu Armen Takhtajan 1983), ed. 2. Clarendon Press, Oxford. Refulio-Rodriguez NF, Olmstead R. 2014. Phylogeny of Lamiidae. American Journal of Botany 101: 287–299. Weigend M, Luebert F, Gottschling M, Couvreur TLP, Hilger HH, Miller JS. 2014. From capsules to nutlets — phylogenetic relationships in the Boraginales. Cladistics 30: 508–518. 395. CONVOLVULACEAE BINDWEED FAMILY Braukmann TWA, Kuzmina M, Stefanović S. 2013. Plastid genome evolution across the genus Cuscuta (Convolvulaceae): two clades within subgenus Grammica exhibit extensive genome loss. Journal of Experimental Botany 64: 977–989. C.U. 2014. Convolvulaceae unlimited. http://convolvulaceae.myspecies.info (accessed July 2014). Eich E. 2008. Solanaceae and Convolvulaceae: secondary metabolites. Springer, Berlin. García MA, Costea M, Kuzmina M, Stefanović S. 2014. Phylogeny, character evolution, and biogeography of Cuscuta (dodders; Convolvulaceae) inferred from coding plastid and nuclear sequences. American Journal of Botany 101: 670–690. García MA, Martín MP. 2007. Phylogeny of Cuscuta subgenus Cuscuta (Convolvulaceae) based on nrDNA ITS and chloroplast trnL intron sequences. Systematic Botany 32: 899–916. Jayasuriya KMGG, Baskin JM, Geneve RL, Baskin CC. 2009. Phylogeny of seed dormancy in Convolvulaceae subfamily Convolvuloideae (Solanales). Annals of Botany 103: 45–63. Manos PS, Miller RE, Wilkin P. 2001. Phylogenetic analysis of Ipomoea, Argyreia, Stictocardia, and Turbina suggests a generalized model of morphological evolution in morning glories. Systematic Botany 26: 585–602. McNeill JR, Kuehl JV, Boore JL, dePamphilis CW. 2007. Complete plastid genome sequences suggest strong selection for the retention of photosynthetic genes in the parasitic plant genus Cuscuta. BMC Plant Biology 7: 57. Refulio-Rodriguez NF, Olmstead R. 2014. Phylogeny

of Lamiidae. American Journal of Botany 101: 287–299. Stefanović S, Austin DF, Olmstead RG. 2003. Classification of Convolvulaceae: a phylogenetic approach. Systematic Botany 28: 791–806. 396. SOLANACEAE NIGHTSHADE FAMILY Aguilar-Meléndez A, Morell PL, Roose ML, Kim SC. 2009. Genetic diversity and structure in semiwild and domesticated chiles (Capsicum annuum; Solanaceae) from Mexico. American Journal of Botany 96: 1190–1202. Avery AG, Satina S, Rietsema J. 1959. The genus Datura. Ronald Press, New York. Bai Y, Lindhout P. 2007. Domestication and breeding of tomatoes: what have we gained and what can we gain in the future? Annals of Botany 100: 1085–1094. Dillon MO, Tu T, Xie L, Quipuscoa Silvestre V, Wen J. 2009. Biogeographic diversification in Nolana (Solanaceae), a ubiquitous member of the Atacama and Peruvian deserts along the western coast of South America. Journal of Systematics and Evolution 47: 457–476. Doncheva T, Berkov S, Philipov S. 2006. Comparative study of the alkaloids in the tribe Datureae and their chemosystematic significance. Biochemical Systematics and Ecology 34: 478–488. Eich E. 2008. Solanaceae and Convolvulaceae: secondary metabolites. Springer, Berlin. Fay MF, Olmstead RG, Richardson JE, Santiago E, Prance GT, Chase MW. 1998. Molecular data support the inclusion of Duckeodendron cestroides in Solanaceae. Kew Bulletin 53: 203–212. Fay MF, Thomas VE, Knapp S. 2007. Mellissia begoniifolia, Solanaceae. Curtis’s Botanical Magazine 24: 243–250. Filipowicz N, Renner SS. 2012. Brunfelsia (Solanaceae): a genus evenly divided between South America and radiations on Cuba and other Antillean islands. Molecular Phylogenetics and Evolution 64: 1–11. Goodspeed TH. 1954. The genus Nicotiana. Chronica Botanica, Waltham. Hawkes JG, Lester RN, Skelding AD (eds). 1979. The biology and taxonomy of the Solanaceae. Linnean Society/Academic Press, London. Hunziker AT. 2001. Genera solanacearum: the genera of Solanaceae illustrated, arranged according to a new system. Gantner, Ruggell. Kelly LJ, Leitch AR, Clarkson JJ, Knapp S, Chase MW. 2012. Reconstructing the complex evolutionary history of wild allopolyploid tobaccos (Nicotiana section Suaveolentes). Evolution 67: 80–94. Knapp S, Bohs L, Nee M, Spooner DM. 2004. Solanaceae — a model for linking genomics with biodiversity. Comparative and Functional Genomics 5: 285–291. Knapp S. 2002. Tobacco to tomatoes: a phylogenetic perspective on fruit diversity in the Solanaceae. Journal of Experimental Botany 53: 2001–2022. Knapp S. 2010. On ‘various contrivances’: pollination, phylogeny and flower form in Solanaceae. Philosophical Transactions of the Royal Society of London B 365: 449–460. Levin RA, Bernadello G, Whiting C, Miller JS. 2011. A new generic circumscription in tribe Lycieae (Solanaceae). Taxon 60: 681–690. Martins TR, Barkman TJ. 2005. Reconstruction of Solanaceae phylogeny using the nuclear gene SAMT. Systematic Botany 30: 435–447.

Olmstead RG, Bohs L, Migid HA, Santiago-Valentin E, García VF, Collier SM. 2008. A molecular phylogeny of Solanaceae. Taxon 57: 1159–1181. Olmstead RG. 2013. Phylogeny and biogeography of Solanaceae, Verbenaceae, and Bignoniaceae: a comparison of continental and intercontinental diversification patterns. Botanical Journal of the Linnean Society 171: 80–102. Ovchinnikova A, Krylova E, Gavrilenko T, Smekalova T, Zhuk M, Knapp S, Spooner DM. 2011. Taxonomy of cultivated potatoes (Solanum section Petota: Solanaceae). Botanical Journal of the Linnean Society 165: 107–155. PBI Solanum Project. 2014 (continuously updated). Solanaceae Source. http://www.solanaceaesource.org/ Perry L, Dickau R, Zarrillo S, Holst I, Pearsall DM, Piperno DR, Berman MJ, Cooke RG, Rademaker K, Ranere AJ, Raymond JS, Sandweiss DH, Scaramelli F, Tarble K, Zeidier JA. 2007. Starch fossils and the domestication and dispersal of chili peppers (Capsicum spp. L.) in the Americas. Science 315: 986–988. Särkinen T, Bohs L, Olmstead RG, Knapp S. 2013. A phylogenetic framework for evolutionary study of the nightshades (Solanaceae): a dated 1000-tip tree. BMC Evolutionary Biology 13: 214. Stern S, Agra MdF, Bohs L. 2011. Molecular delimitation of clades within New World species of “spiny solanums” (Solanum subg. Leptostemon). Taxon 60: 1429–1441. Tewksbury J, Reagan KM, Machnicki NJ, Carlo TA, Haak DC, Calderón-Peñaloza AL, Levey DJ. 2008. Evolutionary ecology of pungency in wild chilies. Proceedings of the National Academy of Sciences of the USA 105: 11808–11811. Tu T, Volis S, Dillon MO, Sun H, Wen J. 2010. Dispersals of Hyoscyameae and Mandragoreae (Solanaceae) from the New World to Eurasia in the early Miocene and their biogeographic diversification within Eurasia. Molecular Phylogenetics and Evolution 57: 1225–1237. Vorontsova MS, Stern S, Bohs L, Knapp S. 2013. African spiny Solanum (subgenus Leptostemon, Solanaceae): a thorny phylogenetic tangle. Botanical Journal of the Linnean Society 173: 176–193. Wu F, Tanksley SD. 2010. Chromosome evolution in the plant family Solanaceae. BMC Genomics 11: 182. Zuckerman L. 1999. The potato: how the humble spud rescued the western world. North Point Press, New York. 397. MONTINIACEAE WILD-CLOVEBUSH FAMILY Carlquist S. 1989. Wood anatomy and relationships of Montinia. Aliso 12: 369–378. Decraene LP, Linder HP, Smets EF. 2000. The questionable relationship of Montinia (Montiniaceae) from a floral ontogenetic and anatomical study. American Journal of Botany 87: 1408–1424. 398. SPHENOCLEACEAE GOOSEWEED FAMILY Carter R, Jones JC, Goddard RH. 2014. Sphenoclea zeylanica (Sphenocleaceae) in North America — dispersal, ecology, and morphology. Castanea 79: 33–50. Erbar C. 1995. On the floral development of Sphenoclea zeylanica (Sphenocleaceae, Camp-anulales)

Plants of the World

741

FURTHER READING — SEM investigations on herbarium material. Botanische Jahrbücher f ür Systematik , Pflanzengeschichte und Pflanzengeographie 117: 469–483. Hirai N, Sakashita S-I, Sano T, Inoue T, Ohigashi H, Premasthira CU, Asakawa Y, Harada J, Fujii Y. 2000. Allelochemicals of the tropical weed Sphenoclea zeylanica. Phytochemistry 55: 131–140. Kausik SB, Subramanyam K. 1946. A contribution to the life history of Sphenoclea zeylanica Gaertn. Proceedings of the Indian Academy of Sciences B, 23: 274–280. Monod T. 1980. A propos du Sphenoclea zeylanica (Sphenocleaceae). Adansonia Séries 2, 20: 147–164. Subramanyam K. 1950. A contribution to our knowledge of the systematic position of the Sphenocleaceae. Proceedings of the Indian Acadedemy of Sciences B, 31: 60–65. 399. HYDROLEACEAE FIDDLELEAF FAMILY Bennett AW. 1870. Review of the genus Hydrolea, with descriptions of three new species. Journal of the Linnean Society of London, Botany 11: 266–279. Constance L, Chuang TI. 1982. SEM survey of pollen morphology and classification in Hydrophyllaceae (waterleaf family). American Journal of Botany 69: 40–53. Davenport LJ. 1988. A monograph of Hydrolea (Hydrophyllaceae). Rhodora 90: 169–208 Di Fulvio TE. 1989. Observaciones embrológicas en especies Argentinas de Hydrolea (Hydrophyllaceae) con especial referencia a la endospermogénesis. Kurtziana 20: 33–64. Di Fulvio TE. 1990. Endospermogénesis y taxonomía de la familia Hydrophyllaceae y su relación con las demas Gamopetalas. Academia Nacional de Ciencias Exactas Físicas y Naturales de Buenos Aires 5: 73–82. 400. PLOCOSPERMATACEAE STAGHORNSHRUB FAMILY Chiang F, Frame D. 1987. The identity of Lithophytum (Loganiaceae, Plocospermeae). Brittonia 39: 260–262. D’Arcy WG, Keating RC. 1973. The affinities of Lithophytum: a transfer from Solanaceae to Verbenaceae. Brittonia 25: 213–225. Leeuwenberg AJM. 1967. Notes on American Loganiaceae I. Revision of Plocosperma Benth. Acta Botanica Neerlandica 16: 56–61. Struwe L, Albert VA, Bremer B. 1994. Cladistics and family-level classification of the Gentianales. Cladistics 10: 175–206. 401. CARLEMANNIACEAE FRAGRANTPRINCESS FAMILY Airy Shaw HK. 1965. A new species of the genus Silvianthus Hook.f., and on the family Carlemanniaceae. Kew Bulletin 19: 507–512. Bentham G. 1853. On three new genera connected with the Indian flora. Journal of Botany (Hooker) 5: 304–309. Bremekamp CEB. 1939. On the position of the genera Carlemannia Benth. and Silvianthus Hook.f. Recueil des Travaux Botanique Néerlandais 36: 372. Tange C. 1998. Silvianthus (Carlemanniaceae) a genus and family new to Thailand. Thai Forest Bulletin 26: 59–65. Yang X, Lu SG, Peng H. 2007. First report of chromosome numbers of the Carlemanniaceae (Lamiales). Journal of Plant Research 120: 707–712.

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402. OLEACEAE OLIVE FAMILY Baas P, Esser PM, Van der Westen MET, Zandee M. 1988. Wood anatomy of the Oleaceae. IAWA Bulletin 9: 103–182. Baldoni L, Guerrero C, Sossey-Aloui K, Abbott AG, Angioilillo A, Lumaret R. 2002. Phylogenetic relationships among Olea species, based on nucleotide variation at a non-coding chloroplast DNA region. Plant Biology 4: 346–351. Besnard G, De Casas RR, Christin P-A, Vargas P. 2009. Phylogenetics of Olea (Oleaceae) based on plastid and nuclear ribosomal DNA sequences: Tertiary climatic shifts and lineage differentiation times. Annals of Botany 104: 143–160. Call VB, Dilcher DL. 1992. Investigations of angiosperms from the Eocene of southeastern North America: samaras of Fraxinus wilcoxiana Berry. Review of Palaeobotany and Palynology 74: 249–266. Fiala JL. 1988. Lilacs, the genus Syringa. Timber Press, Portland. García Verdugo C, Fay MF, Granado Yela C, de Casas RR, Balaguer L, Besnard G, Vargas P. 2009. Genetic diversity and differentiation processes in the ploidy series of Olea europaea L.: a multiscale approach from subspecies to insular populations. Molecular Ecology 18: 454–467. Harborne JB, Green PS. 1980. A chemotaxonomic survey of flavonoids in leaves of the Oleaceae. Botanical Journal of the Linnean Society 81: 155–167. Hong-Wa C, Besnard G. 2013. Intricate patterns of phylogenetic relationships in the olive family as inferred from multi-locus plastid and nuclear DNA sequence analysis: a close-up on Chionanthus and Noronhia (Oleaceae). Molecular Phylogenetics and Evolution 67: 367–378. Hong-Wa C, Besnard G. 2014. Species limits and diversification in the Madagascar olive (Noronhia, Oleaceae). Botanical Journal of the Linnean Society 174: 141–161. Johnson LAS. 1957. A review of the family Oleaceae. Contributions from the New South Wales National Herbarium 2: 395–418. Kim KJ, Jansen RK. 1998. A chloroplast DNA phylogeny of lilacs (Syringa, Oleaceae): plastome groups show a strong correlation with crossing groups. American Journal of Botany 85: 1338–1351. Naghiloo S, Dadpour MR, Gohari G, Endress PK. 2013. Comparative study of inflorescence development in Oleaceae. American Journal of Botany 100: 647–663. Sehr EM, Weber A. 2009. Floral ontogeny of Oleaceae and its systematic implications. International Journal of Botany 170: 845–859. Wallander E. 2013. Systematics and floral evolution in Fraxinus (Oleaceae). Belgische Dendrologie Belge 2012: 38–58. Wallander E, Albert VA. 2000. Phylogeny and classification of Oleaceae based on rps16 and trnL-F sequence data. American Journal of Botany 87: 1827–1841. 403. TETRACHONDRACEAE RUSTWEED FAMILY Jensen SR. 1999. Chemical relationships of Polypremum procumbens, Tetrachondra hamiltonii, and Peltanthera floribunda. Biochemical Systematics and Ecology 28: 45–51. Oxelman B, Backlund B, Bremer B. 1999. Relationships of Buddlejaceae s.l. investigated using parsimony jackknife and branch support

analysis of chloroplast ndhF and rbcL sequence data. Systematic Botany 24: 164–182. Skottsberg C. 1912. Tetrachondra patagonica n. sp. und die systematische Stellung der Gattung. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie Beiblatt 107: 17–26. Wagstaff SJ, Martinsson K. Swenson U. 2000. Divergence estimates of Tetrachondra hamiltonii and T. patagonica (Tetrachondraceae) and their implications for austral biogeography. New Zealand Journal of Botany 38: 587–596. 404. GESNERIACEAE GLOXINIA FAMILY Andersson S. 2006. On the phylogeny of the genus Calceolaria (Calceolariaceae) as inferred from ITS and plastid matK sequences. Taxon 55: 125–137. Bunting GS, Duke A. 1961. Sanango: new Amazonian genus of Loganiaceae. Annals of the Missouri Botanical Garden 48: 269–274. Citerne H, Cronk QCB. 1999. The origin of the peloric Sinningia. New Plantsman 6: 219–222. Christenhusz MJM. 2012. On African violets and Cape primroses — towards a monophyletic Streptocarpus (Gesneriaceae). Phytotaxa 46: 3–9. Clark JL, Herendeen PS, Skog LE, Zimmer EA. 2006. Phylogenetic relationships and generic boundaries in the Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and morphological data. Taxon 55: 313–336. Dendruangboripant J, Cronk QCB, Kokubugata G, Möller M. 2007. Variation and inheritance of nuclear ribosomal DNA clusters in Streptocarpus (Gesneriaceae) and their biological and phylogenetic implications. International Journal of Plant Sciences 168: 455–467. Haston E, Ronse de Craene LP. 2007. Inflorescence and f loral development in Streptocarpus and Saintpaulia (Gesneriaceae) with particular reference to the impact of bracteole suppression. Plant Systematics and Evolution 265: 13–25. Hughes M, MacMaster G, Möller M, Bellstedt DU, Edwards TJ. 2006. Breeding system of a plesiomorphic floral type: an investigation of small flowered Streptocarpus (Gesneriaceae) species. Plant Systematics and Evolution 262: 13–24. Hunziker AT, Di Fulvio E. 1958. Observaciones morfológicas sobre Peltanthera (Loganiaceae), con referencia a su posicion systematica. Boletin de la Academia Nacional de Ciencias. Cordoba (Argentina) 40: 217–228. Jensen SR. 1999. Chemical relationships of Polypremum procumbens, Tetrachondra hamiltonii, and Peltanthera floribunda. Biochemical Systematics and Ecology 28: 45–51. Mayr EM, Weber A. 2006. Calceolariaceae: floral development and systematic implications. American Journal of Botany 93: 327–343. Moore HE 1957. African violets, gloxinias and their relatives: a guide to the cultivated gesneriads. Macmillan, New York. Möller M, Cronk QCB. 2001. Phylogenetic studies in Streptocarpus (Gesneriaceae): reconstruction of biogeographic history and distribution patterns. Systematics and Geography of Plants 71: 545–555. Möller M, Middleton D, Nishii K, Wei YG, Sontag S, Weber A. 2011. A new delineation for Oreocharis incorporating an additional ten genera of Chinese Gesneriaceae. Phytotaxa 23: 1–36. Möller M, Forrest A, Wei YG, Weber A. 2011. A molecular phylogenetic assessment of the advanced Asiatic and Malesian didymocarpoid Gesneriaceae

FURTHER READING with focus on non-monophyletic and monotypic genera. Plant Systematics and Evolution 292: 223–248. Möller M, Clark JL. 2013. The state of molecular studies in the family Gesneriaceae: a review. Selbyana 31: 95–125. Nishii K, Hughes M, Briggs M, Haston E, Christie F, DeVilliers MJ, Hanekom T, Roos WG, Bellstedt DU, Möller M. 2015. Streptocarpus redefined to include all Afro-Malagasy Gesneriaceae: molecular phylogenies prove congruent with geographical distribution and basic chromosome numbers and uncover remarkable morphological homoplasy. Taxon 64: 1243–1274. Perret M, Chautems A, De Araujo AO, Salamin N. 2013. Temporal and spatial origin of Gesneriaceae in the New World inferred from plastid DNA sequences. Botanical Journal of the Linnean Society 171: 61–79. Smith JF. 2000. Phylogenetic resolution within the tribe Episcieae (Gesneriaceae): congruence of ITS and ndhF sequences from parsimony and maximum-likelihood analysis. American Journal of Botany 87: 883–897. Smith JF, Draper SB, Hileman LC, Baum DA. 2004. A phylogenetic analysis within tribes Gloxinieae and Gesnerieae (Gesnerioideae: Gesneriaceae). Systematic Botany 29: 947–958. Weber A. 2013. Pair-flowered cymes in Lamiales: structure, distribution and origin. Annals of Botany 112: 1577–1595. Weber A, Middleton DJ, Forrest A, Kiew R, Lim CL, Rafidah AR, Sontag S, Triboun P, Wei YG, Tao TL, Möller M. 2011. Molecular systematics and remodelling of Chirita and associated genera (Gesneriaceae). Taxon 60: 767–790. Weber A, Clark JL, Möller M. 2013. A new formal classification of Gesneriaceae. Selbyana 31: 68–94. Wiehler H. 1994. A re-examination of Sanango racemosum 4. Its new systematic position in Gesneriaceae. Taxon 43: 625–632. Zimmer EA, Roalson EH, Skog LE, Boggan JK, Idnurm A. 2002. Phylogenetic relationships in the Gesnerioideae (Gesneriaceae) based on nrDNA ITS and cpDNA trnL-F and trnE-T spacer region sequences. American Journal of Botany 89: 296–311. 405. PLANTAGINACEAE SPEEDWELL FAMILY Albach DC, Chase MW. 2001. Paraphyly of Veronica (Veroniceae; Scrophulariaceae): evidence from the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA. Journal of Plant Research 114: 9–18. Albach DC, Chase MW. 2004. Incongruence in Veroniceae (Plantaginaceae): evidence from two plastid and a nuclear ribosomal DNA region. Molecular Phylogenetics and Evolution 32: 183–197. Albach DC, Martínez-Ortega MM, Fischer MA, Chase MW. 2004. Evolution of Veroniceae: a phylogenetic perspective. Annals of the Missouri Botanical Garden 91: 275–302. Albach DC, Martínez-Ortega MM, Fischer MA, Chase MW. 2004. A new classification of the tribe Veroniceae — problems and a possible solution. Taxon 53: 429–452. Albach DC, Meudt HM, Oxelman B. 2005. Piecing together the “new” Plantaginaceae. American Journal of Botany 92: 297–315. Albach DC, Meudt HM. 2010. Phylogeny of Veronica in the Southern and Northern Hemispheres based

on plastid, nuclear ribosomal and nuclear low-copy DNA. Molecular Phylogenetics and Evolution 54: 457–471. Baldwin BG, Kalisz S, Armbruster WS. 2011. Phylogenetic perspectives on diversification, biogeography, and floral evolution of Collinsia and Tonella (Plantaginaceae). American Journal of Botany 98: 731–753. Bello MA, Chase MW, Olmstead R, Rønsted N, Albach D. 2002. The páramo endemic Aragoa is the sister genus of Plantago (Plantaginaceae; Lamiales): evidence from plastid rbcL and nuclear ribosomal ITS sequence data. Kew Bulletin 57: 585–597. Carlson MC. 1957. Monograph of the genus Russelia (Scrophulariaceae). Fieldiana Botany 29(4). Natural History Museum, Chicago. Fay MF, Bennett JR, Dixon KW, Christenhusz MJM. 2010. Parasites, their relationships and the disintegration of Scrophulariaceae sensu lato. Curtis’s Botanical Magazine 26: 286–313. Fritsch PW, Almeda F, Martins AB, Cruz BC, Estes D. 2007. Rediscovery and phylogenetic placement of Philcoxia minensis (Plantaginaceae), with a test of carnivory. Proceedings of the California Academy of Sciences 58: 447–467. Ghebrehiwet M, Bremer B, Thulin M. 2000. Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data. Plant Systematics and Evolution 220: 223–239. Hegelmaier F. 1864. Monographie der Gattung Callitriche. Ebner, Seubert, Stuttgart. Hoggard RK, Kores PJ, Molvray M, Hoggard GD, Broughton DA. 2003. Molecular systematics and biogeography of the amphibious genus Littorella (Plantaginaceae). American Journal of Botany 90: 429–435. Hufford L, McMahon M. 2004. Morphological evolution and systematics of Synthyris and Besseya (Veronicaceae): a phylogenetic analysis. Systematic Botany 29: 716–736. Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisens WJ, Reeves PA. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88: 348–361. Oxelman B, Kornhall P, Olmstead RG, Bremer B. 2005. Further disintegration of Scrophulariaceae. Taxon 54: 411–425. Straw RM. 1966. A redefinition of Penstemon (Scrophulariaceae). Brittonia 18: 80–95. Tank DC, Beardsley PM, Kelchner SA, Olmstead RG. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19: 289–307. Taylor P, Souza VG, Giulietti AM, Harley RM. 2000. Philcoxia: a new genus of Scrophulariaceae with three new species from eastern Brazil. Kew Bulletin 55: 155–163. Wolfe AD, Datwyler SL, Randle CP. 2002. A phylogenetic and biogeographic analysis of the Cheloneae (Scrophulariaceae) based on ITS and matK sequence data. Systematic Botany 27: 138–148. Wolfe AD, Randle CP, Datwyler SL, Morawetz JJ, Arguedas N, Diaz J. 2006. Phylogeny, taxonomic affinities, and biogeography of Penstemon (Plantaginaceae) based on ITS and cpDNA sequence data. American Journal of Botany 93: 1699–1713. 406. SCROPHULARIACEAE FIGWORT FAMILY Backlund M, Oxelman B, Bremer B. 2000. Phylogenetic relationships within the Gentianales based in ndhF and rbcL sequences, with particular

reference to the Loganiaceae. American Journal of Botany 87: 1029–1043. Carlquist SJ. 1997. Wood anatomy of Buddlejaceae. Aliso 15: 41–56. Fay MF, Bennett JR, Dixon KW, Christenhusz MJM. 2010. Parasites, their relationships and the disintegration of Scrophulariaceae sensu lato. Curtis’s Botanical Magazine 26: 286–313. Gándara E, Sosa V. 2013. Testing the monophyly and position of the North American shrubby desert genus Leucophyllum (Scrophulariaceae: Leucophylleae). Botanical Journal of the Linnean Society 171: 508–518. Henrickson J, Flyr LD. 1985. Systematics of Leucophyllum and Eremogeton (Scrophulariaceae). Sida 11: 107–172. Hilliard OM. 1994. The Manuleeae: a tribe of Scrophulariaceae. Edinburgh University Press, Edinburgh. Leeuwenberg AJM. 1979. The Loganiaceae of Africa XVIII. Buddleja L. II. Revision of the African and Asiatic species. Mededelingen van de Landbouwhogeschool te Wageningen 79: 1–163. Lersten NR, Curtis JD. 1997. Anatomy and distribution of foliar idioblasts in Scrophularia and Verbascum (Scrophulariaceae). American Journal Botany 84: 1638–1645. Moore TE, Verboom GA, Cramer MD. 2008. The adaptive significance of leaf size and shape variation in Jamesbrittenia (Scrophulariaceae: Manuleeae). South African Journal of Botany 74: 373. Norman EM. 2000. Buddlejaceae. Flora Neotropica Monograph 81: 1–225. Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisens WJ, Reeves PA. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88: 348–361. Olmstead RG, Reeves PA. 1995. Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndhF sequences. Annals of the Missouri Botanical Garden 82: 176–193. Olmstead RG. 2002. Whatever happened to the Scrophulariaceae? Fremontia 30: 13–22. Oxelman B, Backlund M, Bremer B. 1999. Relationships of the Buddlejaceae s.l. investigated using parsimony jackknife and branch support analysis of chloroplast ndhF and rbcL sequence data. Systematic Botany 24: 164–182. Oxelman B, Kornhall P, Olmstead RG, Bremer B. 2005. Further disintegration of Scrophulariaceae. Taxon 54: 411–425. Roessler H. 1979. Revision der Gattungen Hebenstretia L. und Dischisma Choisy (ScrophulariaceaeSelagineae). Mitteilungen der Botanisches Staatssammlung München 15: 1–160. Tank DC, Beardsley PM, Kelchner SA, Olmstead RG. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19: 289–307. 407. LINDERNIACEAE WISHBONE-FLOWER FAMILY Albach DC, Meudt HM, Oxelman B. 2005. Piecing together the “new” Plantaginaceae. American Journal of Botany 92: 297–315. Barringer K. 1984. Cubitanthus, a new genus of Gesneriaceae from Brazil. Journal of the Arnold Arboretum 65: 145–147. Fischer E, Schäferhoff, Müller K. 2013. The phylogeny of Linderniaceae — the new genus Linderniella, and new combinations within Bonnaya,

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FURTHER READING Craterostigma, Lindernia, Micranthemum, Torenia and Vandellia. Willdenowia 43: 209–238. Perret M, Chautems A, De Araujo AP, Salamin N. 2013. Temporal and spatial origin of Gesneriaceae in the New World inferred from plastid DNA sequences. Botanical Journal of the Linnean Society 171: 61–79. Rahmanzadeh R, Müller K, Fischer E, Bartels D, Borsch T. 2004. The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales). Plant Biology 7: 67–78. Schäferhoff B, Fleischmann A, Fischer E, Albach DC, Borsch T, Heubl G, Müller KF. 2010. Towards resolving Lamiales relationships: insights from rapidly evolving chloroplast sequences. BMC Evolutionary Biology 10: 352. Tank DC, Beardsley PM, Kelchner SA, Olmstead RG. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19: 289–307. 408. BYBLIDACEAE RAINBOWPLANT FAMILY Bruce AN. 1905. On the activities of the glands of Byblis gigantea. Notes from the Royal Botanic Garden Edinburgh 4: 9–14. Carlquist SJ. 1976. Wood anatomy of Byblidaceae. Botanical Gazette 137: 35–38. Conran JG, Christophel DC. 2004. A fossil Byblidaceae seed from Eocene South Australia. International Journal of Plant Sciences 165: 691–694. Conran JG, Dowd JM. 1993. The phylogenetic relationships of the Byblis-Roridula (ByblidaceaeRoridulaceae) complex inferred from 18S rRNA partial sequences. Plant Systematics and Evolution 188: 73–86. Conran JG, Lowrie A, Moyle-Croft J. 2002. A revision of Byblis (Byblidaceae) in southwestern Australia. Nuytsia 15: 11–20. Conran JG. 1996. The embryology and relationship of the Byblidaceae. Australian Systematic Botany 9: 243–254. Hartmeyer I, Hartmeyer S. 2005. Byblis filifolia als echte Karnivore rehabilitiert. Das Taublatt 3: 4–5. Hartmeyer S. 1997. Carnivory of Byblis revisited: a simple method for enzyme testing on carnivorous plants. Carnivorous Plant Newsletter 27: 110–113. Lloyd FE. 1942. The carnivorous plants. Chronica Botanica, Waltham, Massachusetts. Lowrie A, Conran JG. 1998. A taxonomic revision of the genus Byblis (Byblidaceae) in northern Australia. Nuytsia 12: 59–74. McPherson S. 2008. Glistening carnivores. The stickyleaved insect-eating plants. Redfern Natural History Productions, Poole. Müller K, Borsch T, Legendre L, Porembski S, Theisen I, Barthlott W. 2004. Evolution of carnivory in Lentibulariaceae and the Lamiales. Plant Biology 6: 477–490. Płachno BJ, Adamec L, Lichtscheidl IK, Peroutka M, Adlassnig W, Vrba J. 2006. Fluorescence labelling of phosphatase activity in digestive glands of carnivorous plants. Plant Biology 8: 813–820. Rani RS. 1994. Floral anatomy and the affinities of Byblidaceae. Rheedea 4: 144–150. 409. STILBACEAE CANDLESTICKS FAMILY Backlund M, Oxelman B, Bremer B. 2000. Phylogenetic relationships within the Gentianales based in ndhF and rbcL sequences, with particular reference to the Loganiaceae. American Journal of

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Botany 87: 1029–1043. Bremer B, Olmstead RG, Struwe L, Sweere JA. 1994. rbcL sequences support exclusion of Retzia, Desfontainia and Nicodemia from the Gentianales. Plant Systematics and Evolution 190: 213–230. Carlquist SJ. 1986. Wood anatomy of Stilbaceae and Retziaceae: ecological and systematic implications. Aliso 11: 299–316. Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisens WJ, Reeves PA. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88: 348–361. Oxelman B, Kornhall P, Olmstead RG, Bremer B. 2005. Further disintegration of Scrophulariaceae. Taxon 54: 411–425. Raj B. 1993. A contribution to the pollen morphology of Stilbaceae Kunth. Pollen and Spores 25: 395–408. Rourke JP. 2000. A review of generic concepts in the Stilbaceae. Bothalia 30: 9–15. Wagstaff SJ, Olmstead RG. 1997. Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Systematic Botany 22: 165–179. 410. MARTYNIACEAE UNICORNPLANT FAMILY Bretting PK, Nilsson S. 1988. Pollen morphology of the Martyniaceae and its systematic implications. Systematic Botany 13: 51–59. Carlquist SJ. 1987. Wood anatomy of Martyniaceae and Pedaliaceae. Aliso 11: 473–483. Gutierrez R. 2008. Preliminary chloroplast DNA studies on the flowering plant family Martyniaceae (order Lamiales). Journal of the Arizona-Nevada Academy of Science 40: 105–110. Mameli E. 1916. Ricerche anatomiche, fisiologische e biologische sulla Martynia lutea Lindl. Atti dell’Instituto Botanico della Università di Pavia, series 2, 16: 137–188. Phillippi A, Tyrl RJ. 1979. The reproductive biology of Proboscidea louisianica (Martyniaceae). Rhodora 81: 345–361. Płachno BJ, Adamec L, Huet H. 2009. Mineral nutrient uptake from prey and glandular phosphatase activity as a dual test of carnivory in semidesert plants with glandular leaves suspected of carnivory. Annals of Botany 104: 649–654. Rice B. 1999. Testing the appetites of Ibicella lutea and Drosophyllum. Carnivorous Plants Newsletter 28: 40–43. Rice B. 2008. Reassessing commensal-enabled carnivory in Proboscidea and Ibicella? Carnivorous Plants Newsletter 37: 15–29. Schäferhoff B, Fleischmann A, Fischer E, Albach DC, Borsch T, Heubl G, Müller KF. 2010. Towards resolving Lamiales relationships: insights from rapidly evolving chloroplast sequences. BMC Evolutionary Biology 10: 352. Van Eseltine GP. 1929. A preliminary study of the unicorn plants (Martyniaceae). New York Agricultural Experimental Station Technical Bulletin 29: 1–41. Wallace J, McGhee R, Biology Class. 1999. Testing for carnivory in Ibicella lutea. Carnivorous Plant Newsletter 28: 49–50. 411. PEDALIACEAE SESAME FAMILY Carlquist SJ. 1987. Wood anatomy of Martyniaceae and Pedaliaceae. Aliso 11: 473–483. Ihlenfeldt HD. 2010. Pedaliaceae — evolution and phylogeny of the succulent genera. Schumannia 6: 151–182.

McKerral A. 1918. The Burmese Sesamum varieties. Notes on their variation and growth. Journal of the Asiatic Society in Bengal, new series 14: 389–398. Singh SP. 1960. Morphological studies in some members of the family Pedaliaceae — Sesamum indicum DC. Phytomorphology 10: 65–82. Straka H, Ihlenfeldt HD. 1965. Pollenmorphologie und Systematik der Pedaliaceae R.Br. Beiträge zur Biologie der Pflanzen 41: 175–207. 412. ACANTHACEAE BEAR’S-BREECHES FAMILY Balkwill MJ, Balkwill K. 1997. Delimitation and infrageneric classification of Barleria (Acanthaceae). Kew Bulletin 52: 535–573. Borg AJ, McDade LA, Schönenberger J. 2008. Molecular phylogenetics and morphological evolution of Thunbergioideae (Acanthaceae). Taxon 57: 811–822. Borg AJ, Schönenberger J. 2011. Comparative floral development and structure of the black mangrove genus Avicennia and related taxa in the Acanthaceae. International Journal of Plant Sciences 172: 330–340. Carine MA, Scotland RW. 2002. Classification of Strobilanthinae (Acanthaceae): trying to classify the unclassifiable? Taxon 51: 259–279. Carlquist SJ, Zona S. 1988. Wood anatomy of Acanthaceae: a survey. Aliso 12: 201–227. Daniel TF, McDade LA, Manktelow M, Kiel CA. 2008. The “Tetramerium lineage” (Acanthaceae: Acanthoideae: Justicieae): delimitation and intralineage relationships based on cp and nrITS sequence data. Systematic Botany 33: 416–436. Eberling HL. 1924. The origin of the Corinthian capital. The Art Bulletin 6: 75–81. Graham VAW 1988. Delimitation and infrageneric classification of Justicia (Acanthaceae). Kew Bulletin 43: 551–624. Hedrén M, Chase MW, Olmstead RG. 1995. Relationships in the Acanthaceae and related families as suggested by cladistic analysis of rbcL nucleotide sequences. Plant Systematics and Evolution 194: 93–109. Makholela TM, Balkwill K, Manning JC. 2008. Clarification of generic delimitation in Justicia and Siphonoglossa (Acanthaceae). South African Journal of Botany 74: 371–372. Manktelow M. 1996. Phaulopsis (Acanthaceae) — a monograph. Symbolae Botanicae Upsalienses 31(2): 1–184. McDade LA, Daniel TF, Kiel CA. 2008. Toward a comprehensive understanding of phylogenetic relationships among lineages of Acanthaceae s.l. (Lamiales). American Journal of Botany 95: 1136–1152. McDade LA, Daniel TF, Kiel CA., Borg AJ. 2012. Phylogenetic placement, delimitation, and relationships among genera of the enigmatic Nelsonioideae (Lamiales: Acanthaceae). Taxon 61: 637–651. McDade LA, Daniel TF, Masta SE, Riley KM. 2000. Phylogenetic relationships within the tribe Justicieae (Acanthaceae): evidence from molecular sequences, morphology, and cytology. Annals of the Missouri Botanical Garden 87: 435–458. McDade LA, Kiel C, Daniel TF, Tripp EA. 2008. Biogeography of the Acanthaceae. South African Journal of Botany 74: 358. McDade LA, Masta SE, Moody ML, Waters E. 2000. Phylogenetic relationships among Acanthaceae: evidence from two genomes. Systematic Botany 25: 106–121.

FURTHER READING McDade LA, Moody ML. 1999. Phylogenetic relationships among Acanthaceae: evidence from non-coding trnL-trnF chloroplast DNA sequences. American Journal of Botany 86: 70–80. Raj B. 1961. Pollen morphological studies in the Acanthaceae. Grana Palynologica 3: 3–108. Schwarzbach AE, McDade LA. 2002. Phylogenetic relationships of the mang rove family Avicenniaceae based on chloroplast and nuclear ribosomal DNA sequences. Systematic Botany 27: 84–98. Thulin M. 2007. Synopsis of Satanocrater (Acanthaceae). Nordic Journal of Botany 24: 385–388. Tripp EA. 2007. Evolutionary relationships within the species-rich genus Ruellia (Acanthaceae). Systematic Botany 32: 628–649. Tripp EA, Daniel TF, Fatimah S, McDade LA. 2013. Phylogenetic relationships within Ruellieae (Acanthaceae) and a revised classification. International Journal of Plant Sciences 174: 97–137. Tripp EA, Daniel TF, Lendemer JC, McDade LA. 2009. New molecular and morphological insights prompt transfer of Blechum to Ruellia (Acanthaceae). Taxon 58: 893–906. Tripp EA, Fatimah S. 2012. Comparative anatomy, morphology, and molecular phylogenetics of the African genus Satanocrater (Acanthaceae). American Journal of Botany 99: 967–982. Tripp EA, Manos PS. 2008. Is floral specialisation and evolutionary dead end? Pollination system transitions in Ruellia (Acanthaceae). Evolution 62: 1712–1737. Tripp EA, McDade LA. 2014. A rich fossil record yields calibrated phylogeny for Acanthaceae (Lamiales) and evidence for marked biases in timing and directionality of intercontinental disjunctions. Systematic Biology 63: 660–684. Witztum A, Schulgasser K. 1995. The mechanics of seed expulsion in Acanthaceae. Journal of Theoretical Biology 176: 531–542. 413. BIGNONIACEAE TRUMPETVINE FAMILY Chen ST, Guan KY, Zhou ZK, Olmstead R, Cronk QCB. 2005. Molecular phylogeny of Incarvillea (Bignoniaceae) based on ITS and trnL-F sequences. American Journal of Botany 92: 625–633. Gasson P, Dobbins DR. 1991. Wood anatomy of the Bignoniaceae, with a comparison of trees and lianas. IAWA Bulletin, new series 12: 389–417. Gentry AH. 1974. Flowering phenology and diversity in tropical Bignoniaceae. Biotropica 6: 64–68. Gentry AH. 1992. A synopsis of Bignoniaceae ethnobotany and economic botany. Annals of the Missouri Botanical Garden 79: 53–64. Gentry AH. 1990. Evolutionary patterns in Neotropical Bignoniaceae. Memoirs of the New York Botanical Garden 55: 118–129. Grose SW, Olmstead RG. 2007. Taxonomic revisions in the polyphyletic genus Tabebuia s.l. (Bignoniaceae). Systematic Botany 32: 660–670. Kaehler M, Michelangeli FA, Lohmann LG. 2012. Phylogeny of Lundia (Bignoniaceae) based on ndhF and PepC sequences. Taxon 61: 368–380. Li J. 2008. Phylogeny of Catalpa (Bignoniaceae) inferred from sequences of chloroplast ndhF and nuclear ribosomal DNA. Journal of Systematics and Evolution 46: 341–348. Lohmann LG, Bell CD, Calió MF, Winkworth RC. 2013. Pattern and timing of biogeographical history in the Neotropical tribe Bignonieae (Bignoniaceae). Botanical Journal of the Linnean Society 171: 154–170.

Lohmann LG. 2006. Untangling the phylogeny of Neotropical lianas (Bignonieae, Bignoniaceae). American Journal of Botany 93: 304–318. Lohmann LG, Taylor CM. 2014. A new generic classification of tribe Bignonieae (Bignoniaceae). Annals of the Missouri Botanical Garden 99: 348–489. Olmstead R. 2013. Phylogeny and biogeography of Solanaceae, Verbenaceae and Bignoniaceae: a comparison of continental and intercontinental diversification patterns. Botanical Journal of the Linnean Society 171: 80–102. Olmstead RG, Zjhra ML, Lohmann LG, Grose SO, Eckert AJ. 2009. A molecular phylogeny and classification of Bignoniaceae. American Journal of Botany 96: 1731–1743. Pace MR, Angyalossy V. 2013. Wood anatomy and evolution: a case study in the Bignoniaceae. International Journal of Plant Sciences 174: 1014–1048. Suryakanta. 1973. Pollen morphological studies in the Bignoniaceae. Journal of Palynology 9: 45–82. 414. LENTIBULARIACEAE BLADDERWORT FAMILY Adamec L. 2007. Oxygen concentrations inside the traps of the carnivorous plants Utricularia and Genlisea (Lentibulariaceae). Annals of Botany 100: 849–856. Chase MW, Christenhusz MJM, Sanders D, Fay MF. 2010. Murderous plants: Victorian Gothic, Darwin and modern insights into vegetable carnivory. Botanical Journal of the Linnean Society 161: 329–356. Cieslak T, Polepalli JS, White A, Müller K, Borsch T, Barthlott W, Steiger J, Marchant A, Legendre L. 2005. Phylogenetic analysis of Pinguicula (Lentibulariaceae): chloroplast DNA sequences and morphology support several geographically distinct radiations. American Journal of Botany 92: 1723–1736. Degtjareva GV, Casper J, Hellwig F, Schmidt AR, Steiger J, Sokoloff DD. 2006. Morphology and nrITS phylogeny of the genus Pinguicula L. (Lentibulariaceae), with special attention to embryo evolution. Plant Biology 8: 778–790. Fischer E, Porembski S, Barthlott W. 2000. Revision of the genus Genlisea (Lentibulariaceae) in Africa and Madagascar with notes on ecology and phytogeography. Nordic Journal of Botany 20: 291–318. Fleischmann A, Rivadavia F, Gonella PM, Heubl G. 2011. A revision of Genlisea subgenus Tayloria (Lentibulariaceae). Phytotaxa 33: 1–40. Greilhuber J, Borsch T, Müller K, Worberg A, Porembski S, Barthlott W. 2006. Smallest angiosperm genome found in Lentibulariaceae, with chromosomes of bacterial size. Plant Biology 8: 770–777. Jobson RW, Playford J, Cameron KM, Albert VA. 2003. Molecular phylogenetics of Lentibulariaceae inferred from plastid rps16 intron and trnL-F DNA sequences: implictations for character evolution and biogeography. Systematic Botany 28: 157–171. Legendre L. 2002. The genus Pinguicula L. (Lentibulariaceae): an overview. Acta Botanica Gallica 141: 77–95. Lloyd FE. 1942. The carnivorous plants. Chronica Botanica, Waltham. Müller K, Borsch T, Legendre L, Porembski S, Barthlott W. 2006. Recent progress in understanding the evolution of carnivorous Lentibulariaceae (Lamiales). Plant Biology 8: 748–757.

Müller K, Borsch T, Legendre L, Porembski S, Theisen I, Barthlott W. 2004. Evolution of carnivory in Lentibulariaceae and the Lamiales. Plant Biology 6: 477–490. Müller K, Borsch T. 2005. Phylogenetics of Utricularia (Lentibulariaceae) and molecular evolution of the trnK intron in a lineage with high substitution rates. Plant Systematics and Evolution 250: 39–67. Płachno BJ, Kozieradzka-Kiszkurno M, Swiatek P. 2007. Functional ultrastructure of Genlisea (Lentibulariaceae) digestive hairs. Annals of Botany 100: 195–203. Reifenrath K, Theisen I, Schnitzler J, Porembski S, Barthlott W. 2006. Trap architecture in carnivorous Utricularia (Lentibulariaceae). Flora 201: 597–605. Reut MS. 1993. Trap structure of the carnivorous plant Genlisea (Lentibulariaceae). Botanica Helvetica 103: 101–111. Richards JH. 2001. Bladder function in Utricularia purpurea (Lentibulariaceae): is carnivory important? American Journal of Botany 88: 170–176. Rodondi G, Beretta M, Andreis C. 2010. Pollen morphology of alpine butterworts (Pinguicula L., Lentibulariaceae). Review of Palaeobotany and Palynology 162: 1–10. Seine R, Porembski S, Balduin M, Theisen I, Wilbert N, Barthlott W. 2002. Different prey strategies of terrestrial and aquatic species in the carnivorous genus Utricularia (Lentibulariaceae). Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 124: 71–76. Sirová D, Borovec J, Šantrůčková H, Šantrůček J, Vrba J, Adamec L. 2010. Utricularia carnivory revisited: plants supply photosynthetic carbon to traps. Journal of Experimental Botany 61: 99–103. Taylor, P. 1989. The genus Utricularia. Kew Bulletin Additional Series XIV. HMSO, London. 415. SCHLEGELIACEAE HIGUERITO FAMILY Barringer K. 2004. A revision of Gibsoniothamnus L.O.Williams (Schlegeliaceae). Brittonia 56: 213–237. Burger WC, Barringer K. 2000. Schlegeliaceae Reveal. Flora Costaricensis. Fieldiana Botany, new series 41: 69–77. Gasson P, Dobbins DR. 1991. Wood anatomy of the Bignoniaceae, with a comparison of trees and lianas. IAWA Bulletin, new series 12: 389–417. Gentry AH. 1974. Flowering phenology and diversity in tropical Bignoniaceae. Biotropica 6: 64–68. Gentry AH. 1980. Bignoniaceae-part I (Crescentieae and Tourrettieae). Flora Neotropica Monograph 25. 416. THOMANDERSIACEAE WEST-AFRICANBITTERBUSH FAMILY Bremekamp CEB. 1942. The position of the genus Thomandersia Baill. Recueil des Travaux Botaniques Néerlandais 39: 166–175. Heine H. 1966. Révision du genre Thomandersia Baill. (Acanthaceae). Bulletin du Jardin botanique de l’État à Bruxelles 36: 207-248. Ngadjui BT, Dongo E, Ayafor JF, Connolly JD. 1994. Thomandertriol, a triterpenoid from the twigs of Thomandersia laurifolia. Journal of Natural Products 57: 161–163. Sreemadhavan CP. 1977. Diagnosis of some new taxa and some new combinations in Bignoniales. Phytologia 37: 413–416, Wortley AH, Harris DJ, Scotland RW. 2007. On the taxonomy and phylogenetic position of Thomandersia. Systematic Botany 32: 415–444.

Plants of the World

745

FURTHER READING Wortley AH, Scotland RW, Rudall PJ. 2005. Floral anatomy of Thomandersia (Lamiales), with particular reference to the nature of the retinaculum and extranuptial nectaries. Botanical Journal of the Linnean Society 149: 469–482. 417. VERBENACEAE VERVAIN FAMILY Cantino PD. 1992. Evidence for a polyphyletic origin of the Labiatae. Annals of the Missouri Botanical Garden 79: 361–379. Lu-Irving P, Olmstead RG. 2013. Investigating the evolution of Lantaneae (Verbenaceae) using multiple loci. Botanical Journal of the Linnean Society 171: 103–119. Lu-Irving P, O’Leary N, O’Brien A, Olmstead RG. 2014. Resolving the genera Aloysia and Acantholippia within tribe Lantaneae (Verbenaceae), using chloroplast and nuclear sequences. Systematic Botany 39: 644–655. Marx HE, O’Leary N, Yuan YW, Lu-Irving P, Tank DC, Múlgura ME, Olmstead RG. 2010. A molecular phylogeny and classification of Verbenaceae. American Journal of Botany 97: 1647–1663. O’Leary N, Calviño CI, Martínez S, Lu-Irving P, Olmstead RG, Múlgara ME. 2012. Evolution of morphological traits in Verbenaceae. American Journal of Botany 99: 1778–1792. O’Leary N, Yuan YW, Chemisquy A, Olmstead RG. 2009. Reassignment of species of paraphyletic Junellia s.l. to the new genus Mulguraea (Verbenaceae) and new circumscription of genus Junellia: molecular and morphological congruence. Systematic Botany 34: 777–786. Olmstead RG. 2013. Phylogeny and biogeography of Solanaceae, Verbenaceae, and Bignoniaceae: a comparison of continental and intercontinental diversification patterns. Botanical Journal of the Linnean Society 171: 80–102. Peralta P, Múlgara ME, Denham SS, Botta SM. 2008. Revisión del género Junellia (Verbenaceae). Annals of the Missouri Botanical Garden 95: 338–390. Sanders RW. 2001. The genera of Verbenaceae in the southeastern United States. Harvard Papers in Botany 5: 303–358. Thode VA, O’Leary N, Olmstead RG, Freitas LB. 2013. Phylogenetic position of the monotypic genus Verbenoxylum (Verbenaceae) and new combination under Recordia. Systematic Botany 38: 805–817. Wagstaff SJ, Olmstead RG. 1997. Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Systematic Botany 22: 165–179. Yuan YW, Olmstead RG. 2008. A species-level phylogenetic study of the Verbena complex (Verbenaceae) indicates two independent intergeneric chloroplast transfers. Molecular Phylogenetics and Evolution 48: 23–33. 418. LAMIACEAE MINT FAMILY Bendiksby M, Thorbek L, Scheen AC, Lindqvist C, Ryding O. 2011. An updated phylogeny and classification of Lamiaceae subfamily Lamioideae. Taxon 60: 471–484. Bramley GLC, Forest F, De Kok RPJ. 2009. Troublesome tropical mints: re-examining generic limits of Vitex and relations (Lamiaceae) in Southeast Asia. Taxon 58: 500–510. Bräuchler C, Meimberg H, Heubl G. 2010. Molecular phylogeny in Menthinae (Lamiaceae, Nepetoideae, Mentheae) — taxonomy, biogeo-graphy and conflicts. Molecular Phylogenetics and Evolution 55: 501–523. Cantino PD, Sanders RW. 1986. Subfamilial

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Christenhusz, Fay & Chase

classification of Labiatae. Systematic Botany 11: 163–185. Conn BJ, Brown EA, Henwood MJ, Olmstead RG. 2009. Infrageneric phylogeny of Chloantheae (Lamiaceae) based on chloroplast ndhF and nuclear ITS sequence data. Australian Systematic Botany 22: 243–256. Drew BT, Sytsma KJ. 2012. Phylogenetics, biogeography, and staminal evolution in the tribe Mentheae (Lamiaceae). American Journal of Botany 99: 933–953. Grayer RJ. Eckert MR, Veitch NC, Kite GC, Marin PD, Kokubun T, Simmonds MSD, Paton AJ. 2003. The chemotaxonomic significance of two bioactive caffeic acid esters, nepetoidins A and B, in the Lamiaceae. Phytochemistry 64: 519–528. Harley RM, Reynolds T (eds). 1992. Advances in Labiatae science. Royal Botanic Gardens, Kew. Harley RM, Pastore JFB. 2012. A generic revision and new combinations in the Hyptidinae (Lamiaceae), based on molecular and morphological evidence. Phytotaxa 58: 1–55. Li B, Cantino PD, Olmstead RG, Bramley GLC, Xiang CL, Ma ZH, Tan YH, Zhang DX. 2016. A largescale chloroplast phylogeny of the Lamiaceae sheds new light on its subfamilial classification. Scientific Reports 6: 34343. Moon HK, Smets E, Huysmans S. 2010. Phylogeny of the tribe Mentheae (Lamiaceae): the story of molecules and micromorphological characters. Taxon 59: 1065–1076. Paton AJ, Springate D, Suddee S, Otieno D, Grayer RJ, Harley MM, Willis F, Simmonds MSJ, Powell MP, Savolainen V. 2004. Phylogeny and evolution of basils and allies (Ocimeae, Labiatae) based on three plastid DNA regions. Molecular Phylogenetics and Evolution 31: 277–299. Salmaki Y, Zarre S, Ryding O, Scheunert A, Bräuchler C, Heubl G. 2012. Phylogeny of the tribe Phlomideae (Lamioideae: Lamiaceae) with special focus on Eremostachys and Phlomoides: new insights from nuclear and chloroplast sequences. Taxon 61: 161–197. Salmaki Y, Zarre S, Ryding O, Lindqvist C, Bäuchler C, Heubl G, Barber J, Bendiksby M. 2013. Molecular phylogeny of tribe Stachydeae (Lamiaceae subfamily Lamioideae). Molecular Phylogenetics and Evolution 69: 535–551. Scheen AC, Bendiksby M, Ryding O, Mathiesen C, Albert VE, Lindqvist C. 2010. Molecular phylogenetics, character evolution, and suprageneric classification of Lamioideae (Lamiaceae). Annals of the Missouri Botanical Garden 97: 191–217. Steane DA, Mabberley DJ. 1998. Rotheca (Lamiaceae) revived. Novon 8: 204–206. Steane DA, De Kok RPJ, Olmstead RG. 2004. Phylogenetic relationships between Clerodendrum (Lamiaceae) and other ajugoid genera inferred from nuclear and chloroplast DNA sequence data. Molecular Phylogenetics and Evolution 32: 39–45. Tewari DN. 1992. A monograph on teak (Tectona grandis Linn.f.). International Book Distributors, Dehra Dun. Wagstaff SJ, Olmstead RG. 1997. Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Systematic Botany 22: 165–179. Walker JB, Sytsma KJ. 2007. Staminal evolution in the genus Salvia (Lamiaceae): molecular phylogenetic evidence for multiple origins of the staminal level. Annals of Botany 100: 375–391. Walker JB, Sytsma KJ, Treutlein J, Wink M. 2004. Salvia (Lamiaceae) is not monophyletic:

implications for the systematics, radiation, and ecological specializations of Salvia and tribe Mentheae. American Journal of Botany 91: 1115–1125. Wester P, Claßen-Bockhoff R. 2011. Pollination syndromes of New World Salvia species with special reference to bird pollination. Annals of the Missouri Botanical Garden 98: 101–155. Whitten WM. 1981. Pollination ecology of Monarda didyma, M. clinopodia, and hybrids (Lamiaceae) in the southern Appalachian Mountains. American Journal of Botany 68: 435–442. Wilson TC, Conn BJ, Henwood MJ. 2012. Molecular phylogeny and systematics of Prostanthera (Lamiaceae). Australian Systematic Botany 25: 341–352. Wink M. 2003. Evolution of secondary metabolites from an ecological and molecular phylogenetic perspective. Phytochemistry 64: 3–19. Yuan YW, Mabberley DJ, Steane DA, Olmstead RG. 2010. Further disintergration and redefinition of Clerodendrum (Lamiaceae): implications for understanding the evolution of an intriguing breeding strategy. Taxon 59: 125–133. 419. MAZACEAE CUPFLOWER FAMILY Argue CL. 1984. Pollen morphology in Dodartia, Lancea, Leucocarpus, and Mazus and an analysis of pollen morphotypes in the Mimuleae (Scrophulariaceae). Canadian Journal of Botany 62: 1287–1297. Deng T, Zhang XS, Kim C, Zhang JW, Zhang DG, Volis S. 2016. Mazus sunhangii (Mazaceae), a new species discovered in central China appears to be highly endangered. PLoS ONE 11: e0163581. 420. PHRYMACEAE LOPSEED FAMILY Argue CL. 1980. Pollen morphology in the genus Mimulus (Scrophulariaceae) and its taxonomic significance. American Journal of Botany 67: 68–87. Argue CL. 1984. Pollen morphology in Dodartia, Lancea, Leucocarpus, and Mazus and an analysis of pollen morphotypes in the Mimuleae (Scrophulariaceae). Canadian Journal of Botany 62: 1287–1297. Barker WR, Nesom GL, Beardsley PM, Fraga NS. 2012. A taxonomic conspectus of Phrymaceae: a narrowed circumscription for Mimulus, new and resurrected genera, and new names and combination. Phytoneuron 2012-39: 1–60. Beardsley PM, Barker WR. 2005. Patterns of evolution in Australian Mimulus and related genera (Phrymaceae-Scrophulariaceae): a molecular phylogeny using chloroplast and nuclear sequence data. Australian Systematic Botany 18: 61–73. Beardsley PM, Olmstead RG. 2002. Redefining Phrymaceae: the placement of Mimulus, tribe Mimuleae, and Phryma. American Journal of Botany 89: 1093–1102. Beardsley PM, Schoenig S, Whittall JB, Olmstead RG. 2004. Patterns of evolution in western North American Mimulus (Phrymaceae). American Journal of Botany 91: 474–489. Bradshaw HD Jr, Wilbert SM, Otto KG, Schemske DW. 1995. Genetic mapping of floral traits associated with reproductive isolation in monkeyflowers (Mimulus). Nature 376: 762–765. Grant AL. 1924. A monograph of the genus Mimulus. Annals of the Missouri Botanical Garden 11: 99–389. Holm T. 1913. Phryma leptostachya L., a morphological study. Botanical Gazette 56: 306–318.

FURTHER READING Nie ZL, Sun H, Beardsley PM, Olmstead RG, Wen J. 2006. Evolution of biogeographic disjunction between eastern Asia and eastern North America in Phryma (Phrymaceae). American Journal of Botany 93: 1343–1356. Oxelman B, Kornhall P, Olmstead RG, Bremer B. 2005. Further disintegration of Scrophulariaceae. Taxon 54: 411–425. Schemske DW, Bradshaw HD. 1999. Pollinator preference and evolution of floral traits in monkeyflowers (Mimulus). Proceedings of the National Academy of Sciences of the USA 96: 11910–11915. Tank DC, Beardsley PM, Kelchner SA, Olmstead RG. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19: 289–307. Whittall JB, Carlson ML, Beardsley PM, Meinke RJ, Liston A. 2006. The Mimulus moschatus alliance (Phrymaceae): molecular and morphological phylogenetics and their conservation implications. Systematic Botany 31: 380–397. 421. PAULOWNIACEAE EMPRESS-TREE FAMILY Campbell DH. 1930. The relationships of Paulownia. Bulletin of the Torrey Botanical Club 57: 47–50. Erbar C, Gülden C. 2011. Ontogeny of the flowers in Paulownia tomentosa — a contribution to the recognition of the resurrected monogeneric family Paulowniaceae. Flora 206: 205–218. Hu SY. 1958. A monograph of the genus Paulownia. Quarterly Journal of the Taiwan Museum 12: 1–54. Kirkham T, Fay MF. 2009. Paulownia kawakamii. Curtis’s Botanical Magazine 26: 111–119. Maheshwari JK. 1961. The genus Wightia Wall. in India with a discussion on its systematic position. Bulletin of the Botanical Survey of India 3: 31–35. Smiley CJ. 1961. A record of Paulownia in the Tertiary of North America. American Journal of Botany 48: 175–179. Steenis CGGJ van. 1949. Notes on the genus Wightia (Scrophulariaceae). Bulletin du Jardin Botanique de Buitenzorg, séries III, 18: 213–227. Wei Z. 1989. Pollen morphology of Wightia and its taxonomic significance. Acta Metallurgica Sinica 11: 1–3. 422. OROBANCHACEAE BROOMRAPE FAMILY Albach DC, Li HQ, Zhao N, Jensen SR. 2007. Molecular systematics and phytochemistry of Rehmannia (Scrophulariaceae). Biochemical Systematics and Ecology 35: 293–300. Albach DC, Yan K, Rosendal Jensen SR, Li HQ. 2009. Phylogenetic placement of Triaenophora (formerly Scrophulariaceae) with some implications for the phylogeny of Lamiales. Taxon 58: 749–756. Bennett JR, Mathews S. 2006. Phylogeny of the parasitic plant family Orobanchaceae inferred from phytochrome A. American Journal of Botany 93: 1039–1051. Bolliger M. 1992. Monographie der Gattung Odontites (Scrophulariaceae) sowie der verwandten Gattungen Macrosyringion, Odontitella, Bornmuellerantha und Bartsiella. Willdenowia 26: 37–168. Dong LN, Wang H, Wortley AH, Lu L, Li DZ. 2013. Phylogenetic relationships in the Pterygiella complex (Orobanchaceae) inferred from molecular and morphological evidence. Botanical Journal of the Linnean Society 171: 491–507. Ernst WR. 1972. Floral morphology and systematics of Lamourouxia (Scrophulariaceae: Rhinanthoideae). Smithsonian Contributions of Botany 6: 1–63.

Fay MF, Bennett JR, Dixon KW, Christenhusz MJM. 2010. Parasites, their relationships and the disintegration of Scrophulariaceae sensu lato. Curtis’s Botanical Magazine 26: 286–313. Gussarova G, Popp M, Vitek E, Brochmann C. 2008. Molecular phylogeny and biogeography of the bipolar Euphrasia (Orobanchaceae): recent radiations in an old genus. Molecular Phylogenetics and Evolution 48: 444–460. Hjertson ML. 1995. Taxonomy, phylogeny and biogeography of Lindenbergia (Scrophulariaceae). Botanical Journal of the Linnean Society 119: 265–321. Li XD, Zan YY, Li JQ, Yang SZ. 2008. A numerical taxonomy of the genera Rehmannia and Triaenophora (Scrophulariaceae). Journal of Systematics and Evolution 46: 730–737. Manen JF, Habashi C, Jeanmonod D, Park JM, Schneeweiss GM. 2004. Phylogeny and intraspecific variability of holoparasitic Orobanche (Orobanchaceae) inferred from plastid rbcL sequences. Molecular Phylogenetics and Evolution 33: 482–500. McNeal JR, Bennett JR, Wolfe AD, Mathews S. 2013. Phylogeny and origins of holoparasitism in Orobanchaceae. American Journal of Botany 100: 971–983. Morawetz JJ, Randle CP, Wolfe AD. 2010. Phylogenetic relationships within the tropical clade of Orobanchaceae. Taxon 59: 416–426. Morawetz JJ, Wolfe AD. 2009. Assessing the monophyly of Alectra and its relationship to Melasma (Orobanchaceae). Systematic Botany 34: 561–569. Musselman LJ, Dickison WC. 1975. The structure and development of the haustorium in parasitic Scrophulariaceae. Botanical Journal of the Linnean Society 70: 183–212. Musselman LJ. 1980. The biology of Striga, Orobanche, and other root-parasitic weeds. Annual Review of Phytopathology 18: 463–489. Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisens WJ, Reeves PA. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88: 348–361. Oxelman B, Kornhall P, Olmstead RG, Bremer B. 2005. Further disintegration of Scrophulariaceae. Taxon 54: 411–425. Park JM, Manen JF, Colwell AE, Schneeweiss GM. 2008. A plastid gene phylogeny of the non-photosynthetic parasitic Orobanche (Orobanchaceae) and related genera. Journal of Plant Research 121: 365–376. Park JM, Manen JF, Schneeweiss GM. 2007. Horizontal gene transfer of a plastid gene in the non-photosynthetic flowering plants Orobanche and Phelipanche (Orobanchaceae). Molecular Phylogenetics and Evolution 43: 974–985. Press MC, Phoenix GK. 2005. Effects of climate change on parasitic plants: the root hemiparasitic Orobanchaceae. Folia Geobotanica 40: 205–216. Rix M. 1987. The genus Rehmannia. Plantsman 8: 193–195. Schneeweiss GM, Weiss H. 2003. Polyploidy in Aeginetia indica L. (Orobanchaceae). Cytologia 68: 15–17. Tank DC, Beardsley PM, Kelchner SA, Olmstead RG. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19: 289–307. Tank DC, Egger JM, Olmstead RG. 2009. Phylogenetic classification of subtribe Castillejinae (Orobanchaceae). Systematic Botany 34: 182–197.

Tank DC, Olmstead RG. 2009. The evolutionary origin of a second radiation of annual Castilleja (Orobanchaceae) species in South America: the role of long distance dispersal and allopolyploidy. American Journal of Botany 96: 1907–1921. Těšitel J, Říha O, Svobodová Š, Malinová T, Štech M. 2010. Phylogeny, life history evolution and biogeography of the rhinanthoid Orobanchaceae. Folia Geobotanica 45: 347– 367. Wimpee CF, Wrobel RL, Garin DK. 1991. A divergent plastid genome in Conopholis americana, an achlorophyllous parasitic plant. Plant Molecular Biology 17: 161–166. Wolfe AD, Randle CP, Liu L, Steiner KE. 2005. Phylogeny and biogeography of Orobanchaceae. Folia Geobotanica 40: 115–134. Wolfe AD, Randle CP. 2001. Relationships within and among species of the holoparasitic genus Hyobanche (Orobanchaceae) inferred from ISSR banding patterns and nucleotide sequences. Systematic Botany 26: 120–130. Xia Z, Wang YZ, Smith JF. 2009. Familial placement and relations of Rehmannia and Triaenophora (Scrophulariaceae s.l.) inferred from five gene regions. American Journal of Botany 96: 519–530. Young ND, Steiner KE, dePamphilis CW. 1999. The evolution of parasitism in Scrophulariaceae/ Orobanchaceae: plastid gene sequences refute an evolutionary transition series. Annals of the Missouri Botanical Garden 86: 876–893. Zhang RX, Li MX, Jia ZP. 2008. Rehmannia glutinosa: review of botany, chemistry and pharmacology. Journal of Ethnopharmacology 117: 199–214. 423. STEMONURACEAE BUFF-BEECH FAMILY Byng JW, Bernardini B, Joseph JA, Chase MW, Utteridge TMA. 2014. Phylogenetic relationships of Icacinaceae, focusing on the vining genera. Botanical Journal of the Linnean Society 176: 277–294. Kårehed J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88: 2259–2274. Lens F, Kårehed J, Baas P, Jansen S, Rabaey D, Huysmans S, Hamann T, Smets E. 2008. The wood anatomy of the polyphyletic Icacinaceae s.l., and their relationships within asterids. Taxon 57: 525–552. Schori M, Lowry PP II, Schatz GE. 2013. A revision of the genus Grisollea (Stemonuraceae). Systematic Botany 38: 497–506. 424. CARDIOPTERIDACEAE CITRONELLA FAMILY Byng JW, Bernardini B, Joseph JA, Chase MW, Utteridge TMA. 2014. Phylogenetic relationships of Icacinaceae, focusing on the vining genera. Botanical Journal of the Linnean Society 176: 277–294. Kårehed J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88: 2259–2274. Kong DR, Peng H, Liang HX. 2002. A new type of embryo sac in Cardiopteris and its systematic implication. Acta Botanica Sinica 44: 496–498. Kong DR, Schori M, Lu SG, Li L, Peng H. 2014. Floral development of Cardiopteris, with emphasis on gynoecial structure and ovular morphology. Journal of Systematics and Evolution 52: 629–642. Lens F, Kårehed J, Baas P, Jansen S, Rabaey D, Huysmans S, Hamann T, Smets E. 2008. The wood anatomy of the polyphyletic Icacinaceae

Plants of the World

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FURTHER READING s.l., and their relationships within asterids. Taxon 57: 525–552. Tobe H. 2011. Floral structure of Cardiopteris (Cardiopteridaceae) with special emphasis on the gynoecium: systematic and evolutionary implications. Journal of Plant Research 125: 261–369. 425. PHYLLONOMACEAE FLOWERING-LEAF FAMILY Dickinson TA, Sattler R. 1974. Development of the epiphyllous inflorescence of Phyllonoma integerrima (Turcz.) Loes.: implications for comparative morphology. Botanical Journal of the Linnean Society 69: 1–13. Mori SA, Kallunki JA. 1977. A revision of the genus Phyllonoma (Grossulariaceae). Brittonia 29: 69–83. Rusby HH. 1905. Phyllonomaceae. North American Flora 22: 191. Tobe H. 2013. Floral morphology and structure of Phyllonoma (Phyllonomaceae): systematic and evolutionary implications. Journal of Plant Research 126: 709–718.

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revisited with molecular data. Kew Bulletin 55: 341–347. Selbach-Schnadelbach A, Cavalli S, Manen JF, Coelho GC, Teixeira de Souza-Chies T. 2009. New information for Ilex phylogenetics based on the plastid psbA-trnH integenic spacer (Aquifoliaceae). Botanical Journal of the Linnean Society 159: 182–193. Tsang ACW, Corlett RT. 2005. Reproductive biology of the Ilex species (Aquifoliaceae) in Hong Kong, China. Canadian Journal of Botany 83: 1645–1654.

426. HELWINGIACEAE FLOWERING-RAFTS FAMILY Dickinson TA, Sattler R. 1975. Development of the epiphyllous inflorescence of Helwingia japonica (Helwingiaceae). American Journal of Botany 62: 962–973. Hara H, Kurosawa S. 1975. A revision of the genus Helwingia. Bulletin, University Museum, University of Tokyo 8: 393–413. Iwashina T, Kamenosono K, Hatta H. 1997. Flavonoid glycosides of Aucuba japonica and Helwingia japonica (Cornaceae): phytochemical relationship with the genus Cornus. Journal of Japanese Botany 72: 337–346.

428. ROUSSEACEAE PUTAWETA FAMILY Gustafsson MHG, Bremer K. 1997. The circumscription and systematic position of Carpodetaceae. Australian Systematic Botany 10: 855–862. Hansen DM, Müller CB. 2009. Invasive ants disrupt gecko pollination and seed dispersal of the endangered plant Roussea simplex in Mauritius. Biotropica 41: 202–208. Hansen DM. 2005. Pollination of the enigmatic Mauritian endemic Roussea simplex (Rousseaceae): birds or geckos? Ecotropica 11: 69–72. Lundberg J, Bremer K. 2003. A phylogenetic study of the order Asterales using one morphological and three molecular data sets. International Journal of Plant Sciences 164: 553–578. Lundberg J. 2001. The asteralean affinity of the Mauritian Roussea (Rousseaceae). Botanical Journal of the Linnean Society 137: 267–276. Magdalena C, Couch C, Tredwell E. Roussea simplex. ht t p://w w w.kew.org /science - conser vat ion / plants-fungi/roussea-simplex Praglowski J, Grafström E. 1985. The genus Carpodetus (Escalloniaceae): a pollen morphological enigma. Grana 24: 11–21.

427. AQUIFOLIACEAE HOLLY FAMILY Baas P. 1975. Vegetative anatomy and the affinities of Aquifoliaceae, Sphenostemon, Phelline, and Oncotheca. Blumea 22: 311–407. Copeland HF. 1964. Structural notes on hollies (Ilex aquifolium and I. cornuta, family Aquifoliaceae). Phytomorphology 13: 455–464. Cuénoud P, Del Pero Martinez MA, Loizeau PA, Spichiger R, Andrews S, Manen JF. 2000. Molecular phylogeny and biogeography of the genus Ilex L. (Aquifoliaceae). Annals of Botany 85: 111–122. Galle FC. 1997. Hollies: the genus Ilex. Timber Press, Portland. González AM, Tarragó JR. 2009. Anatomical structure and secretion compounds in nine Ilex species from southern South America. Botanical Journal of the Linnean Society 160: 197–210. Lee DE, Lee WG, Mortimer N. 2001. Where and why have all the flowers gone? Depletion and turnover in the New Zealand Cenozoic angiosperm flora in relation to palaeogeography and climate. Australian Journal of Botany 49: 341–356. Manen JF, Barriera G, Loizeau PA, Naciri Y. 2010. The history of extant Ilex species (Aquifoliaceae): evidence of hybridisation within a Miocene radiation. Molecular Phylogenetics and Evolution 57: 961–977. Martin HA. 1977. The history of Ilex (Aquifoliaceae) with special reference to Australia: evidence from pollen. Australian Journal of Botany 25: 655–673. Powell M, Savolainen V, Cuénoud P, Manen JF, Andrews S. 2000. The mountain holly (Nemopanthus mucronatus: Aquifoliaceae)

429. CAMPANULACEAE BELLFLOWER FAMILY Antonelli A. 2008. Higher level phylogeny and evolutionary trends in Campanulaceae subfam. Lobelioideae: molecular signal overshadows morphology. Molecular Phylogenetics and Evolution 46: 1–18. Antonelli A. 2009. Have giant lobelias evolved several times independently? Life form shifts and historical biogeography of the cosmopolitan and highly diverse subfamily Lobelioideae (Campanulaceae). BMC Biology 7: 82. Batterman MRW, Lammers TG. 2004. Branched foliar trichomes of Lobelioideae (Campanulaceae) and the infrageneric classification of Centropogon. Systematic Botany 29: 448–458. Carlquist S. 1962. Ontogeny and comparative anatomy of thorns of Hawaiian Lobeliaceae. American Journal of Botany 49: 413–419. Carlquist S. 1969. Wood anatomy of Lobelioideae (Campanulaceae). Biotropica 1: 47–72. Carolin RC. 1967. The concept of the inflorescence in the order Campanulales. Proceedings of the Linnean Society of New South Wales 92: 7–26. Chapman JL. 1966. Comparative palynology in Campanulaceae. Transactions from the Kansas Academy of Sciences 69: 197–200. Cosner ME, Jansen RK, Lammers TG. 1994. Phylogenetic relationships in the Campanulales based on rbcL sequences. Plant Systematics and Evolution 190: 79–95. Cupido CN, Prebble JM, Eddie WMM. 2013. Phylogeny of southern African and Australasian wahlenbergioids (Campanulaceae) based on ITS and trnL-F sequence data: implications for a

Christenhusz, Fay & Chase

reclassification. Systematic Botany 38: 523–535. Dupont YL, Hansen DM, Rasmussen JT, Olesen JM. 2004. Evolutionary changes in nectar sugar composition associated with switches between bird and insect pollination: the Canarian birdflower element revisited. Functional Ecology 18: 670–676. Eddie WM, Shulkina T, Gaskin JF, Haberle RC, Jansen RK. 2004. The phylogeny of Campanulaceae s. str. inferred from ITS sequences of nuclear ribosomal DNA. Annals of the Missouri Botanical Garden 90: 544–576. Erbar C, Leins P. 1989. On the early floral development and the mechanisms of secondary pollen presentation in Campanula, Jasione and Lobelia. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 111: 29–55. Givnish TJ, Millam KJ, Mast AR, Paterson TB, Theim TJ, Hipp AJ, Henss JM, Smith JF, Woods KR, Sytsma KJ. 2009. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales: Campanulaceae). Proceedings of the Royal Society B 276: 407–416. Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic relationships of the Asteraceae Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). American Journal of Botany 82: 250–265. Haberle R, Dang A, Lee T, Peñaflor C, Cortes-Burns H, Oestreich A, Raubeson L, Cellinese N, Edwards EJ, Kim ST, Eddie WMM, Jansen RK. 2009. Taxonomic and biogeographic implications of a phylogenetic analysis of the Campanulaceae based on three chloroplast genes. Taxon 58: 715–734. Hansen DM, Beer K, Müller CB. 2006. Mauritian coloured nectar no longer a mystery: a visual signal for lizard pollinators. Biology Letters 2: 165–168. Hong DY. 1995. The geography of the Campanulaceae: on the distribution centres. Acta Phytotaxonomica Sinica 33: 521–536. Knox EB, Muasya AM, Muchhala N. 2008. The predominantly South American clade of Lobeliaceae. Systematic Botany 33: 462–468. Lancucka-Srodoniowa M. 1977. New herbs described from the Tertiary of Poland. Acta Palaeobotanica 18: 37–44. Olesen JM, Rønsted N, Tolderlund U, Cornett C, Mølgaard P, Madsen J, Jones CG, Olsen CE. 1998. Mauritian red nectar remains a mystery. Nature 393: 529. Olesen JM, Alarcón M, Ehlers BK, Aldasoro JJ, Roquet C. 2012. Pollination, biogeography and phylogeny of oceanic island bellf lowers (Campanulaceae). Perspectives in Plant Ecology, Evolution and Systematics 14: 169–182. Roquet C, Sanmartín I, García-Jacas N, Sáez L, Susanna A, Wikström H, Aldasoro JJ. 2009. Reconstructing the history of Campanulaceae s. str. with a Bayesian approach to molecular dating and dispersal-vicariance analysis. Molecular Phylogenetics and Evolution 52: 575–587. Tank DC, Donoghue MJ. 2010. Phylogeny and phylogenetic nomenclature of the Campanulidae based on an expanded sample of genes and taxa. Systematic Botany 35: 425–441. Tobe H, Morin NR. 1996. Embryology and circumscription of Campanulaceae and Campanulales: a review of the literature. International Journal of Plant Research 109: 425–435.

FURTHER READING 430. PENTAPHRAGMATACEAE SCORPION’STAIL FAMILY Carlquist S. 1997. Pentaphragma: a unique wood and its significance. IAWA Bulletin, new series 18: 3–12. Dunbar A. 1978. Pollen morphology and taxonomic position of the genus Pentaphragma Wall. (Pentaphragmataceae). Grana 17: 141–147. Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic relationships of the Asteraceae Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). American Journal of Botany 82: 250–265. Kapil RN, Vijayaraghavan MR. 1965. Embryology of Pentaphragma horsfieldii (Miq.) Airy Shaw with a discussion of the systematic position of the genus. Phytomorphology 15: 93–102. Vogel S. 1998. Remarkable nectaries: structure, ecology, organophyletic perspectives. IV. Miscellaneous cases. Flora 193: 225–248. 431. STYLIDIACEAE TRIGGERPLANT FAMILY Armbruster WS, Edwards ME, Debevec EM. 1994. Character displacement generates assemblage structure of western Australian triggerplants (Stylidium). Ecology 75: 315–329. Bronckers F, Steiner F. 1972. A l’étude morphologique du pollen de la familles des Stylidiaceae. Grana 12: 1–22. Carlquist S. 1969. Studies in Stylidiaceae: new taxa, field observations, evolutionary tendencies. Aliso 7: 13–64. Carlquist S. 1981. Types of cambial activity and wood anatomy of Stylidium (Stylidiaceae). American Journal of Botany 68: 778–785. Carlquist S, Lowrie A. 1989. Two new species of Stylidium from Western Australia. Phytologia 67: 368–376. Carolin RC. 1960. Floral structure and anatomy in the family Stylidiaceae Swartz. Proceedings of the Linnean Society of New South Wales 85: 189–196. Darnowski DW, Carroll DM, Płachno B, Kabanoff E, Cinnamon E. 2006. Evidence of protocarnivory in triggerplants (Stylidium spp.; Stylidiaceae). Plant Biology 8: 805–812. Erbar C. 1992. Floral development of two species of Stylidium (Stylidiaceae) and some remarks on the systematic position of the family Stylidiaceae. Canadian Journal of Botany 70: 258–271. Erickson R. 1958. Triggerplants. Paterson Brokensha, Perth. Findlay GP, Findlay N. 1989. The structure of the column in Stylidium. Australian Journal Botany 37: 81–101. Glenny D. 2009. A revision of the genus Forstera (Stylidiaceae). New Zealand Journal of Botany 47: 285–315. Laurent N, Bremer B, Bremer K. 1998. Phylogeny and generic interrelationships of the Stylidiaceae (Asterales), with a possible extreme case of floral paedomorphosis. Systematic Botany 23: 289–304. Wagstaff SJ, Wege J. 2002. Patterns of diversification in New Zealand Stylidiaceae. American Journal of Botany 89: 865–874. 432. ALSEUOSMIACEAE TOROPAPA FAMILY Burtt BL. 1949. Studies in the Ericales, IX. The taxonomic position of Wittsteinia. Kew Bulletin 3: 493–495. Dickison WC. 1986. Wood anatomy and affinities of the Alseuosmiaceae. Systematic Botany 11: 214–221.

Dickison WC. 1989. Stem and leaf anatomy of the Alseuosmiaceae. Aliso 12: 567–578. Gardner RO. 1978. The species of Alseuosmia (Alseuosmiaceae). New Zealand Journal of Botany 16: 271–277. Kårehed J, Lundberg J, Bremer B, Bremer K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae and Phellinaceae. Systematic Botany 24: 660–682. Lundberg J, Bremer K. 2003. A phylogenetic study of the order Asterales using one morphological and three molecular data sets. International Journal of Plant Sciences 164: 553–578. Van Steenis CGGJ. 1984. A synopsis of Alseuosmiaceae in New Zealand, New Caledonia, Australia, and New Guinea. Blumea 29: 387–394. 433. PHELLINACEAE CORKFRUIT FAMILY Baas P. 1975. Vegetative anatomy and the affinities of Aquifoliaceae, Sphenostemon, Phelline, and Oncotheca. Blumea 22: 311–407. Kårehed J, Lundberg J, Bremer B, Bremer K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae and Phellinaceae. Systematic Botany 24: 660–682. Lundberg J, Bremer K. 2003. A phylogenetic study of the order Asterales using one morphological and three molecular datasets. International Journal of Plant Sciences 164: 553–578. 434. ARGOPHYLLACEAE SILVERLEAF FAMILY Eyde RH. 1966. Systematic evolution of the flower and fruit of Corokia. American Journal of Botany 53: 833–847. Kårehed J, Lundberg J, Bremer B, Bremer K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae and Phellinaceae. Systematic Botany 24: 660–682. Lobova TA. 1997. Seed morphology and anatomy in the genera Argophyllum and Corokia (Argophyllaceae). Botanicheskii Zhurnal 82: 68–78. Webb CJ. 1994. Pollination, self-incompatibility, and fruit production in Corokia cotoneaster (Escalloniaceae). New Zealand Journal of Botany 32: 385–392. Zemann M. 1907. Studien zu einer Monographie der Gattung Argophyllum Forst. Annalen des Naturhistorichen Museums, Wien 22: 270–292. 435. MENYANTHACEAE BOGBEAN FAMILY Armstrong JE. 2002. Fringe science: are the corollas of Nymphoides (Menyanthaceae) flowers adapted for surface tension interactions? American Journal of Botany 89: 362–365. Bohm BA, Nicholls KW, Ornduff R. 1986. Flavonoids of the Menyanthaceae: intra- and interfamilial relationships. American Journal of Botany 73: 204–213. Chuang TI, Ornduff R. 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79: 1396–1406. Dulberger R, Ornduff R. 2000. Stigma morphology in distylous and non-heterostylous species of Villarsia (Menyanthaceae). Plant Systematics of Evolution 25: 171–184. Erbar C. 1997. Fieberklee unde Seekanne - Enzianoder Aster-verwandt? Zur Blütenentwicklung und systematischen Stellung der Menyanthaceae. Botanische Jahrbücher f ür Systematik , Pflanzengeschichte und Pflanzengeographie 119: 115–135.

Maheshwari Devi H. 1962. Embr yological studies in Gentianaceae (Gentianoideae and Menyanthoideae). Proceedings of the Indian Academy of Sciences B, 56: 195–216. Ornduff R. 1966. The origin of dioecism from heterostyly in Nymphoides (Menyanthaceae). Evolution 20: 309–314. Richards JH, Dow M, Troxler T. 2010. Modeling Nymphoides architecture: a morphological analysis of Nymphoides aquatica (Menyanthaceae). American Journal of Botany 97: 1761–1771. Tippery NP, Les DH. 2008. Phylogenetic analysis of the internal transcribed spacer (ITS) region in Menyanthaceae using predicted secondary stucture. Molecular Phylogenetics and Evolution 49: 526–537. Tippery NP, Les DH. 2011. Phylogenetic relationships and morphological evolution in Nymphoides (Menyanthaceae). Systematic Botany 36: 1101–1113. Tippery NP, Les DH, Padgett DJ, Jacobs SWL. 2008. Generic circumscription in Menyanthaceae: a phylogenetic evaluation. Systematic Botany 33: 598–612. 436. GOODENIACEAE FANFLOWER FAMILY Carlquist S. 1969. Wood anatomy of Goodeniaceae and the problem of insular woodiness. Annals of the Missouri Botanical Garden 56: 358–390. Carolin RC. 1959. Floral structure and anatomy in the family Goodeniaceae Dumort. Proceedings of the Linnean Society of New South Wales 84: 252–255. Carolin RC. 1966. Seeds and fruit of the Goodeniaceae. Proceedings of the Linnean Society of New South Wales 91: 58–83. Carolin RC. 1978. The systematic relationship of Brunonia. Brunonia 1: 9–29. Carolin RC, Rajput MTM, Morrison D. 1992. Goodeniaceae. Flora of Australia 35: 4–351. Cave RL, Birch CJ, Hammer GL, Erwin JE, Johnston ME. 2010. Floral ontogeny in Brunonia australis (Goodeniaceae) and Calandrinia speciosa (Portulacaceae). Australian Journal of Botany 58: 61–69. Erbar C, Leins P. 1988. Studien zur Blütenentwicklung und Pollenpräsentation bei Brunonia australis Smith (Brunoniaceae). Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 110: 263–282. Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic relationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). American Journal of Botany 82: 250–265. Gustafsson MHG, Backlund A, Bremer B. 1996. Phylogeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae. Plant Systematics and Evolution 199: 217–242. Gustafsson MHG, Grafström E, Nilsson S. 1997. Pollen morphology of the Goodeniaceae and comparisons with related families. Grana 36: 185–207. Hansen HV. 1997. Studies in the Goodeniaceae and the Brunoniaceae with a discussion of their relationships to Asteraceae and Calyceraceae. Nordic Journal of Botany 17: 495–510. Howarth DG, Gustafsson MHG, Baum DA, Mortley TJ. 2003. Phylogenetics of the genus Scaevola (Goodeniaceae): implications for dispersal patterns across the Pacific Basin and colonisation of the Hawaiian Islands. American Journal of Botany 90: 915–923.

Plants of the World

749

FURTHER READING Jabaily RS, Shepherd KA, Gustafsson MHG, Sage LW, Krauss SL, Howarth DG, Motley TJ. 2012. Systematics of the Austral-Pacific family Goodeniaceae: establishing a taxonomic and evolutionary framework. Taxon 61: 419–436. Jabaily RS, Shepherd KA, Gardner AG, Gustafsson MHG, Howarth DG, Motley TJ. 2014. Historical biogeography of the predominantly Australian plant family Goodeniaceae. Journal of Biogeography 41: 2057–2067. Peacock WJ. 1963. Chromosome numbers and cytoevolution in the Goodeniaceae. Proceedings of the Linnean Society of New South Wales 88: 8–27.  437. CALYCERACEAE BALSAMLEAF FAMILY Bohm BA, Reid A, DeVore M, Stuessy TF. 1995. Flavonoid chemistry of Calyceraceae. Canadian Journal of Botany 73: 1962–1965. Bremer K, Backlund A, Sennblad B, Swenson U, Andreasen K, Hjertson M, Lundberg J, Backlund M, Bremer B. 2001. A phylogenetic analysis of 100+ genera and 50+ families of euasterids based on morphological and molecular data with notes on possible higher level morphological synapomorphies. Plant Systematics and Evolution 229: 137–169. Carlquist S, DeVore ML. 1998. Wood anatomy of Calyceraceae with reference to ecology, habit, and systematic relationships. Aliso 17: 63–76. DeVore M. 1991. The occurrence of Acicarpha tribuloides (Calyceraceae) in eastern North America. Rhodora 93: 26–35. Erbar C. 1993. Studies on the floral development and pollen presentation in Acicarpha tribuloides with a discussion of the systematic position of the family Calyceraceae. Botanische Jahrbücher f ür Systematik , Pf lanzengeschichte und Pflanzengeographie 115: 325–350. Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic relationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). American Journal of Botany 82: 250–265. Hansen HV. 1992. Studies in the Calyceraceae with a discussion of its relationship to Compositae. Nordic Journal of Botany 12: 63–75. 438. ASTERACEAE DAISY FAMILY Baldwin BG, Wessa BL, Panero JL. 2002. Nuclear rDNA evidence for major lineages of helenioid Heliantheae (Compositae). Systematic Botany 27: 161–198. Barker MS, Kane NC, Matvienko M, Kozik A, Michemore RW, Knapp SJ, Rieseberg LH. 2008. Multiple paleopolyploidizations during the evolution of the Compositae reveal parallel patterns of duplicate gene retention after millions of years. Molecular Biology and Evolution 25: 2445–2455. Barreda V, Palazzesi L, Tellería MC, Katinas L, Crisci JV. 2010. Fossil pollen indicates an explosive radiation of basal asteracean lineages and allied families during Oligocene and Miocene times in the Southern Hemisphere. Review of Palaeobotany and Palynology 160: 102–110. Barreda V, Palazzesi L, Katinas L, Crisci JV, Tellería MC, Bremer K, Passala MG, Bechis F, Corsolini R. 2012. An extinct Eocene taxon of the daisy family (Asteraceae): evolutionary, ecological and biogeographical implications. Annals of Botany 109: 127–134. Bayer RJ, Starr JR. 1998. Tribal phylogeny of the Asteraceae based on two non-coding chloroplast

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sequences, the trnL intron and trnL/trnF intergenic spacer. Annals of the Missouri Botanical Garden 85: 242–256. Bremer K. 1994. Asteraceae: cladistics and classification. Timber Press, Portland. Bremer K, Gustafsson MHG. 1997. East Gondwana ancestry of the sunflower alliance of families. Proceedings of the National Academy of Sciences of the USA 94: 9188–9190. Bremer K, Friis EM, Bremer B. 2004. Molecular phylogenetic dating of asterid flowering plants shows Early Cretaceous diversification. Systematic Biology 53: 496–505. Carlquist S. 1966. Wood anatomy of Compositae: a summary with comments on factors controlling wood evolution. Aliso 6: 25–44. Carlquist S. 1976. Tribal interrelationships and phylogeny of the Asteraceae. Aliso 8: 465–492. Cronquist AJ. 1955. Phylogeny and taxonomy of the Compositae. American Midland Naturalist 53: 478–511. Eldenäs PK, Källersjö M, Anderberg AA. 1999. Phylogenetic placement and circumscription of tribes Inuleae s.str. and Plucheeae (Asteraceae): evidence from sequences of chloroplast gene ndhF. Molecular Phylogenetics and Evolution 13: 50–58. Funk VA, Bayer RJ, Keeley S, Chan R, Watson L, Gemeinholzer B, Schilling EE, Panero JL, Baldwin BG, Garcia-Jacas NT, Susanna A, Jansen RK. 2005. Everywhere but Antarctica: using a supertree to understand the diversity and distribution of the Compositae. Biologiske Skrifter 55: 343–373. Funk VA, Susanna A, Stuessy TF, Bayer RJ (eds). 2009. Systematics, evolution, and biogeography of Compositae. International Association for Plant Taxonomy, Vienna. Gustafsson MHG, Bremer K. 1995. Morphology and phylogenetic relationships of the Asteraceae Calyceraceae, Campanulaceae, Goodeniaceae and related families (Asterales). American Journal of Botany 82: 250–265. Heywood VH, Harborne JB, Turner BL (eds).1977. The biology and chemistry of the Compositae. Academic Press, London. Hind DJN, Jeffrey C, Pope GV (eds). 1995. Advances in Compositae systematics. Royal Botanic Gardens, Kew. Hind DJN, Beentje HJ (eds). 1996. Compositae: systematics. Proceedings of the International Compositae Conference. Royal Botanic Gardens, Kew. Jansen RK, Michaels HJ, Palmer JD. 1991. Phylogeny and character evolution in the Asteraceae based on chloroplast DNA restriction site mapping. Systematic Botany 16: 98–115. Jansen RK, Palmer JD. 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proceedings of the National Academy of Sciences of the USA 84: 5818–5822. Jeffrey C. 2004. Systema compositarum (asteracearum) nova. Botanicheskii Zhurnal 89: 1817–1822. Katinas L, Crisci JV, Hoch P, Tellería MC, Apodaca MJ. 2013. Trans-oceanic dispersal and evolution of early composites (Asteraceae). Perspectives in Plant Ecology, Evolution and Systematics 15: 269–280. Kim KJ, Choi KS, Jansen RK. 2005. Two chloroplast DNA inversions originated simultaneously during the early evolution of the sunflower family (Asteraceae). Molecular Biology and Evolution 22: 1783–1792.

Li WP, Yang FS, Jivkova T, Yin GS. 2012. Phylogenetic relationships and generic delimitation of Eurasian Aster (Asteraceae: Astereae) inferred from ITS, ETS and trnL-F sequence data. Annals of Botany 109: 1341–1357. Panero JL, Funk VA. 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proceedings of the Biological Society of Washington 115: 909–922. Panero JL, Funk VA. 2008. The value of sampling anomalous taxa in phylogenetic studies: major clades of the Asteraceae revisited. Molecular Phylogenetics and Evolution 47: 757–782. Park DS, Potter D. 2013. A test of Darwin’s naturalisation hypothesis in the thistle tribe shows that close relatives make bad neighbours. Proceedings of the National Academy of Sciences of the USA 29: 17915–17920. Pelser PB, Veldkamp JF, Van der Meijden R. 2006. New combinations in Jacobaea Mill. (Asteraceae: Senecioneae). Compositae Newsletter 44: 1–11. Pelser PB, Nordenstam B, Kadereit JW, Watson LE. 2007. An ITS phylogeny of tribe Senecioneae (Asteraceae) and a new delimitation of Senecio L. Taxon 56: 1077–1104. Robinson H. 1999. Revisions of Paleotropical Vernonieae (Asteraceae). Proceedings of the Biological Society of Washington 112: 220–247. Robinson H. 1999. Generic and subtribal classification of American Vernonieae. Smithsonian Contributions in Botany 89: i–iv, 1–116. Stuessy TF, Spooner DM. 1988. The adaptive and phylogenetic significance of receptacular bracts in the Compositae. Taxon 37: 114–126. Thomas MM, Rudall PJ, Ellis AG, Savolainen V, Glover BJ. 2009. Development of a complex floral trait: the pollinator-attracting petal spots of the beetle daisy, Gorteria diffusa (Asteraceae). American Journal of Botany 96: 2184–2196. Timme RE, Simpson BB, Linder CR. 2007. High resolution phylogeny for Helianthus (Asteraceae) using the 18S-26S ribosomal DNA external transcribed spacer. American Journal of Botany 94: 1837–1852. Torices R. 2010. Adding time-calibrated branch lengths to the Asteraceae supertree. Journal of Systematic Evolution 48: 271–278. Vallès J, Torrell M. Garnatje T, Garcia-Jacas N, Vilatersana R, Susanna A. 2003. The genus Artemisia and its allies: phylogeny of the subtribe Artemisiinae (Asteraceae, Anthemidae) based on nucleotide sequences of nuclear ribosomal DNA internal transcribed spacers (ITS). Plant Biology 5: 274–284. Vallès J, Canela MA, Garcia S, Hidalgo O, Pellicer J, Sánchez-Jiménez I, Siljak-Yakovlev S, Vitales D, Garnatje T. 2013. Genome size variation and evolution in the family Asteraceae. Caryologia 66: 221–235. 439. ESCALLONIACEAE CURRYPLANT FAMILY Bremer K, Friis EM, Bremer B. 2004. Molecular phylogenetic dating of asterid flowering plants shows Early Cretaceous diversification. Systematic Biology 53: 496–505. Fernando Alonso JL, Amaya JA. 1991. Historia del nombre genérico Escallonia Mutis ex L. fil. Caldasia 16: 317–326. Hibsch-Jetter C, Soltis DE, McFarlane TD. 1997. Phylogenetic analysis of Eremosyne pectinata (Saxifragaceae s.l.) based on rbcL sequence data. Plant Systematics and Evolution 204: 225–232. Lundberg J. 2001. Phylogenetic studies in the

FURTHER READING euaster-ids II with particular reference to Asterales and Escalloniales. Acta Universitatis Upsaliensis, Uppsala. Sede SM, Dürnhöfer SI, Morello S, Zapata F. 2013. Phylogenetics of Escallonia (Escalloniaceae) based on plastid DNA sequence data. Botanical Journal of the Linnean Society 173: 442–451. Sleumer H. 1968. Die Gattung Escallonia (Saxifragaceae). Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, afdeling Natuurwetenschappen, 2e reeks 58: 1–146. Stern WL 1974. Comparative anatomy and systematics of woody Saxifragaceae. Escallonia. Botanical Journal of the Linnean Society 68: 1–20. Zapata F. 2013. A multilocus phylogenetic analysis of Escallonia (Escalloniaceae): diversification in montane South America. American Journal of Botany 100: 526–545. 440. COLUMELLIACEAE ANDEAN-HOLLY FAMILY Hasselberg GBE. 1937. Zur Morphologie des vegetativen Sprosses der Loganiaceen. Symbolae Botanicae Upsalienses 2(3): 1–170. Stern WL, Warcup JH, Eyde RH. 1969. Comparative anatomy and relationships of Columelliaceae. Journal of the Arnold Arboretum 50: 36–75. 441. BRUNIACEAE BUTTONBUSH FAMILY Carlquist S. 1978. Wood anatomy of Bruniaceae: correlations with ecology, phylogeny, and organography. Aliso 9: 323–364. Claßen-Bockhoff R. 2000. Inf lorescences in Bruniaceae, with general comments on inflorescences in woody plants. Opera Botanica Belgica 12: 5–310. Claßen-Bockhoff R, Oliver EG, Hall AV, Quint M. 2011. A new classification of the South African endemic family Bruniaceae based on molecular and morphological data. Taxon 60: 1138–1155. Hall AV. 1987. Evidence of a Cretaceous alliance for the Bruniaceae. South African Journal of Science 83: 58–59. Hall AV. 1988. Systematic palynology of the Bruniaceae. Botanical Journal of the Linnean Society 96: 285–296. Pillans NS. 1947. A revision of Bruniaceae. Journal of South African Botany 11: 121–206. Quint M, Claßen-Bockhoff R. 2006. Floral ontogeny, petal diversity and nectary uniformity in Bruniaceae. Botanical Journal of the Linnean Society 152: 459–477. Quint M, Claßen-Bockhoff R. 2006. Phylogeny of Bruniaceae based on matK and ITS sequence data. International Journal of Plant Sciences 167: 135–146. Quint M, Claßen-Bockhoff R. 2008. Ancient or recent? Insights into the temporal evolution of Bruniaceae. Organisms, Diversity and Environment 8: 193–304. 442. PARACRYPHIACEAE POSSUMWOOD FAMILY Baas P. 1975. Vegetative anatomy and the affinities of Aquifoliaceae, Sphenostemon, Phelline, and Oncotheca. Blumea 22: 311–407. Cameron KM. 2003. On the phylogenetic position of the New Caledonian endemic families Pa r a c r y ph ia c e a e, O nc ot he ca c e a e, a nd Strasburgeriaceae: a comparison of molecules and morphology. Botanical Review 68: 428–443. Carlquist S. 2012. How wood evolves: a new synthesis. Botany 90: 901–940.

Dickison WC, Baas P. 1977. The morphology and relationships of Paracryphia (Paracryphiaceae). Blumea 23: 417–438. Friis EM, Pedersen KR, Endress PK. 2013. Floral structure of extant Quintinia (Paracryphiales, Campanulids) compared with the Late Cretaceous Silvianthemum and Bertilanthus. International Journal of Plant Sciences 174: 647–664. Savinov IA. 2003. Comparative carpology of the genus Sphenostemon (Sphenostemonaceae) in the context of its taxonomy and phylogeny. Botanicheskii Zhurnal 88: 5–16. Tank DC, Donoghue MJ. 2010. Phylogeny and phylogenetic nomenclature of the Campanulidae based on an expanded sample of genes and taxa. Systematic Botany 35: 425–441. 443. ADOXACEAE ELDER FAMILY Backlund A, Bremer B. 1997. Phylogeny of the Asteridae s. str. based on rbcL sequences, with particular reference to Dipsacales. Plant Systematics and Evolution 207: 225–254. Clement WL, Arakaki M, Sweeney PW, Edwards EJ, Donoghue MJ. 2014. A chloroplast tree for Viburnum (Adoxaceae) and its implications for phylogenetic classification and character evolution. American Journal of Botany 101: 1029–1049. Clement WL, Donoghue MJ. 2011. Dissolution of Viburnum section Megalotinus (Adoxaceae) of southeast Asia and its implications for morphological evolution and biogeography. International Journal of Plant Sciences 172: 559–573. Donoghue MJ, Eriksson T, Reeves PA, Olmstead RG. 2001. Phylogeny and phylogenetic taxonomy of Dipsacales, with special reference to Sinadoxa and Tetradoxa (Adoxaceae). Harvard Papers in Botany 6: 459–479. Erbar C. 1994. Contributions to the affinities of Adoxa from the viewpoint of f loral development. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 116: 259–282. Hara H. 1983. A revision of Caprifoliaceae of Japan with reference to allied plants in other districts and the Adoxaceae. Ginkgoana 5: 1–336. Jacobs B, Donoghue MJ, Bouman F, Huysmans S, Smets E. 2008. Evolution and phylogenetic importance of endocarp and seed characters in Viburnum (Adoxaceae). International Journal of Plant Sciences 169: 409–431. Jacobs B, Huysmans S, Smets E. 2010. Evolution and systematic value of fruit and seed characters in Adoxaceae (Dipsacales) Taxon 59: 850–866. Roels P, Smets E. 1994. A comparative floral ontogenetical study between Adoxa moschatellina and Sambucus ebulus. Belgian Journal of Botany 127: 235–254. Schmerler SB, Clement WL, Beaulieu JM, Chatelet DS, Sack L, Donoghue MJ, Edwards EJ. 2012. Evolution of leaf form correlates with tropicaltemperate transitions in Viburnum (Adoxaceae). Proceedings of the Royal Society B, 279: 3853–3860. 444. CAPRIFOLIACEAE HONEYSUCKLE FAMILY Backlund A.1996. Phylogeny of the Dipsacales. Acta Universitatis Upsaliensis, Uppsala. Backlund A, Pyck N. 1998. Diervillaceae and Linnaeaceae, two new families of caprifolioids. Taxon 47: 657–661. Bell CD, Edwards EJ, Kim ST, Donoghue MJ. 2001. Dipsacales phylogeny based on chloroplast DNA

sequences. Harvard Papers in Botany 6: 481–499. Bell CD, Donoghue MJ. 2005. Dating the Dipsacales: comparing models, genes, and evolutionary implications. American Journal of Botany 92: 284–296. Benko-Iseppon AM, Morawetz W. 2000. Viburnales: cytological features and a new circumscription. Taxon 49: 5–16. Carlquist S. 1982. Wood anatomy of Dipsacaceae. Taxon 31: 443–450. Christenhusz MJM. 2013. Twins are not alone: a recircumscription of Linnaea (Caprifoliaceae). Phytotaxa 125: 25–32. Christy M. 1923. The common teasel as a carnivorous plant. Journal of Botany 61: 33–45. Jacobs B, Geuten K, Pyck N, Huysmans S, Jansen S, Smets E. 2011. Unraveling the phylogeny of Heptacodium and Zabelia (Caprifoliaceae): an interdisciplanary approach. Systematic Botany 36: 231–252. Jacobs B, Pyck N, Smets E. 2010. Phylogeny of the Linnaea clade: are Abelia and Zabelia closely related? Molecular Phylogenetics and Evolution 57: 741–752. Landrein S, Prenner G. 2013. Unequal twins? Inflorescence evolution in the twinflower tribe Linnaeeae (Caprifoliaceae s.l.). International Journal of Plant Sciences 174: 200–233. Landrein S, Prenner G, Chase MW, Clarkson JJ. 2012. Abelia and relatives: phylogenetics of Linnaeeae (Dipsacales-Caprifoliaceae s.l.) and a new interpretation of their inflorescence morphology. Botanical Journal of the Linnean Society 169: 692–713. Ogata K. 1988. Wood anatomy of the Caprifoliaceae of Japan. IAWA Bulletin new series 9: 299–316. Ogata K. 1991. Wood anatomy of Zabelia (Caprifoliaceae): evidence for generic recognition. IAWA Bulletin new series 12: 111–121. Zhang ZY, Zhou ZK, Gu ZJ. 2002. Karyomorphology of Heptacodium (Caprifoliaceae s. str.) and its phylogenetic implications. Taxon 51: 499–505. 445. PENNANTIACEAE KAIKOMAKO FAMILY Gardner RO, De Lange PJ. 2002. Revision of Pennantia (Icacinaceae), a small isolated genus of Southern Hemisphere trees. Journal of the Royal Society of New Zealand 32: 669–695. Kårehed J. 2001. Multiple origin of the tropical forest tree family Icacinaceae. American Journal of Botany 88: 2259–2274. Kårehed J. 2003. The family Pennantiaceae and its relationship to Apiales. Botanical Journal of the Linnean Society 141: 1–24. Lens F, Kårehed J, Baas P, Jansen S, Rabaey S, Huysmans S, Hamann T, Smets E. 2008. The wood anatomy of polyphyletic Icacinaceae s.l., and their relationships within asterids. Taxon 57: 525–552. 446. TORRICELLIACEAE IVY-PALM FAMILY Meller B. 2006. Comparative investigation of modern and fossil Torricellia fruits — a disjunctive element in the Miocene and Eocene of Central Europe and the U.S.A. Beiträge zur Paläontologie 30: 315–327. Philipson WR, Stone BC. 1980. The systematic position of Aralidium Miq. — a multidisciplinary study. Taxon 29: 391–416. Plunkett GM, Chandler GT, Lowry II PP, Pinney SM, Sprenkle TS. 2004. Recent advances in understanding Apiales and a revised classification. South African Journal of Botany 70: 371–381. Nicolas AN, Plunkett GM. 2014. Diversification times and biogeographic patterns in Apiales. Botanical Review 80: 30–58.

Plants of the World

751

FURTHER READING Schatz GE, Lowry II PP, Wolf AE. 1999. Endemic families of Madagascar. I. A synoptic revision of Melanophylla Baker (Melanophyllaceae). Adansonia Séries 3, 20: 233–242. 447. GRISELINIACEAE KAPUKA FAMILY Baas P, Wheeler E, Chase MW. 2000. Dicotyledonous wood anatomy and the APG system of angiosperm classification. Botanical Journal of the Linnean Society 134: 3–17. Dillon MO, Muñoz-Schick M. 1993. A revision of the dioecious genus Griselinia (Griseliniaceae) including a new species from the Atacama desert of northern Chile. Brittonia 45: 261–274. 448. PITTOSPORACEAE CHEESEWOOD FAMILY Cayzer LW, Crisp MD, Telford IRH. 2000. Revision of Pittosporum (Pittosporaceae) in Australia. Austalian Systematic Botany 13: 845–902. Cayzer LW, Crisp MD, Telford IRH. 2000. Auranticarpa, a new genus of Pittosporaceae from northern Australia. Austalian Systematic Botany 13: 903–917. Chandler GT, Plunkett GM, Pinney SM, Cayzer LW, Gemmill CEC. 2007. Molecular and morphological agreement in Pittosporaceae: phylogenetic analysis with nuclear ITS and plastid trnL-trnF sequence data. Australian Systematic Botany 20: 390–401. Gemill CEC, Allan GJ, Wagner WL, Zimmer EA. 2002. Evolution of insular Pacific Pittosporum (Pittosporaceae): origin of the Hawaiian radiation. Molecular Phylogenetics and Evolution 22: 31–42. Nicolas AN, Plunkett GM. 2014. Diversification times and biogeographic patterns in Apiales. Botanical Review 80: 30–58. 449. ARALIACEAE IVY FAMILY Chandler GT, Plunkett GM. 2004. Evolution in Apiales: nuclear and chloroplast markers together in (almost) perfect harmony. Botanical Journal of the Linnean Society 144: 123–147. Frodin DG, Lowry PP II, Plunkett GM. 2010. Scheff lera (Araliaceae): taxonomic history, overview and progress. Plant Diversity and Evolution 128: 561–595. Henwood MJ, Lu-Irving P, Perkins AJ. 2010. Can molecular systematics provide insights into aspects of the reproductive biology of Trachymene Rudge (Araliaceae)? Plant Diversity and Evolution 128: 85–110. Lowry PP II, Plunkett GM, Wen J. 2003. Generic relationships in Araliaceae: looking into the crystal ball. South African Journal of Botany 70: 382–392. Lowry PP II, Plunkett GM, Frodin DG. 2013. Revision of Plerandra (Araliaceae). I. A synopsis of the genus with an expanded circumscription and a new infrageneric classification. Brittonia 65: 42–61. Nicolas AN, Plunkett GM. 2009. The demise of subfamily Hydrocotyloideae (Apiaceae) and the re-alignment of its genera across the entire order Apiales. Molecular Phylogenetics and Evolution 53: 134–151. Nicolas AN, Plunkett GM. 2014. Diversification times and biogeographic patterns in Apiales. Botanical Review 80: 30–58. Plunkett GM, Chandler GT, Lowry PP II, Pinney SM, Sprenkle TS. 2004. Recent advances in understanding Apiales and a revised classification. South African Journal of Botany 70: 371–381. Plunkett GM, Lowry PP II. 2010. Paraphyly and polyphyly in Polyscias sensu lato: molecular

752

Christenhusz, Fay & Chase

evidence and the case for recircumscribing the “pinnate genera” of Araliaceae. Plant Diversity and Evolution 128: 23–54. Sokoloff D, Oskolski AA, Remizowa MV, Nuraliev MS. 2007. Flower structure and development in Tupidanthus calyptratus (Araliaceae): an extreme case of polymery among asterids. Plant Systematics and Evolution 268: 209–234. Valcárcel V, Fiz-Palacios O, Wen J. 2014. The origin of the early differentiation of ivies (Hedera L.) and the radiation of the Asian palmate group (Araliaceae). Molecular Phylogenetics and Evolution 70: 492–503. Wen J, Plunkett GM, Mitchell AD, Wagstaff SJ. 2000. The evolution of Araliaceae: a phylogenetic analysis based on ITS sequences of nuclear ribosomal DNA. Systematic Botany 26:144–167. 450. MYODOCARPACEAE MOUSEFRUIT FAMILY Lowry PP II. 1986. A systematic study of Delarbrea Vieill. (Araliaceae). Allertonia 4: 169–201. Oskolski AA, Lowry PP II, Richter HG. 1997. Systematic wood anatomy of Myodocarpus, Delarbrea, and Pseudosciadium (Araliaceae). Adansonia sér. 3, 19: 61–75. Plunkett GM, Chandler GT, Lowry PP II, Pinney SM, Sprenkle TS. 2004. Recent advances in understanding Apiales and a revised classification. South African Journal of Botany 70: 371–381. 451. APIACEAE CARROT FAMILY Andersson L, Kocsis M, Eriksson R. 2006. Relationships of the genus Azorella (Apiaceae) and other hydrocotyloids inferred from sequence variation in three plastid markers. Taxon 55: 270–280. Banasiak L, Piwczynski M, Ullinski T, Downie SR, Watson MF, Shakya B, Spalik K. 2013. Dispersal patterns in space and time: a case study of Apiaceae subfamily Apioideae. Journal of Biogeography 40: 1324–1335. Calviño CI, Tilney PM, Van Wyk BE, Downie SR. 2006. A molecular phylogenetic study of southern African Apiaceae. American Journal of Botany 93: 1828–1847. Calviño CI, Martínez SG, Downie SR. 2008. Morphology and biogeography of Apiaceae subfamily Saniculoideae as inferred by phylogenetic analysis of molecular data. American Journal of Botany 95: 196–214. Calviño CI, Martínez SG, Downie SR. 2010. Unraveling the taxonomic complexity of Eryngium L. (Apiaceae, Saniculoideae): phylogenetic analysis of 11 non-coding cpDNA loci corroborates rapid radiation. Plant Diversity and Evolution 128: 137–149. Magee AR, Calviño CI, Liu M, Downie SR, Tilney PM, Van Wyk BE. 2010. New tribal delimitations for the early diverging lineages of Apiaceae subfamily Apioideae. Taxon 59: 567–580. Morison R. 1672. Plantarum Umbelliferarum Distributio Nova, per Tabulas Cognationis et Affinitatis, ex Libra Naturae observata et detecta. Nicolas AN, Plunkett GM. 2009. The demise of subfamily Hydrocotyloideae (Apiaceae) and the re-alignment of its genera across the entire order Apiales. Molecular Phylogenetics and Evolution 53: 134–151. Nicolas AN, Plunkett GM. 2012. Untangling generic limits in Azorella, Laretia, and Mulinum (Apiaceae: Azorelloideae): insights from phylogenetic and biogeography. Taxon 61: 826–840.

Nicolas AN, Plunkett GM. 2014. Diversification times and biogeographic patterns in Apiales. Botanical Review 80: 30–58. Oskolski AA, Van Wyk BE. 2008. Systematic and phylogenetic value of wood anatomy in Heteromorpheae (Apiaceae, Apioideae). Botanical Journal of the Linnean Society 158: 569–583. Oskolski AA, Van Wijk BE. 2010. Wood and bark anatomy of Centella: scalariform perforation plates support an affinity with the subfamily Mackinlayoideae (Apiaceae). Plant Systematics and Evolution 289: 127–135. Spalik K, Piwczynski M, Danderson CA, KurzynaMlynik R, Bone TS, Downie SR. 2010. Amphitropic amphiantarctic disjunctions in Apiaceae subfamily Apioideae. Journal of Biogeography 37: 1977–1994. Spalik K, Banasiak Ł, Feist MAE, Downie SR. 2014. Recurrent short-distance dispersal explains wide distributions of hydrophytic umbellifers (Apiaceae tribe Oenantheae). Journal of Biogeography 41: 1559–1571.

GENERAL REFERENCES

GENERAL REFERENCES Angiosperm Phylogeny Group. 1998. An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85: 531–553. Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: 399–436. Angiosperm Phylogeny Group. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105–121. Angiosperm Phylogeny Group. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20. Baas P, Wheeler E, Chase MW. 2000. Dicotyledonous wood anatomy and the APG system of angiosperm classification. Botanical Journal of the Linnean Society 134: 3–17. Backlund A, Bremer K. 1998. To be or not to be — principles of classification and monotypic plant families. Taxon 47: 391–401. Beentje H. 2010. The Kew plant glossary, an illustrated dictionary of plant terms. Royal Botanic Gardens, Kew. Bower FO. 1908. The origin of a land flora. A theory based upon the facts of alternation. Macmillan, London. Brummitt RK. 1992. Vascular plant families and genera. Royal Botanic Gardens, Kew. Bridson D, Forman L. 1992. The herbarium handbook, revised edition. Royal Botanic Gardens, Kew. Byng JW. 2015. The flowering plants handbook: a practical guide to families and genera of the world. Plant Gateway, Hertford. http://www.plantgateway.com. Carlquist S. 1974. Island biology. Columbia University Press, New York.

Chanderbali AS, Berger BA, Howarth DG, Soltis PS, Soltis DE. 2016. Evolving ideas on the origin and evolution of flowers: new perspectives in the genomic era. Genetics 202: 1255–1265. Chase MW, Christenhusz MJM, Sanders D, Fay MFR. 2009. Murderous plants: Victorian Gothic, Darwin and modern insights into vegetable carnivory. Botanical Journal of the Linnean Society 161: 329–356. Chase MW, Reveal JL. 2009. A phylogenetic classification of the land plants to accompany APG III. Botanical Journal of the Linnean Society 161: 122–127. Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall MR, Price RA, Hills HG, Qiu Y-L, Kron KA, Rettig JH, Conti E, Palmer JD, Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD, Hedrén M, Gaut BS, Jansen RK, Kim K-J, Wimpee CF, Smith JF, Furnier GR, Strauss SH, Xiang Q-Y, Plunkett GM, Soltis PS, Swensen SM, Williams SE, Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn Jr GH, Graham SW, Barrett SCH, Dayanandan S, Albert VA. 1993. Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. Annals of the Missouri Botanical Garden 80: 528–580. Christenhusz MJM, Chase MW. 2013. Biogeographical patterns of plants in the Neotropics - dispersal rather than plate tectonics is most explanatory. Botanical Journal of the Linnean Society 170: 277–286. Christenhusz MJM, Chase MW. 2014. Trends and concepts in fern classification. Annals of Botany 113: 571–594. Christenhusz MJM, Fay MF, Chase MW. 2011. Linear sequence of vascular plants. Phytotaxa 19. Christenhusz MJM, Reveal J, Farjon A, Gardner MF, Mill RR, Chase MW. 2011. A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70.

Christenhusz MJM, Vorontsova MS, Fay MF, Chase MW. 2015. Results from an online survey of family delimitation in angiosperms and ferns: recommendations to the Angiosperm Phylogeny Group for thorny problems in plant classification. Botanical Journal of the Linnean Society 178: 501–528. Crepet WL, Nixon KC, Gandolfo MA. 2004. Fossil evidence and phylogeny: the age of major angiosperm clades based on mesofossil and macrofossil evidence form Cretaceous deposits. American Journal of Botany 91: 1666–1682. Cronquist A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York. Czaja AT. 1978 Structure of starch grains and the classification of vascular plant families. Taxon 27: 463–470. Dahlgren RMT, Clifford HT, Yeo PF. 1985. The families of the monocotyledons. Structure, evolution, and taxonomy. Springer, Berlin. Davis GL. 1966. Systematic embryology of the angiosperms. Wiley, New York. Endress PK. 1994. Diversity and evolutionary biology of tropical flowers. Cambridge University Press, Cambridge. Endress PK, Baas P, Gregory M. 2000. Systematic plant morphology and anatomy — 50 years of progress. Taxon 49: 401–434. Engler A, Prantl K (eds). 1887–1915. Die natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mirwirkung zahlreicher hervorragender Fachgelehrten. 33 parts, 23 volumes. Engelmann, Leipzig. Erdtman G. 1952. Pollen morphology and plant taxonomy. Angiosperms. Almqvist, Wiksell, Stockholm. Fay MF, Bennett JR, Dixon KW, Christenhusz MJM. 2010. Parasites, their relationships and the disintegration of Scrophulariaceae sensu lato. Curtis’s Botanical Magazine 26: 286–313. Plants of the World

753

GENERAL REFERENCES

Foster AS, Gifford EM Jr. 1974. Comparative morphology of vascular plants, 2nd edition. Freeman, San Francisco. Govaerts R (ed.) 2015. World checklist of selected plant families. Facilitated by the Royal Botanic Gardens, Kew. http:// apps.kew.org/wcsp/ (retrieved November 2015).

Kubitzki K. 1998. The families and genera of vascular plants. Volume III. Flowering plants. Monocotyledons: Lilianae (except Orchidaceae). Springer, Heidelberg (edited by Kubitzki K).

Hardy NB, Cook LG. 2012. Testing for ecological limitation of diversification: a case study using parasitic plants. American Naturalist 180: 438–449.

Kubitzki K. 1998. The families and genera of vascular plants. Volume IV. Flowering plants. Monocotyledons, Alismatanae and Commelinanae (except Gramineae). Springer, Heidelberg (edited by Kubitzki K).

Haston E, Richardson JE, Stevens PF, Chase MW, Harris DJ. 2009. The linear Angiosperm Phylogeny Group (LAPG) III: a linear sequence of the families in APG III. Botanical Journal of the Linnean Society 161: 128–131. Hertweck K, Kinney M, Stuart S, Maurin O, Mathews S, Chase MW, Gandolfo M, Pires JC. 2015. Phylogenetics, divergence times, and diversification from three genomics partitions in monocots. Botanical Journal of the Linnean Society 178: 375–393. Heywood VH. (ed.) 1978. Flowering plants of the world. Mayflower Books, New York. Heywood VH, Brummitt RK. Culham A, Seberg O. 2007. Flowering plant families of the world. Kew Publishing, Kew. Hickman JC. (ed.) 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley. In ‘t Veld T, Van der Veen F. 1981. Het groene mirakel. Ploegsma, Amsterdam. Judd WS, Campbell CS, Kellogg ES, Stevens PF, Donoghue MJ. 2015. Plant systematics, a phylogenetic approach, 4th edition. Sinauer, Sunderland. Kenrick P, Crane PR. 1997. The origin and early evolution of plants on land. Nature 389: 33-39. Kubitzki K. 1990. The families and genera of vascular plants. Volume I. Pteridophytes and gymnosperms. Springer, Heidelberg (edited by Kramer KU and Green PS). Kubitzki K. 1993. The families and genera of vascular plants. Volume II. Flowering 754

plants. Dicotyledons: magnoliid, hamamelid and caryophyllid families. Springer, Heidelberg (edited by Kubitzki K, Rohwer JG and Bittrich V).

Christenhusz, Fay & Chase

Kubitzki K. 2003. The families and genera of vascular plants. Volume V. Flowering plants. Dicotyledons: Malvales, Capparales and non-betalain Caryophyllales. Springer, Heidelberg (edited by Kubitzki K and Bayer C). Kubitzki K. 2004. The families and genera of vascular plants. Volume VI. Flowering plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales. Springer, Heidelberg (edited by Kubitzki K). Kubitzki K. 2004. The families and genera of vascular plants. Volume VII. Flowering plants. Dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae). Springer, Heidelberg (edited by Kadereit JW). Kubitzki K. 2007. The families and genera of vascular plants. Volume VIII. Flowering plants. Eudicots: Asterales. Springer, Heidelberg (edited by Kadereit JW and Jeffrey C). Kubitzki K. 2007. The families and genera of vascular plants. Volume IX. Flowering plants. Eudicots: Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae alliance, Passifloraceae alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae. Springer, Heidelberg (edited by Kubitzki K). Kubitzki K. 2011. The families and genera of vascular plants. Volume X. Flowering plants. Eudicots: Sapindales, Cucurbitales, Myrtaceae. Springer, Heidelberg (edited by Kubitzki K).

Kubitzki K. 2014. The families and genera of vascular plants. Volume XI. Flowering plants. Eudicots: Malpighiales. Springer, Heidelberg (edited by Kubitzki K). Leins P, Erbar C. 2010. Flower and fruit. Schweizerbart, Stuttgart. Maas PJM, Westra LYT. 1993. Neotropical plant families. Koeltz Scientific Books, Königstein. Mabberley DJ. 2008. Mabberley’s plant book, 3rd ed. Cambridge University Press, Cambridge. McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Marhold K, Prado J, Prud’homme van Reine WF, Smith GF, Wiersema JH, Turland NJ. 2012. International code of nomenclature for algae, fungi, and plants (Melbourne Code). Regnum Vegetabile 154. Koeltz Scientific Books, Königstein. Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J. 2000. Biodiversity hotspots for conservation priorities. Nature 403: 853–858. Niklas K. 2016. Plant evolution: an introduction to the history of life. University of Chicago Press, Chicago. Nyffeler R, Eggli U. 2010. An up-to-date familial and suprafamilial classification of succulent plants. Bradleya 28: 125–144. Quattrocchi U. 1999-2012. CRC World dictionary of plant names: common names, scientific names, eponyms, synonyms, and etymology. CRC Press, Boca Raton, London, New York, Washington. Five vols. Qiu YL, Lee JY, Bernasconi-Quadroni F, Soltis DE, Soltis PS, Zanis M, Zimmer E, Chen Z, Savolainen V, Chase MW. 1999. The earliest angiosperms: evidence from mitochondrial, plastid and nuclear genomes. Nature 402: 404–407. Qiu YL, Li L, Wang B, Xue JY, Hendry TA, Li RQ, Brown JW, Liu Y, Hudson GT, Chen ZD. 2010. Angiosperm phylogeny inferred from sequences of four mitochondrial genes. Journal of Systematics and Evolution 48: 391–425. Radcliffe-Smith A. 1998. Three-language list of botanical name components. Royal Botanic Gardens, Kew.

GENERAL REFERENCES Raghavendra AS, Sage R (eds). 2011. C4 photosynthesis and related CO2 concentrating mechanisms. Springer, Dordrecht. Reveal JL. 2010. A checklist of familial and suprafamilial names for extant vascular plants. Phytotaxa 6: 1–402. Richards AJ. 1997. Plant breeding systems. Chapman and Hall, London. Ruhfel BR, Gitzendanner MA, Soltis PS, Soltis DE, Burleigh JG. 2014. From algae to angiosperms — inferring phylogeny of green plants (Viridiplantae) from 360 plastid genomes. BMC Evolutionary Biology 14: 23. Savolainen V, Fay MF, Albach DC, Backlund A, Van der Bank M, Cameron KM, Johnson SA, Lledó MD, Pintaud JC, Powell M, Sheahan MC, Soltis DE, Soltis PS, Weston P, Whitten WM, Wurdack KJ, Chase MW. 2000. Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bulletin 55: 257–309. Simmonds NW. 1976. Evolution of crop plants. Longman, London. Simpson MG. 2006. Plant systematics. Elsevier Academic Press, London. Soltis DE, Smith SA, Cellinese N, Wurdack KJ, Tank DC, Brockington SF, RefulioRodriguez NF, Walker JB, Moore MJ, Carlsward BS, Bell CD, Latvis M, Crawley S, Black C, Diouf D, Xi Z, Rushworth CA, Gitzendanner MA, Sytsma KJ, Qiu YL, Hilu KW, Davis CC, Sanderson MJ, Beaman RS, Olmstead RG, Judd WS, Donoghue MJ, Soltis PS. 2011. Angiosperm phylogeny: 17 genes, 640 taxa. American Journal of Botany 98: 704–740. Soltis DE, Soltis PS, Endress P, Chase MW, Manchester S, Judd W, Majure L, Mavrodiev E. 2017. Phylogeny and evolution of the angiosperms. University of Chicago Press, Chicago. Soltis PS, Soltis DE, Chase MW. 1999. Angiosperm phylogeny inferred from multiple genes as a research tool for comparative biology. Nature 402: 402– 404. Stace C. 2010. New flora of the British Isles, third edition. Cambridge University Press, Cambridge.

Stearn W. 1966. Botanical Latin. History, grammar, syntax, terminology and vocabulary. Nelson, London. Stebbins GL. 1967. Variation and evolution in plants. Columbia University Press, New York. Stevens PF. 2001 onwards. Angiosperm phylogeny website. Version 12, July 2012 [and more or less continuously updated since]. http://www.mobot.org/MOBOT/ research/APweb/, accessed 30 November 2015. Sun Y, Moore MJ, Zhang S, Soltis PS, Soltis DE, Zhao T, Meng A, Li X, Li J, Wang H. 2016. Phylogenomic and structural analyses of 18 complete plastomes across all families of early-diverging eudicots, including an angiosperm-wide analysis of IR gene content evolution. Molecular Phylogenetics and Evolution 96: 93–101. Takhtajan A. 1980. Outline of the classification of the flowering plants (Magnoliophyta) Botanical Review 46: 225–359. Takhtajan A. 1997. Diversity and classification of flowering plants. Columbia University Press, New York.

plant classification in systematic arrangements in botanic gardens and herbaria. Botanical Journal of the Linnean Society 172: 127–141. Willis K, McElwain J. 2014. The evolution of plants, 2nd edition. Oxford University Press, Oxford. Zeng L, Zhang Q, Sun R, Kong H, Zhang N, Ma H. 2014. Resolution of deep angiosperm phylogeny using conserved nuclear genes and estimates of divergence times. Nature Communications 5: 4956. Zohary D, Hopf M, Weiss E. 2012. Domestication of plants in the Old World. The origin and spread of domesticated plants in south-west Asia, Europe, and the Mediterranean Basin, fourth edition. Oxford University Press, Oxford. Zomlefer W. 1994. Guide to the flowering plant families. University of North Carolina Press, Chapel Hill. Zona S, Christenhusz MJM. 2015. Littertrapping plants: filter-feeders of the plant kingdom. Botanical Journal of the Linnean Society 179: 554–586.

Taylor TN, Taylor EL, Krings M. 2008. Paleobotany: the biology and evolution of fossil plants, 2nd edition. Academic Press (Elsevier), Burlington, London, San Diego, New York. Thompson JN. The coevolutionary process. University of Chicago Press, Chicago. Thompson JN. Relentless evolution. University of Chicago Press, Chicago. Thorne RF. 1992. Classification and geography of the flowering plants. Botanical Review 58: 225–348. Voogd HH, Van der Steen JC. 1965. Plantkunde deel 1. Versluys, Amsterdam. Wanntorp L, Ronse De Craene (ed.) 2011. Flowers on the tree of life. Cambridge University Press, Cambridge. Watson L, Dallwitz MH. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval, version 22 July 2014. http://delta-intkey.com. Wearn JA, Chase MW, Mabberley DJ, Couch C. 2013. Utilising a phylogenetic Plants of the World

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INDEX This index includes scientific names of genera (in italics), sub-classes, orders, common names, useful plants and products. When a generic name is also used as a common name only the generic name is listed, unless the common name applies to more than one genus. Plant family names are indexed separately on the front and back endpapers of the book.

Aa  158 Aaronsohnia  603 Abarema  253 Abatia  329 Abdra  416 Abdulmajidia  486 Abelia  622 Abeliophyllum  546 Abelmoschus  389 abeokuta coffee  519 Abies  79–80, 529 abiu 490 Abobra  287 Abolboda  198 Abolbodoideae 198 Aboriella  275 Abrahamia  370 Abroma  389 Abronia  461 Abrophyllum  589 Abrotanella  603 Abrus  251, 258–9 absinthe  610–11, 636 Abuta  218 Abutilon  390 abutilon 393 Acacia  15, 101, 250, 253, 256, 259 Acaciella  253 Acaena  263 acai 178 Acalypha  334 Acalyphoideae 334 Acampe  155 Acamptopappus  603 Acanthella  355 Acanthephippium  155 Acanthocarpus  174 Acanthocephalus  603 Acanthocereus  469 Acanthochlamydoideae  137 Acanthochlamys  137 Acanthocladium  603 Acanthocladus  262 Acanthodesmos  603 Acanthogilia  485 Acanthoideae 564 Acantholepis  603 Acantholimon  433 Acantholippia  573 Acanthomintha  576 Acanthonema  551 Acanthopale  564 Acanthophoenix  177 Acanthophyllum  447 Acanthoprasium  576 Acanthopsis  564 Acanthorrhinum  553 Acanthoscyphus  434 Acanthosicyos  287

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Acanthospermum  603 Acanthostachys  195 Acanthostelma  564 Acanthostyles  603 Acanthosyris  427 Acanthothamnus  294 Acanthus  564–6 Acareosperma  245 Acarpha  601 Acaulimalva  390 Acca  351 Accara  351 Acentra  325 Acer  373 Aceranthus  219 Aceratium  300 Aceratorchis  158 acerola 318 Acharia  324 Achatocarpus  449 Achetaria  553 Achillea  603, 610, 613 Achimenes  551 achira 188 Achlydosa  158 Achlyphila  198 Achlys  219 Achnophora  603 Achnopogon  603 achocha 288 Achuaria  375 Achyrachaena  603 Achyranthes  450 Achyrocline  603 Achyronychia  447 Achyropappus  603 Achyropsis  450 Achyrospermum  576 Acianthera  155 Acianthus  158 Acicarpha  601 Acidanthera  167 Acidocroton  335 Acidonia  226 Acidosasa  208 Acidoton  334 Acilepidopsis  603 Acilepis  603 Acineta  155 Acioa  322 Aciotis  355 Aciphylla  632, 637 Acis  171 Acisanthera  355 Ackama  299 ackee 373–4 Acleisanthes  461 Acmadenia  375 Acmanthera  318 Acmella  603, 610 Acmispon  251

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Acmopyle  82 Acnistus  536 Acoelorrhaphe  177 Acokanthera  527 Acomis  603 Aconitella  220 Aconitum  220, 222 Aconogonon  434 Acorales  10–11, 117, 132 Acorites  117 acorns  264, 278, 280, 319, 322, 425 Acorus  117 Acosmium  251 Acostia  208 Acourtia  603 Acrachne  208 Acradenia  375 Acranthera  517 Acridocarpus  318 Acriopsis  155 Acrisione  603 Acritochaete  208 Acritopappus  603 Acrobotrys  517 Acrocarpus  253 Acroceras  208 Acrocomia  177 Acrodon  457 Acroglochin  450 Acrolophia  155 Acronema  632 Acronychia  375 Acrophyllum  299 Acropogon  390 Acrorchis  155 Acrosanthes  457 Acrostichum  51 Acrosynanthus  517 Acrothamnus  509 Acrotome  576 Acrotrema  231 Acrotriche  509 Acrymia  576 Acsmithia  299 Actaea  220, 222 Actephila  341 Actinanthella  430 Actinidia  504 Actinobole  603 Actinocarya  531 Actinocephalus  198 Actinocheita  370 Actinocladum  208 Actinodaphne  112 Actinodium  351 Actinokentia  177 Actinolema  632 Actinorhytis  177 Actinoschoenus  202 Actinoscirpus  202

Actinostachys  35–6 Actinostemma  287 Actinostemon  335 Actinostrobus  84 Actinotus  632, 637 Acunaeanthus  517 Acunniana  603 Acuston  416 Acystopteris  56 Adamantinia  155 Adansonia  390 Adelia  334 Adelobotrys  355 Adelostemma  527 Adelostigma  603 Adelphia  318 Adenandra  375 Adenanthellum  603 Adenanthera  253, 256, 258 Adenanthos  226 Adenarake  307 Adenaria  346 Adenia  327 Adenium  527 Adenoa  327 Adenocalymma  566 Adenocarpus  251 Adenocaulon  603 Adenochilus  158 Adenochloa  208 Adenocline  335 Adenocritonia  603 Adenodolichos  251 Adenoglossa  603 Adenogramma  462 Adenolobus  250 Adenoon  603 Adenopeltis  335 Adenophaedra  334 Adenophora  592–3 Adenophyllum  603 Adenopodia  253 Adenorandia  517 Adenosciadium  632 Adenosma  553 Adenostemma  603 Adenostoma  263 Adenostyles  603 Adenothamnus  603 Adesmia  251 Adiantum  50, 54 Adina  517 Adinandra  489 Adiscanthus  375 Adlumia  216 Adolphia  270 Adonidia  177 Adonis  220, 222 Adoxa  619, 620, 621 Adriana  334

Adrorhizon  155 adzuki bean  255 Aechmea  195 Aedesia  603 Aegialitidoideae 433 Aegialitis  433 Aegiceras  494 Aegilops  208, 209–10 Aeginetia  583–4 Aegiphila  576 Aegle  375 Aeglopsis  375 Aegokeras  632 Aegopodium  632, 636 Aegopogon  208 Aeluropus  208 Aenhenrya  158 Aenictophyton  251 Aenigmatanthera  318 Aenigmopteris  67 Aeollanthus  576, 578 Aeonium  240, 241 Aequatorium  603 Aerangis  155 Aeranthes  155 Aerides  155 Aerva  450 Aeschynanthus  551 Aeschynomene  251 Aesculus  373 Aetanthus  430 Aëtheolirion  180 Aetheorhiza  603 Aetheorhyncha  155 Aethephyllum  457 Aethionema  416 Aethusa  632, 637 Aëtoxylon  396 Aextoxicon  420 Afgekia  251 Afraegle  375 Aframmi  632 Aframomum  191, 193 African basil  578 African cabbage  414 African cotton  392 African honeysuckle  560 African linden  393 African mahogany  381 African mango  303 African marigold  610 African oil palm  178 African pear  490 African rice  210 African tulip tree  568 African violets  551 African yam bean  255 Afroaster  603 Afrobrunnichia  434 Afrocalathea  190 Afrocanthium  517

Afrocarpus  82 Afrocarum  632 Afrofittonia  564 Afroguatteria  106 Afrolicania  322 Afroligusticum  632 Afrolimon  433 Afropteris  52 Afrosciadium  632 Afroscirpoides  202 Afrosison  632 Afrothismia  134 Afrotrewia  334 Afrotrichloris  208 Afrotrilepis  202 Afrotysonia  531 Afrovivella  241 Afzelia  250, 258 Agalinis  583–4 Agalmyla  551 Aganisia  155 Aganonerion  527 Aganope  251 Aganosma  527 Agapanthoideae 170–1 Agapanthus  171 Agapetes  509 Agarista  509 agarwood 396 Agastache  576–8 Agastachys  226 Agasyllis  632 Agatea  325 Agathis  81 Agathisanthemum  517 Agathosma  375 Agave  173–4 agave nectar  174 Agavoideae 173–4 Agdestidoideae 459 Agdestis  459 Agelaea  296 Agelanthus  430 Agenium  208 Ageratella  603 Ageratina  603, 614 Ageratinastrum  603 Ageratum  603, 613 Agiortia  509 Aglaia  381 Aglaodorum  119 Aglaonema  119–20 Aglossorrhyncha  155 Agnesia  208 Agonandra  424 Agonis  351 Agoseris  603 Agouticarpa  517 Agrianthus  603 Agrimonia  263 Agriophyllum  450

INDEX Agropyron  208 Agropyropsis  208 Agrostemma  447 Agrostis  208, 211 Agrostistachys  334 Agrostocrinum  168 Agrostophyllum  155 Agrostopoa  208 Aguiaria  390 ahaloth 396 Ahernia  324 Aichryson  241 Aidia  517 Aidiopsis  517 Aidomene  527 Ailanthus  380 Ainsliaea  603 Ainsworthia  632 Aiouea  112 Aiphanes  177 Aira  208 Airopsis  208 Airosperma  517 Airyantha  251 Aistopetalum  299 Aizoanthemum  457 Aizoon  457 Ajania  603, 613 Ajaniopsis  603 ajmud 636 Ajuga  576, 578 Ajugoideae 575 Ajugoides  576 ajwain 636 Akania  402 Akeassia  603 akebi 217 Akebia  217 Akrosida  390 Akschindlium  251 Alafia  527 Alamania  155 Alangium  479 Alania  160 Alantsilodendron  253 Alatoseta  603 albarco 487 Alberta  517 Albertinia  603 Albertisia  218 Albidella  122 Albizia  101, 253, 259 Albraunia  553 Albuca  174 Alcantarea  195 Alcea  390 Alchemilla  15, 263, 267 Alchornea  334 Alchorneopsis  334 Aldama  603 alder buckthorn  271 Aldina  251 Aldrovanda  437 Alectra  583 Alectryon  373 alehoof 577 Aleisanthia  517 Aleisanthiopsis  517 Alepidea  632 Alepidocline  603 Alepis  430 Aletes  632 Aletris  133 Aleurites  335 Alexa  251 alexanders 635

Alexfloydia  208 Alexgeorgea  204 alfalfa  255, 257 Alfaroa  280 Alfredia  603 algerita 220 Algernonia  335 Algrizea  351 Alhagi  251, 256 Alibertia  517 Alicia  318 Aliciella  485 Aliella  603 Aligera  623 Alisma  122 Alismatales  10–11, 115, 118, 150 alison 419 Alistilus  251 alkanet 532 Alkanna  531 Alkekengi  536 Allagopappus  603 Allagoptera  177 Allamanda  527 Allanblackia  309 Allantoma  486 Allantospermum  303 Allardia  603 Alleizettella  517 Allenrolfea  450 Allexis  325 alley cropping  258 Alliaria  416 Allioideae  170–1, 173 Allionia  461 Allium  8, 15, 170–1 Allmania  450 Allmaniopsis  450 Allocassine  294 Allocasuarina  281 Allocephalus  603 Allocheilos  551 Allochrusa  447 Alloeochaete  208 Alloispermum  603 Allolepis  208 Allomaieta  355 Allomarkgrafia  527 Allomorphia  355 Alloneuron  355 Allonia  235 Allophyllum  485 Allophylus  373 Alloplectus  551 Allopterigeron  603 Allosanthus  372 Alloschemone  119 Allosidastrum  390 Allosorus  53 Allospondias  370 Allostigma  551 Allosyncarpia  351 Alloteropsis  208 Allotropa  509 Allowissadula  390 Allowoodsonia  527 Alloxylon  226 allspice  107, 535 Alluaudia  465 Alluaudiopsis  465 Almaleea  251 Almeidea  375 almond 565 almondettes 371 Almutaster  603

Alniphyllum  500 Alnus  282 Alocasia  119–20 Alococarpum  632 Aloë  168, 170 aloes  116, 168 aloewood 396 Aloiampelos  168 Aloidendron  168 Aloinopsis  457 Alomia  603 Alomiella  603 Alonsoa  555 Alopecurus  208 Alophia  167 Aloysia  573 Alphandia  335 Alphitonia  270 Alphonsea  106 alpine clubmoss  19 alpine everlasting  614 alpine pennycress  419 Alpinia  191, 193 Alpinioideae 191 Alrawia  173–4 Alseis  517 Alseodaphne  112 Alseuosmia  597 Alsobia  551 Alsodeiopsis  513 Alsomitra  287 Alsophila  39, 43 Alstonia  527 Alstroemeria  144–5 Altensteinia  158 Alternanthera  450, 452 Althaea  390 Althenia  129 Altingia  234 Altoparadisium  208 alula 593 alunqua 527 Aluta  351 Alvaradoa  364 Alvesia  576 Alvimia  208 Alvimiantha  270 Alvordia  603 Alyogyne  390 Alysicarpus  251 Alyssoides  416 Alyssopsis  416 Alyssum  416 Alyxia  527 ama-cha 474 Amaioua  517 Amalocalyx  527 Amana  150 amanatsu 378 Amanoa  341 Amaracarpus  517 Amaranthoideae  450 Amaranthus  450 Amaroria  380 Amarula 370 Amaryllidoideae 170–1 Amaryllis  171 amaryllis 171 Amasonia  576 amatungulu 527 Amauria  603 Amauriopsis  603 ambarella 371 Ambassa  603 Ambavia  106 Ambavioideae 106

Ambelania  527 Amberboa  603, 613 Ambilobea  368 Amblyanthopsis  494 Amblyanthus  494 Amblygonocarpus  253 Amblynotus  531 Amblyocarpum  603 Amblyolepis  603 Amblyopappus  604 Amblysperma  603 Ambongia  564 Amborella  16, 88 Amborellales  10–11, 88 Amborellanae 88 Amboroa  604 Ambrella  155 ambrette 392 Ambrosia  604, 614 Ambrosiinae  602 Ambrosina  119 Amburana  251 Ameghinoa  603 Amelanchier  263 Amellus  604, 613 Amentotaxus  86 America  604 American barberry  220 American blackcurrant 238 American brooklime  554 American dittany  577 Ameroglossum  558 Amesiella  155 Amesiodendron  373 amethyst hyacinth  174 Amethystea  576, 578 Amherstia  250, 256, 259 Amianthium  143 Amicia  251 Amischotolype  180 Amitostigma  158 Ammandra  177 Ammannia  346 Ammi  632, 637 Ammobium  604 Ammocharis  171 Ammochloa  208 Ammodaucus  632, 636 Ammodendron  251 Ammoides  632 Ammophila  208, 211 Ammopiptanthus  251 Ammoselinum  632 Ammosperma  416 Ammothamnus  251 Amolinia  604 Amomum  191, 193 Amomyrtella  351 Amomyrtus  351 Amoreuxia  397 Amorimia  318 Amorpha  251, 259 Amorphophallus  119–20 Amorphospermum  490 Ampelaster  604 Ampelocalamus  208 Ampelocera  272 Ampelocissus  245 Ampelodesmos  208 Ampelopsis  245 Ampelosycios  287 Ampelozizyphus  270 Amperea  334 Amphiachyris  604 Amphiasma  517

Amphiblemma  355 Amphibolia  457 Amphibolis  131 Amphicarpaea  251 Amphicarpum  208 Amphidasya  517 Amphiglossa  604 Amphilophium  567, 568 Amphimas  251 Amphipappus  604 Amphipetalum  467 Amphiphyllum  197 Amphipogon  208 Amphipterygium  370 Amphirrhox  325 Amphiscirpus  202 Amphitecna  566–7 Amphithalea  251 Amphoricarpos  604 Amphorocalyx  355 Amphorogyne  427 Amphorogyneae 427 Amritsar gum  256 Amsinckia  531 Amsonia  527 Amydrium  119 Amyema  430 Amylotheca  430 Amyrea  334 Amyris  375 Amyxa  396 Anabaena  38 Anabasis  450 Anacampseros  468 Anacamptis  158 Anacardioideae 370 Anacardium  370 Anacolosa  422 Anacyclus  604, 610, 613 Anadelphia  208 Anadenanthera  253 Anadendrum  119 Anaectocalyx  355 Anagyris  251 Anakasia  630 Anamaria  553 Anamirta  218 Ananas  195 Ananthacorus  54 Ananthura  604 Anaphalioides  604 Anaphalis  604, 613 Anaphyllopsis  119 Anaphyllum  119 Anarrhinum  553 Anarthria  202, 204 Anarthrioideae 204 Anarthrophyllum  251 Anastatica  21, 416 Anastrabe  560 Anathallis  155 Anatropanthus  527 Anaxagorea  106 Anaxagoreoideae 106 Anaxeton  604 Ancathia  604 Anchietea  325 Anchomanes  119 Anchonium  416 anchovy pear  487 Anchusa  531 Anchusella  531 Ancistrachne  208 Ancistragrostis  208 Ancistranthus  564 Ancistrocarphus  604

Ancistrocarpus  389 Ancistrocarya  531 Ancistrochilus  155 Ancistrocladus  442 Ancistrorhynchus  155 Ancistrothyrsus  327 Ancylobothrys  527 Andean wax palm  178 Anderbergia  604 Andersonia  509 Andinia  155 Andira  251 andiroba seed  381 Andrachne  341 Andradea  461 Andreadoxa  375 Andringitra  390 Androcalymma  251 Androcorys  158 Androcymbium  146 Andrographis  564 Androlepis  195 Andromeda  509 Andromischus  241 Andropogon  208, 211 Andropterum  208 Androsace  494 Androsiphonia  327 Androstachys  539 Androstephium  174 Androstoma  509 Androtium  370 Androtrichum  202 Androya  555 Andryala  604 Andrzeiowskya  416 Anechites  527 Aneilema  180 Anelsonia  416 Anemarrhena  174 Anemia  36 Anemioideae 36 Anemocarpa  604 Anemoclema  220 Anemone  220, 222 anemone  222, 474 Anemonidium  220 Anemonopsis  220 Anemopaegma  567 Anemopsis  98 Anerincleistus  355 Anetanthus  551 Anethum  632, 636 Anetium  54 Aneulophus 306 Angadenia  527 angel’s trumpets  540 Angeldiazia  604 Angelica  632, 635, 637 angelica  632, 635, 637 Angelesia  322 Angelonia  553 Angianthus  604 Anginon  632 Angiopteris  27 angiosperms  10–11, 87 Angkalanthus  564 Angolaea  311 Angophora  351 Angoseseli  633 Angostura  375 Angostylis  334 Angraecopsis  155 Angraecum  155 Anguloa  155 Angylocalyx  251

Plants of the World

757

INDEX Ania  155 Aniba  112 Anigozanthos  184 Anisacanthus  564 Anisadenia  337 anise  577, 610, 612, 635, 636 anise hyssop  577 Aniseia  534 Aniselytron  208 Anisocampium  60 Anisocapparis  413 Anisocarpus  604 Anisochaeta  604 Anisochilus  576 Anisocoma  604 Anisocycla  218 Anisodontea  390 Anisodus  536 Anisomeles  576 Anisomeria  459 Anisopappus  604 Anisopetalum  564 Anisophyllea  284 Anisopogon  208 Anisoptera  400 Anisopus  527 Anisosperma  287 Anisotes  564 Anisothrix  604 Anisotoma  527 Anisotome  633 Ankyropetalum  447 Anna  551 annatto 397 Annea  250 Annesijoa  335 Anneslea  489 Annesorhiza  633 Annickia  106 Annona  106 Annonoideae 106 annual black-eyed Susan 613 Anoda  390 Anodendron  527 Anodiscus  551 Anodopetalum  299 Anoectochilus  158 Anogeissus  346 Anogramma  52 Anomacanthus  564 Anomalluma  527 Anomatheca  167 Anomochloa  206 Anomochloöideae  206 Anomospermum  218 Anomostachys  335 Anonidium  106 Anoplocaryum  531 Anopteris  52 Anopterus  615–16 Anopyxis  305 Anotea  390 Anredera  466 Ansellia  155 ansy mustard  419 Antarctic hair grass  206, 447 Antarctic pearlwort  447 Antegibbaeum  457 Antennaria  604, 613 Anteremanthus  604 Anthaenantia  208 Anthaenantiopsis  208 Anthemis  604, 612–3

758

Anthephora  208 Anthericopsis  180 Anthericum  174 Antheroporum  251 Antherostele  517 Antherothamnus  555 Antherotoma  355 Anthobolus  424 Anthocarapa  381 Anthocercis  536 Anthochlamys  450 Anthochortus  204 Anthocleista  523 Anthodiscus  313 Anthodon  294 Anthogonium  155 Anthonotha  250 Anthopteropsis  509 Anthopterus  509 Anthorrhiza  517 Anthosachne  208 Anthospermopsis  517 Anthospermum  517 Anthostema  335 Anthotium  600 Anthotroche  536 Anthoxanthum  208 Anthriscus  633, 636 Anthurium  119–20 Anthyllis  251, 259 Antiaris  274 Antiaropsis  274 Anticharis  555 Anticlea  143 Antidaphne  427 Antidesma  340 Antidesmatoideae 340 Antigonon  435 Antillanthus  604 Antillia  604 Antimima  457 Antinoria  208 Antiotrema  531 Antiphiona  604 Antiphytum  531 Antirhea  517 Antirrhinum  553 Antistrophe  494 Antithrixia  604 Antizoma  218 Antonia  523 Antopetitia  251 Antrocaryon  370 Antrophyum  54 Anubias  119 Anvillea  604 Anzhengxia  416 Aoranthe  517 Aosa  476 Aotus  251 Apacheria  363 Apalochlamys  604 Aparisthmium  334 Apassalus  564 Apatesia  457 Apatophyllum  294 Apeiba  389 Apera  208 Apetahia  592–3 Aphaenandra  517 Aphaerema  329 Aphanactis  604 Aphanamixis  381 Aphananthe  273 Aphandra  177 Aphanelytrum  208

Christenhusz, Fay & Chase

Aphanisma  450 Aphanocalyx  250 Aphanocarpus  517 Aphanopetalum  241 Aphanopleura  633 Aphanosperma  564 Aphanostephus  604 Aphelandra  564–5 Aphelia  204 Aphloia theiformis  359 Aphragmus  416 Aphyllanthes  173–4 Aphyllanthoideae 173–4 Aphylloclados  603 Aphyllodium  251 Aphyllorchis  155 Apiales 625 Apiastrum  633 Apinagia  311 Apioideae 632 Apiopetalum  630, 633 Apios  251, 256 Apium  633, 635, 637 Aplanodes  416 Aplectrum  155 Apluda  208 Apoballis  119 Apocaulon  375 Apochiton  208 Apoclada  208 Apocopis  208 Apocynoideae 527 Apocynum  527 Apodanthera  287 Apodanthes  283 Apodasmia  204 Apodicarpum  633 Apodiscus  340 Apodolirion  171 Apodostigma  294 Apodytes  514 Apollonias  112 Apomuria  517 Aponogeton  126 Apophyllum  413 Apoplanesia  251 Apopyros  604 Aporosa  340 Aporostylis  158 Aporrhiza  373 Aposeris  604 Apostasia  154 Apostasioideae  152, 154 Apostates  604 Apowollastonia  604 Appendicula  155 Appendicularia  355 Appertiella  125 apple mango  371 apple of paradise  4, 188 applemint 577 apples  536, 541 apples of Sodom  541 Appunia  517 apricot  75, 102, 264–5 Aptandra  422 Aptandroideae 422 Aptenia  457 Apteranthes  527 Apteria  134 Apterokarpos  370 Apterosperma  496 Aptosimum  555 Apuleia  251 Apurimacia  251 Aquifoliales 585

Aquifolium  589 Aquilaria  396 Aquilegia  220, 222 arabica coffee  519 Arabidella  416 Arabidopsis  416 Arabis  416 Aracamunia  158 Arachis  251, 254 Arachniodes  66 Arachnis  155 Arachnitis  142 Arachnothryx  517 Araeococcus  195 Arafoe  633 Aragoa  553 Aralia  629–31 Aralidium  627 Aralioideae 629–30 Araliopsis  375 Arapatiella  253 araththa 193 Aratitiyopea  198 Araucaria  81 Araucariales  10–11, 81 Araujia  527 Arbelaezaster  604 Arberella  208 arborvitae 84 Arbutoideae 509 Arbutus  509 Arcangelista  218 Arceuthobium  427 Archaeamphora  501 Archaeanthus  103 Archaefructus  89 Archaeopteris  72 Archanthemis  604 Archboldiodendron  489 Archeozostera  128 Archeria  509 Archiatriplex  450 Archibaccharis  604 Archiboehmeria  275 Archidendron  253 Archidendropsis  253 Archirhodomyrtus  351 Archiserratula  604 Archontophoenix  177 Archytaea  308 Arctagrostis  208 Arctanthemum  604, 613 Arctic raspberry  266 Arctium  604, 610, 615 Arctogeron  604 Arctomecon  216 Arctophila  208 Arctopus  633 Arctostaphylos  509 Arctotheca  604 Arctotis  604, 613 Arctottonia  99 Arctoüs  509 Arcuatopterus  633 Arcyosperma  416 Arcytophyllum  517 Ardisia  494 Ardisiandra  494 Areca  99, 177, 179 Arecales 175 Arecoideae 177 Aremonia  263 Arenaria  447 Arenga  177 Arenifera  457 Arethusa  155

Arfeuillea  373 argan oil  491 Argania  490 Argemone  216 Argentipallium  604 Argocoffeopsis  517 Argomuellera  334 Argophyllum  598 Argostemma  517 Argylia  567 Argyranthemum  604, 613 Argyreia  534 Argyrocytisus  251, 259 Argyrodendron  390 Argyroderma  457 Argyroglottis  604 Argyrolobium  251 Argyroxiphium  604, 615 Argythamnia  334 Arida  604 Aridarum  119 Arillastrum  351 Ariocarpus  469 Ariopsis  119 Aripuana  522 Arisaema  119–20 Arisarum  119–21 Aristaloë  168 Aristea  167 Aristeguietia  604 Aristeoideae 167 Aristida  208 Aristidoideae 206 Aristocapsa  434 Aristogeitonia  539 Aristolochia  101 Aristolochioideae 101 Aristotelia  300 Arivela  414 Arjona  428 Armatocereus  469 Armeria  433 Armoracia  416 Arnaldoa  603 Arnebia  531 Arnhemia  396 Arnica  604, 611 arnica 611 Arnicastrum  604 Arnicratea  294 Arnocrinum  168 Arnoglossum  604 Arnoseris  604 Aroa  253 Aroideae 119 aromatic aster  614 Aronia  263 Arophyton  119 Arpitium  633 Arpophyllum  155 arracacha 635 Arracacia  633, 635 Arrhenatherum  208 Arrhenechthites  604 Arrojadoa  469 Arrojadocharis  604 arrow poison  101, 135, 218, 374 arrowleaf elephant ear  120 arrowroot  188, 190 Arrowsmithia  604 Artabotrys  106 Artanema  558 Artedia  633 Artemisia  603, 604, 610–1, 614

Artemisiella  604 Artemisiopsis  604 Arthraerva  450 Arthragrostis  208 Arthraxon  208 Arthrobotrya  66 Arthrocereus  469 Arthrochilus  159 Arthroclianthus  251 Arthrocnemum  450 Arthrophytum  451 Arthropodium  174 Arthropogon  208 Arthropteris  67–8 Arthrostemma  355 Arthrostylidium  208 Arthrostylis  202 Arthrotaxis  84 Artia  527 Artocarpus  274 Artorima  155 Arum  119–20 Aruncus  263 Arundina  155 Arundinaria  208 Arundinella  208 Arundinoideae 206 Arundo  208 Arundoclaytonia  208 Arytera  373 asafoetida 636 Asanthus  604 Asarina  553 Asaroideae 101 Asarum  101 Ascarina  114 Aschersoniodoxa  416 Aschistanthera  355 Asciadium  633 Ascidieria  155 Ascidiogyne  604 Asclepiadoideae 527 Asclepias  527 Ascolepis  202 Ascotheca  564 Asemeia  262 Asepalum  583 ash  265, 278, 300, 306, 353, 381, 546–7, 610 Ashtonia  340 Asian bell tree  568 Asian pennywort  636 Asian rice  210 Asimina  106 Asketanthera  527 Askidiosperma  204 Aspalathus  251, 256 Asparagales  10–11, 147, 150–1 Asparagoideae 173–4 Asparagus  173–4 asparagus fern  174 asparagus pea  255 Aspasia  155 Aspazoma  457 Asperuginoides  416 Asperugo  531 Asperula  517 Asphodeline  168, 170 Asphodeloideae 168 Asphodelus  168, 170 Aspicarpa  318 Aspidistra  174 Aspidocarya  218 Aspidogyne  159 Aspidopterys  318

INDEX Aspidosperma  527 Aspidostemon  112 Aspidotis  53 aspirin  266, 330, 509 Asplenioideae 55–8 Aspleniopsis  52 Asplenium  55, 57–8 Asplundia  139 Asplundianthus  604 assegai wood  477 Asta  416 Astartea  351 Astelia  161 Astephanus  527 Aster  603, 604, 610, 613, 615 Asterales 589 Asteranthe  106 Asteranthera  551 Asteranthos  486 Asteridea  604 Asteriscus  604 Asterogyne  177 Asterohyptis  576 Asteroideae 602–3 Asterolasia  375 Asteromyrtus  351 Asteropeia  444–5 Asteropsis  604 Asteropyrum  220 Asterostemma  527 Asterostigma  119 Asterothamnus  604 Asterotheca  27 Asterotrichion  390 Asthenochloa  208 Astianthus  567 Astiella  517 Astilbe  239 Astilboides  239 Astiria  390 Astomaea  633 Astonia  122 Astraea  335 Astragalus  15, 250–1, 256 Astranthium  604 Astrantia  633, 637 Astrebla  208 Astridia  457 Astripomoea  534 Astrocalyx  355 Astrocaryum  177 Astrocasia  341 Astrococcus  334 Astrodaucus  633 Astroloba  168 Astroloma  509 Astronia  355 Astronidium  355 Astronium  370 Astrophytum  469 Astrothalamus  275 Astrotricha  630 Astrotrichilia  381 Astus  351 Astydamia  633 Asyneuma  592–3 Asystasia  564–5 Atalantia  375 Atalaya  373 Atamisquea  413 Atelanthera  416 Ateleia  251 Athamanta  633, 636 Athanasia  604

Athenaea  536 Atherandra  527 Atherosperma  109 Athertonia  226 Athrixia  604 Athroisma  604 Athroostachys  208 Athyana  373 Athyrioideae 55–6, 59–60 Athyrium  55, 60 Athysanus  416 atki 494 Atkinsonia  430 Atractantha  208 Atractocarpus  517 Atractogyne  517 Atractylis  604 Atractylodes  604 Atraphaxis  435 Atrichantha  604 Atrichodendron  536 Atrichoseris  604 Atriplex  451 Atropa  536 Atropanthe  536 Atropatenia  416 Atroxima  262 Attalea  177 Attilaea  370 Atuna  322 aubergine 538 Aubletiana  334 Aubregrinia  490 Aubrevillea  253 Aubrieta  416 Aucoumea  368 Aucuba  515 Audouinia  618 Auerodendron  270 Augea  247 Augouardia  250 Augusta  517 Augustea  447 Aulacocalyx  517 Aulacospermum  633 Aulandra  490 Aulax  226 Aulea  311 Aulonemia  208 Aulosepalum  159 Aulotandra  191 Auranticarpa  629 Aurantioideae 375 Aureliana  536 Aureolaria  583–4 Aurinia  416 Australian bluebell  629 Australian frangipani  629 Australian ginger  193 Australian holly  336 Australian lime  376 Australian soap tree  271 Australina  275 Australluma  527 Australopyrum  208 Austrian foxglove  554 Austrobaileya  92 Austrobaileyales 10–11, 88, 92 Austrobaileyanae 88 Austrobrickellia  604 Austrobryonia  287 Austrobuxus  539 Austrocactus  469 Austrocedrus  84

Austrochloris  208 Austrocritonia  604 Austroderia  208 Austrodolichos  251 Austroeupatorium  604 Austrofestuca  208 Austrogramme  52 Austromatthaea  111 Austromuellera  226 Austromyrtus  351 Austropeucedanum  633 Austrosteenisia  251 Austrosynotis  604 Austrotaxus  86 Autana  311 Autranella  490 autumn crocus  146 Autumnalia  633 Auxemma  531 Auxopus  155 Avellanita  334 Avena  208, 210 Averrhoidium  373 Avicennia  563–5 Avicennioideae 564 avocado  108, 112 Avonia  468 awapuhi 193 Axinaea  355 Axinandra  356 Axiniphyllum  604 Axonopus  208, 211 Axyris  451 ayahuasca  318, 520 Ayapana  604 Ayapanopsis  604 Ayenia  389 Aynia  604 Azadirachta  381 azafancillo 335 Azalea  509 Azanza  389–90 Azara  329 azarole 266 Azilia  633 Azima  408 Azolla  38 Azolloideae 38 Azorella  629, 633, 637 Azorelloideae 632 Azorina  592–3 Aztecaster  604 Aztekium  469   Babiana  167 Babingtonia  351 baby tears  558 baby’s breath  448 bacaba 178 Baccaurea  340 Baccharis  604, 613 Baccharoides  604 Bachmannia  413 Backhousia  351 Bacopa  553 Bactris  177 bacuri 310 Badiera  262 Badilloa  604 Badula  494 Badusa  517 Baeckea  351 Baeometra  146 Baeriopsis  604 Bafodeya  322 Bafutia  604

Bagassa  274 Bahama whitewood  95 Baharuia  527 Bahia  604 Bahianthus  604 Bahiella  527 Bahiopsis  604 Baijiania  287 Baikiaea  250 Baileya  604 Baileyoxylon  324 Baillonella  490 Baillonia  573 Baimashania  416 Baissea  527 baitoa 272 Bajacalia  604 Bakerella  430 Bakeridesia  390 Bakerolimon  433 Bakoa  119 bakupari 310 Balaka  177 Balakata  335 Balanites  247 Balanophora  425 Balanops 319 balau 400 Balaustion  351 Balbisia  344 baldmoney 637 Balduina  604 Balfourodendron  375 Balgoya  262 Baliospermum  335 Ballantinia  416 Ballochia  564 Ballota  576, 578 Balmea  517 Baloghia  335 Baloskion  204 balsa wood  393 balsam apple  288 Balsamocarpon  253, 257 Balsamocitrus  375 Balthasaria  489 Baltimora  604 bambangan 371 bambara bean  255 Bambekea  287 bamboo  1, 202, 206, 211, 220 bamboo shoots  221 Bambusa  208, 211 Bambusoideae 208 Bamiania  433 Bampsia  558 banana  116, 188 banana passionfruits  328 Banara  329 baneberry 222 Banisteriopsis  318 Banksia  226 baobab  391, 393 Baphia  251 Baphiastrum  251 Baphiopsis  251 Baptisia  251, 259 Baptorhachis  208 Baratranthus  430 Barbacenia  137 Barbaceniopsis  137 Barbados cherry  318 Barbados gooseberry  470 Barbara’s buttons  614 Barbarea  416

Barbaretta  184 barberry 200 Barbeuia  455, 459 Barbieria  251 Barbosella  155 Barcella  177 Barclaya  91 Bardotia  583 Barjonia  527 Barkeria  155 Barkleyanthus  604 Barklya  250 Barleria  564–5 Barleriola  564 barley 210 Barnadesia  603 Barnadesioideae 602–3 Barnardia  174 Barnebya  318 Barnebydendron  250 Barnebyella  251 Barneoudia  220 Barnhartia  262 Barongia  351 Baronia  370 Baroniella  527 Barringtonia  486 Barrosoa  604 Barteria  327 Barthea  355 Barthlottia  555 Bartholina  159 Bartholomaea  329 Bartlettia  604 Bartlettina  604, 614 Bartonia  522 Bartsia  583 Bartsiella  583 Basananthe  327 Basedowia  604 Basella  466 Baseonema  527 Bashania  208 basil 577 basil thyme  577 Basiphyllaea  155 Basisperma  351 Basistemon  553 Baskervilla  159 Bassecoia  623 Basselinia  177 Bassia  451 bastard balm  578 bastard hemp  291 bastard marula  371 Bastardia  390 Bastardiastrum  390 Bastardiopsis  390 batako plum  330 batat  534, 535 Batemannia  155 Batesia  253 Batesimalva  390 bat-flower  135 Bathiorhamnus  270 Bathysa  517 Batis  408 Batocarpus  274 Batopedina  517 Batopilasia  604 Baudouinia  251 Bauera  299 Bauhinia  249–50, 255–7, 259 Baumia  583 bauno 371

Baxteria  175 Bay  112–13, 267 bay rum oil  353 Bayabusua  287 bayberry 279 Baynesia  527 Bdallophytum  386 beach naupaka  600 beach plum  265 bear’s breeches  565 beard lily  150 beard tongue  555 Beaucarnea  173–4 Beaufortia  351 Beaumontia  527 Beauprea  226 Beaupreopsis  226 Beautempsia  413 beauty bush  625 beautyberry 578 Bebbia  604 Beccarinda  551 Beccariophoenix  177 Bechium  604 Beckmannia  208 Beckwithia  220 Beclardia  155 Becquerelia  202 Bedfordia  604 beebalm 578 beech  277, 278 Beesia  220 beet 451 beetleweed 499 beetroot 451 Begonia  15, 291–2 Beguea  373 Behaimia  251 Behnia  174 Behuria  355 Beilschmiedia  112 Beirnaertia  218 Beiselia  368 Bejaranoa  604 Bejaria  509 Belamcanda  167 Belemia  461 Belencita  413 bell peppers  538–9 Bellendena  226 Bellendenoideae 226 Bellevalia  174 Bellida  604 Belliolum  96 Bellis  604, 613 Bellium  604 Belloa  604 Bellonia  551 bells of Ireland  578 Bellucia  355 bellwort 146 Beloglottis  159 Belonophora  517 Belosynapsis  180 Bembicia  329 Bencomia  263 Benevidesia  355 Benguellia  576 beni seed  262 Benin pepper  99 Benincasa  287 Benitoa  604 Benjaminia  553 Benkara  517 Bennettiodendron  329 Bennettitales 72

Plants of the World

759

INDEX bennettites  73 Benoicanthus  564 Benoistia  335 Bensoniella  239 Benstonea  140 Benthamia  159 Benthamiella  536 Benthamina  430 Bentinckia  177 Bentleya  629 Benzingia  155 benzoin 500 Benzonia  517 Bequaertia  294 Berardia  604 Berberidoideae 219 Berberidopsidales 420 Berberidopsis  421 Berberis  219–20 Berchemia  270 Berchemiella  270 Berenice  592–3 bergamot 378 bergamot oil  377 bergamot orange  377 Bergenia  239 Bergera  375 Bergeranthus  457 Bergerocactus  469 Bergeronia  251 Berghesia  517 Bergia  316 Berhautia  430 Berkheya  604, 613 Berlandiera  604, 613 Berlinia  250 Bermuda grass  211 Bernardia  334 Berneuxia  499 Bernouilla  390 Berroa  604 Berrya  389 Bersama  344 Berteroa  416 Bertholletia  486 Bertiera  517 Bertilia  604 Bertilianthus  618 Bertolonia  355 Bertya  335 Berula  633 Berylsimpsonia  603 Berzelia  618 Beschorneria  174 Besleria  551 Bessera  174 Bessey, Charles Edwin 552 Beta  451 betel nut  99, 178–9, 520 Bethencourtia  604 Betonica  576 betony 578 betsa-betsa 560 Betula  282 Betuloideae 282 Bewsia  208 Beyeria  335 Bhesa  316 Bhidea  208 Bhutanthera  159 Bia  334 Biarum  119 Bicuiba  102 Bidens  604, 613 Bidoupia  159

760

Biebersteinia  365 Bienertia  451 Biermannia  155 Bifora  633 Bifrenaria  155 Bigelowia  604 Bignonia  567–8 Bijlia  457 Bikinia  250 Bikkia  517 Bilacunaria  633 bilberry 509 bilimbi 297 Billardiera  629 Billbergia  195 Billburttia  633 Billia  373 Billieturnera  390 Billolivia  551 billy buttons  614 bindi 391 bindweed 533 Biophytum  297 Biovularia  570 Bipinnula  159 birch  211, 282 bird pepper  538 bird’s eye bush  308 bird’s-foot trefoil  257 bird-of-paradiseflower  185 birthwort 101 Bisboeckelera  202 Bischofia  340 biscuitroots 635 Biscutella  416 Biserrula  251 Bisglaziovia  355 Bisgoeppertia  522 Bishopalea  604 Bishopanthus  604 Bishopiella  604 Bishovia  604 Bismarckia  177 Bistella  533 bistort 436 Bistorta  434 bitinga rubber  528 bitter apple  288 bitter berry  449, 538 bitter cucumber  288 bitter orange  376 bitter root  464 bitterwood 380 Bituminaria  251 Bivinia  329 Bivonaea  416 Bixa  397 Bizonula  373 Blachia  335 black bindweed  435 black cabbage  614 black cardamom  193 black cumin  635 black ginger  193 black guava  520 black nightshade  538 black pepper  99, 494 black salsify  611 black sapote  492 black sesame  578 black turtle beans  254 black wood  492 Blackallia  271 blackberry  266, 274 blackcurrant 218

Christenhusz, Fay & Chase

black-eyed pea  255 black-eyed Susan  563, 565, 614 blackhaw 620 Blackstonia  522 bladdernut 361 Blainvillea  604 Blakea  355 Blakeanthus  604 Blakiella  604 Blanchetia  604 Blanchetiodendron  253 Blancoa  184 Blandfordia  160 blanket flowers  614 Blastemanthus  307 Blastus  355 Bleasdalea  226 Blechnoideae  55–6, 60–1 Blechnum  39, 42, 55, 61, 67 Blechum  564 bleeding heart  216 Bleekrodea  274 Blennodia  416 Blennosperma  604 Blennospora  604 Blepharandra  318 Blepharidachne  208 Blepharidium  517 Blepharipappus  604 Blepharis  564 Blepharispermum  604, 612 Blepharistemma  305 Blepharizonia  604 Blepharocalyx  351 Blepharocarya  370 Blepharodon  527 Blepharoneuron  208 Blephilia  576, 578 blessed thistle  611 Bletia  155 Bletilla  155 Blighia  373 Blighiopsis  373 blindness tree  336 Blinkworthia  534 blite 452 Blitum  451 Blomia  373 blood orange  376 bloodberry 460 bloodleaf 452 bloodroot 216 Bloomeria  174 Blossfeldia  469 Blotiella  46, 49 blue buttons  625 blue curls  578 blue daisy  614 blue diamods  419 blue dicks  174 blue gem  558 blue gum  532 blue lilly pilly  352 blue poppy  216 blue potato bush  541 blue sage  565 bluebeard 578 bluebell  6–7, 174, 629 blueberry  509–10 blueberry ash  300 bluebottle 613 blueboys 578 blue-eyed Mary  532

bluemink 614 Blumea  604, 612 Blumenbachia  476 Blumeodendron  334 Blumeopsis  604 Blutaparon  450 Blysmus  202 Blyxa  125 Bobartia  167 Bobea  517 Bobgunnia  251 Bocagea  106 Bocageopsis  106 Bocconia  215–6 Bocoa  251 Bocquillonia  334 Boea  551 Boeberastrum  604 Boeberoides  604 Boechera  416 Boehmeria  275 Boeica  551 Boelckea  553 Boenninghausenia  375 boer-bean 255 Boerhavia  461 Boerlagea  355 Boerlagella  490 Boesenbergia  192–3 bog oak  278 bog-myrtle 279 Bognera  119 Bogoria  155 Bogotá tea  498 Boholia  517 bois meduse  308 Bolandia  604 Bolandra  239 Bolanosa  604 Bolanthus  447 Bolax  633 Bolbitis  66–7 Bolboschoenus  202 boldo 111 Boldoa  461 bolo bolo  392 Boltonia  604, 613 Bolusafra  251 Bolusanthus  251 Bolusia  251 Bolusiella  155 bolwarra 105 Bomarea  144–5 Bombacoideae 390 Bombacopsis  389 Bombax  390 Bombycilaena  604 Bommeria  53 Bonamia  534 Bonania  335 Bonannia  633 Bonatea  159 bonavist bean  255 Bonellia  494 boneset  613, 615 Bonetiella  370 Bongardia  219 Bonia  208 Boninia  375 Bonnaya  558 Bonnetia  308 Bonplandia  485 Bontia  555 Bonyunia  523 boojum tree  484 Boophone  171

Boopis  601 Boquila  217 borage 531–2 Boraginales 529 Borago  531 Borassodendron  177 Borassus  177 Borderea  134 Borismene  218 borlotti beans  254 Borneacanthus  564 Borneocola  192 Borneodendron  335 Borneosicyos  287 Bornmuellera  416 Bornmuellerantha  583 borodina 538 Borodinia  416 Borodiniopsis  416 Boronella  375 Boronia  375 Borrichia  604 Borsczowia  451 Borthwickia  412 Borya  159–60 Boschia  389 Boschniakia  583 Boscia  413 Bosea  450 Bosistoa  375 Bosqueiopsis  274 Bossiaea  251 Boston fern  63 Boswellia  368 Bothriochloa  208 Bothriocline  604 Bothriospermum  531 Bothriospora  517 Botryarrhena  517 Botryophora  334 Botschantzevia  416 Bottegoa  375 bottle gourd  288, 299 bottletree 393 Boucerosia  527 Bouchardatia  375 Bouchea  573 Bouchetia  536 Bouea  370 Bougainvillea  461 boundary tree  568 Bourbon vanilla  152 Bourreria  531 Bousigonia  527 Bouteloua  208, 211 Boutiquea  106 Boutonia  564 Bouvardia  517 Bowdichia  251 Bowenia  74–5 bower plant  568 Bowiea  173–4 Bowkeria  560 Bowkerieae 560 Bowlesia  633 Bowringia  251 boxthorn 539 boxwood 541 Boyania  355 Boykinia  239 boysenberry 266 Brabejum  226 Brachanthemum  604 Brachiaria  208, 210 Brachionidium  155 Brachionostylum  604

Brachistus  536 Brachybotrys  531 Brachycaulos  239 Brachycereus  469 Brachychiton  390 Brachychloa  208 Brachyclados  603 Brachycorythis  159 Brachycylix  250 Brachyelytrum  208 Brachyglottis  604 Brachylaena  603, 612 Brachylepis  450 Brachyloma  509 Brachylophon  318 Brachynema  422 Brachyotum  355 Brachypeza  155 Brachypodium  208 Brachypus  416 Brachyscias  633 Brachyscome  604, 613 Brachysiphon  358 Brachysola  576 Brachystegia  250 Brachystele  159 Brachystelma  527 Brachystemma  447 Brachystephanus  564 Brachystigma  583 Brachythrix  604 Brachytome  517 Bracisepalum  155 bracken 48–9 Brackenridgea  307 Bradea  517 Braemia  155 Brahea  177 Brainea  155 bramble  266, 460 Brandisia  581, 583 Brandzeia  250 Brasenia  90 Brasilianthus  355 Brasilicereus  469 Brasiliocroton  335 brass buttons  614 Brassaiopsis  630 Brassavola  155 Brassia  155 Brassiantha  294 Brassica  416 Brassicales 401 Brassiophoenix  177 Braunblanquetia  553 Braunsia  457 Bravaisia  564 Braya  416 Brayopsis  416 Brazilian cress  610 Brazilian nutmeg.  112 Brazilian red-cloak  565 Brazilian sassafras  113 brazilwood 257 Brazoria  576 Brazzeia  486 brazzein 410 bread wheat  210 breadfruit 274 breadnut 274 Bredemeyera  262 Bredia  355 Bremeria  517 Brenandendron  604 Brenania  517 Brenierea  250

INDEX Breonadia  517 Breonia  517 Bretschneidera  402 Breviea  490 Brewcaria  195 Brexia  294 Brexiella  294 Brickellia  604 Brickelliastrum  604 bridal wreath  344 Bridelia  341 Bridgesia  373 Briggsiopsis  551 Brighamia  592, 593–4 brighteyes 610 Brillantaisia  564–5 Brimeura  174 Briquetia  390 Briza  208 broad bean  254, 259, 577 Brocchia  604 Brocchinia  195–6, 198, 570 broccoli 418 Brochoneura  102 Brodiaea  174 Brodiaeoideae 173–4 Brodriguesia  250 Brombya  375 Bromelia  195–6 Bromheadia  155 Bromuniola  208 Bromus  208 Brongniartia  251 Bronwenia  318 Brookea  553 brookweed 494 broom  253, 256, 258, 357, 452, 509, 510 Brosimum  274 Broughtonia  155 Broussaisia  473 Broussonetia  274 Browallia  536 Brownanthus  457 Brownea  250, 259 Browneopsis  250 Browningia  469 Brownleea  159 Brownlowia  389 Brownlowioideae 389 Brucea  380 Brugmansia  536 Bruguiera  305 Bruinsmia  500 Brunellia  301 Brunfelsia  536 Brunia  617–18 Bruniales 616 Brunnera  531 Brunnichia  435 Brunonia  599–600 Brunoniella  564 Brunsvigia  171 Brussels sprouts  401, 418 Bruxanelia  517 Brya  251, 258 Bryanthus  509 Bryaspis  251 Brylkinia  208 Bryobium  155 Bryocarpum  494 Bryomorphe  604 Bryonia  287 bryophyte 3 buah merah  140

Bubbia  96 Bucephalandra  119 Buchanania  370 Buchenavia  346 Buchholzia  413 Buchlomimus  208 Buchnera  582–3 Buchnerodendron  324 Buchtienia  159 Buckinghamia  226 Buckleya  427 Buckollia  527 Bucquetia  355 Budawangia  509 Buddha belly plant  336 Buddha’s hand  378 Buddleja  555 Buergersiochloa  208 buffaloberry 269 Bufonia  447 Buforrestia  180 bugle 578 bugloss 532 Bukiniczia  433 bukir 331 Bulbine  168, 170 Bulbinella  168 Bulbocodium  146 Bulbophyllum  15, 152, 155 Bulbostylis  202 bulgur 209 bullace 264 bullet wood  491 Bulleyia  155 Bullockia  517 bullwort 637 Bulnesia  247 bumba 256 bumble tree  413 Bunchosia  318 Bungarimba  517 Bungea  583 Bunias  416 Bunium  633, 635 Buphthalmum  604, 613 Bupleurum  633, 637 Burasaia  218 burbark 392 Burbridgea  191 Burchardia  146 Burchellia  517 Burckella  490 Burdachia  318 burdock  610, 615 Burkartia  603 Burkea  253, 256–7 Burkillanthus  375 Burkilliodendron  251 Burmannia  134 Burmannieae  134 bur-marigold 613 Burmeistera  592 Burmese grape  341 Burnatia  122 burnet 266 burnet rose  267 Burnettia  159 Burretiodendron  390 Burretiokentia  177 Bursaria  629 Bursera  368 Burttdavya  517 Burttia  296 burweed 392 busanguli 371

bush anemone  474 bush currants  355 bush honeysuckle  625 bush mallow  393 bush marguerites  613 bush pepper  505 bush tomato  539 Bussea  253 busy lizzy  480 butcher’s broom  174 Butea  251 Butia  177 Butomopsis  122–3 Butomus  123 butter bean  254 butterbur 612 buttercup 222 butterfly bush  557 butterfly vine  336 butternut 280 butternut squash  287 butterwort 570 Buttonia  583 Buxales 227 Buxus  228 Byblis  559, 596 Byrsanthus  329 Byrsonima  318 Byrsonimoideae 318 Byrsophyllum  517 Bystropogon  576 Bythophyton  558 Byttneria  389 Byttnerioideae 389   caa-ehe 611 Caamembeca  262 Caatinganthus  604 cabbage  178, 418 cabbage on a stick  593 cabbage tree  174 cabello de angel  289 Cabobanthus  604 Cabomba  90 Cabralea  381 Cabreriella  604 Cacalia  603–4, 608 Cacaliopsis  604 Caccinia  531 cachaça 211 Cachrys  633 Cacosmia  604 cactus 470 cactus fruit  470 Cadaba  413 Cadellia  260 Cadia  251 Cadiscus  604 cadushi 470 Caesalpinia  253, 257, 259 Caesalpinioideae  249–50, 253 Caesia  168 Caesulia  604 caihuba 102 Cailliella  355 Caiophora  476 Cajanus  251, 255 Cakile  416 calabash  289, 328, 567 calabrese 418 calabura 387 Calacanthus  564 Caladenia  159 Caladium  119–20 calafate 220

Calamagrostis  208 calambac 396 Calamites  24, 72 Calamoideae 177 calamondin 376 Calamovilfa  208 Calamus  177 Calandrinia  464 Calanthe  155 Calanthea  413 Calantica  329 Calanticaria  604 Calathodes  220 Calatola  514 Calcareoboea  551 Calceolaria  551 Calceolarioideae 551 Calcicola  318 Calciphila  527 Calciphilopteris  53 Caldcluvia  299 Calderonella  208 Caldesia  122 Calea  604, 612 Caleana  159 Calectasia  175 Calendula  604, 611, 613 Calepina  416 Calia  251 Calibrachoa  536 calico flower  101 Calicorema  450 Calicotome  251 California  342 Californian poppy  216 Caliphruria  171 Calla  119 calla lilies  120 Callaeum  318 Callerya  251 Calliandra  253, 259 Calliandropsis  253 Callianthe  390 Callianthemum  220 Callicarpa  575–6, 578 Callicarpoideae 576 Callicephalus  604 Callichilia  527 Callichlamys  567 Callicoma  299 Calligonum  435 Callilepis  604 Callipeltis  517 Calliphysalis  536 Callirhoe  390 Callisia  180 Callistachys  251 Callistemon  351 Callistephus  604, 613 Callisthene  350 Callitriche  553 Callitris  84 Callitropsis  84 Callopsis  119 Callostylis  155 Callothlaspi  416 Calluna  509 Calobota  251 Calocedrus  84 Calocephalus  604 Calochilus  159 Calochlaena  39, 41, 44 Calochone  517 Calochortus  150 Calocrater  527 Calodecaryia  381

Calodendrum  375 Calolisianthus  522 Calomeria  604 Caloncoba  324 Calophaca  251 Calophyllum  310 Calopis  204 calopo 257 Calopogon  155, 251 Calopogonium  251, 257 Calopyxis  346 Calorezia  603 Calorophus  204 Calostemma  171 Calostephane  604 Calotesta  604 Calothamnus  351 Calotis  604 Calotropis  527 Calpocalyx  253 Calpurnia  251 Caltha  220, 222 Caluera  155 Calvoa  355 Calycacanthus  564 Calycadenia  604 Calycera  601 Calycobolus  534 Calycocarpum  218 Calycogonium  355 Calycolpus  351 Calycopeplus  335 Calycophyllum  517 Calycophysum  287 Calycopteris  346 Calycorectes  351 Calycoseris  604 Calycosia  517 Calycosiphonia  517 Calycotropis  447 Calyculogygas  390 Calydorea  167 Calymmanthera  155 Calymmanthium  469 Calymmatium  416 Calypso  155 Calyptocarpus  604 Calyptraemalva  390 Calyptranthera  527 Calyptranthes  351 Calyptrion  325 Calyptrocalyx  177 Calyptrocarya  202 Calyptrochilum  155 Calyptrochloa  208 Calyptrogenia  351 Calyptrogyne  177 Calyptronoma  177 Calyptrosciadium  633 Calyptrotheca  465 Calytrix  351 Camarea  318 Camarotea  564 Camassia  174 Cambessedesia  355 Camchaya  604 camel bush  532 Camelina  416 camelina oil  419 Camelinopsis  416 Camellia  496 Cameraria  527 Camissonia  349 Camissoniopsis  349 Camoensia  251 Camonea  534

Campanula  591, 592–3 Campanuloideae 591–2 Campanulorchis  155 Campestigma  527 Camphor 611–12 camphor wood  112, 400, 613 Camphorosma  451 camphorwood 113 Campimia  355 Campnosperma  370 Campomanesia  351 Campovassouria  604 Campsiandra  253 Campsidium  567–8 Campsis  567–8 Camptacra  604 Camptandra  191–2 Camptocarpus  527 Camptolepis  372–3 Camptoloma  555 Camptorrhiza  146 Camptosema  251 Camptosorus  58 Camptostemon  390 Camptostylus  324 Camptotheca  472 Campuloclinium  604 Campylanthus  553 Campylocentrum  155 Campyloneurum  70 Campylopetalum  370 Campylopetalum  370 Campylosiphon  134 Campylospermum  307 Campylostachys  560 Campylostemon  294 Campylotropis  251 Campynema  141 Campynemanthe  141 Canacomyrica  279 Canadanthus  604 Canadian aspen  330 Canadian waterweed  125 Cananga  106 Canaria  633 Canarian bellflower  592 Canarina  592–3 Canarium  368 canary creeper  403 Canary Island guadil  535 Canastra  208 Canavalia  251, 255 Canbya  216 Cancrinia  604 Cancriniella  604 candelilla 335 candle tree  568 candlenut 335 candlewood 368 Candolleodendron  251 candy carrot  636 candytuft 419 Canella  95 Canellales  10–11, 95 Canephora  517 canistel 490 Canistropsis  195 Canistrum  195 Canna  188 Cannabis  273 cannabis  273, 392, 540, 610 cannibal chutney  538 cannibal’s tomato  538 Cannomois  204

Plants of the World

761

INDEX cannonball tree  487 canola 419 canola oil  419 Canotia  294 Canscora  522 Canscorinella  522 Cansjera  424 cantaloupe 288 Canthium  517 Cantinoa  576 Cantleya  585–6 Cantua  485 Capanea  551 Capanemia  155 caparrosa 461 Cape cowslip  174 Cape gooseberry  538 Cape honeysuckle  368 Cape ivy  613 Cape myrtle  494 Cape pondweed  126 Cape primroses  551 Capelio  604 Capeobolus  202 Capeochloa  208 caper  247, 401, 403, 413, 611 Caperonia  334 Caphexandra  413 Capillipedium  208 Capirona  517 Capitanopsis  576 Capitularina  202 Capnoides  216 Capnophyllum  633 Capparicordis  413 Capparis  413 Capraria  555 Caprifolioideae 623 Caprifolium  625 Capsella  416 Capsicodendron  95 Capsicophysalis  536 Capsicum  536 capucin 491 Capurodendron  490 Capuronia  346 Capuronianthus  381 Caputia  604 Caracasia  481 Caragana  251, 259 caraguá 195 Caraipa  310 Carajasia  517 Carallia  305 Caralluma  525, 527 carambola 297 Carapa  381 Carapichea  517 caraway 636 caraway thyme  578 Cardamine  416 Cardenasiodendron  370 Cardiandra  473 cardillo 611 Cardiochilos  155 Cardiochlamys  534 Cardiocrinum  150 Cardionema  447 Cardiopetalum  106 Cardiopteris  586 Cardiospermum  373 cardoon  610, 470 Cardopatium  604 Carduncellus  604 Carduoideae 602

762

Carduus  604, 610, 623 Cardwellia  225–6 Carex  15, 200, 202 Careya  486 Caribea  461 Carica  405 caricature plant  565 Carinavalva  416 Cariniana  486 Carissa  527 Carlemannia  545 Carlephyton  119 Carlesia  633 Carlina  604 Carlowrightia  564 Carlquistia  604 Carludovica  139 Carmichaelia  251, 259 Carminatia  604 Carnarvonia  226 carnation 448 carnauba palm  178 Carnauba wax  178 Carnegiea  469 carnivory  125, 167, 196, 198, 239, 300, 328, 343, 419, 436, 437, 438, 439, 440, 441, 448, 461, 462, 483, 500, 501, 502, 503, 521, 529, 541, 552, 555, 558–9, 561, 569–70, 584, 596, 625 caroá 195 carob 255 Carolina allspice  107 Carolus  318 carom seeds  636 Caropsis  633 Carpacoce  517 Carpanthea  457 Carpentaria  177 Carpenteria  473 Carpesium  604 Carpha  202 Carphalea  517 Carphephorus  604 Carphochaete  604 Carpinus  282 Carpinus   282 Carpobrotus  457 Carpodetus  589 Carpodiptera  389 Carpolobia  262 Carpotroche  324 Carpoxylon  177 Carramboa  604 Carrichtera  416 Carrierea  329 carrion plant  528 Carrissoa  251 Carronia  218 carrot 634–5 Carruanthus  457 Carruthersia  527 Carsonia  414 Carterella  517 Carthamus  604, 612 Cartonema  180 Cartonematoideae 180 Carum  633, 636 Carvalhoa  527 Carya  280 Caryocar  313 Caryodaphnopsis  112

Christenhusz, Fay & Chase

Caryodendron  334 Caryomene  218 Caryophyllales 431 Caryophyllif lora  447 Caryopteris  575–6, 578 Caryota  177 Caryotophora  457 casabanana 288 Casasia  517 Cascadia  239 Cascaronia  251 Casearia  329 cashew  254, 370–1 Casimirella  513 Casimiroa  375 cassava 335 Casselia  573 Cassia  251, 259 cassia bark  112 Cassine  294 Cassinia  604, 614 Cassinopsis  513 Cassiope  509 Cassipourea  305 cassumunar ginger  193 Cassytha  112–13 Castanea  278 Castanedia  604 Castanopsis  278 Castanospermum  251, 255 Castanospora  373 Castela  380 Castellia  208 Castelnavia  311 Castilla  274 Castilleja  583–4 cast-iron plant  174 castor oil  335, 419 Castratella  355 Castrilanthemum  604 Casuarina  281 cat tail  452 cat thyme  578 cat whiskers  414 cat’s claw  520 cat’s claw creeper  568 cat’s foot  614 cat’s tail  336 Catabrosa  208 Catacolea  204 Catalepidia  226 Catalepis  208 Catalpa  566–8 Catamixis  603 Catananche  604, 613 Catanthera  355 Catapodium  208 Catasetum  155 Catatia  604 catchfly  448 Catenulina  416 Catesbaea  517 Catha  294 Catharanthus  527 Cathariostachys  208 Cathaya  80 Cathayanthe  551 Cathcartia  216 Cathedra  422 Cathestecum  208 Cathetostemma  527 Cathormion  253 catnip 578 Catocoryne  355 Catoferia  576

Catolesia  604 Catolobus  416 Catophractes  567 Catopsis  195–6 Catostemma  390 Cattleya  152, 155 Cattleyella  155 catuaba 306 Catunaregam  517 Caucaea  155 Caucalis  633 Caucanthus  318 Caucasalia  604 Caudanthera  527 Caularthron  155 cauliflower  418 Caulokaempferia  192 Caulophyllum  219 Caustis  202 Cautleya  192 Cavacoa  335 Cavalcantia  604 Cavanillesia  390 Cavea  604 Cavendishia  509 Caxamarca  604 Cayaponia  287 Caylusea  412 Cayratia  244–5 Ceanothus  271 Cearanthes  171 Cecarria  430 Cecropia  275 Cedar Bay cherry  352 cedar cup  393 Cedrela  84, 380–1 Cedrelinga  253 Cedreloideae 381 Cedrelopsis  375 cedro macho  384 Cedronella  576 Cedrus  80, 84 Ceiba  390 Celastrales 292 celastrifolia  487 Celastroideae 294 Celastrus  294 celeriac 635 Celerina  564 celery  39, 635–7 Celianella  340 Celiantha  522 Celmisia  604, 613 Celosia  450 Celtis  273 Cenarrhenes  226 Cenchrus  208 Cenocentrum  390 Cenolophium  633 Cenostigma  253 Centaurea  603–4, 611, 613 Centaurium  522 Centaurodendron  604 Centauropsis  604 Centaurothamnus  604 Centella  629, 633, 636 Centema  450 Centemopsis  450 Centipeda  604 Centotheca  208 Centradenia  355 Centradeniastrum  355 Centranthera  583 Centranthus  623, 625 Centrapalus  604

Centratherum  604 Centrochloa  208 Centroglossa  155 Centrolepidoideae 202–4 Centrolepis  204 Centrolobium  251 Centromadia  604 Centronia  355 Centropodia  208 Centropogon  592 Centrosema  251 Centrostachys  450 Centrostegia  434 Centrostigma  159 Centrothecoideae 206 Cephalacanthus  564 Cephalanthera  155 Cephalantheropsis  155 Cephalanthus  517 Cephalaralia  630 Cephalaria  623, 625 Cephalipterum  604 Cephalocarpus  202 Cephalocereus  469 Cephalocroton  334 Cephalohibiscus  390 Cephalomappa  334 Cephalopappus  603 Cephalopentra  287 Cephalophyllum  457 Cephalopodum  633 Cephalorrhizum  433 Cephalorrhyncus  603 Cephalosorus  604 Cephalosphaera  102 Cephalostachyum  208 Cephalostemon  197 Cephalotaxus  86 Cephalotomandra  461 Ceraria  465 Cerastium  447 Ceratandra  159 Ceratiola  509 Ceratiosicyos  324 Ceratocapnos  216 Ceratocarpus  451 Ceratocaryum  204 Ceratocentron  155 Ceratocephala  220 Ceratocnemum  416 Ceratogyne  604 Ceratolacis  311 Ceratolimon  433 Ceratolobus  177 Ceratonia  253, 255, 259 Ceratopetalum  299 Ceratophyllales 212 Ceratophyllum  212 Ceratopteridoideae 50, 51, 52 Ceratopteris  50–1 Ceratopyxis  517 Ceratosanthes  287 Ceratostema  509 Ceratostigma  433 Ceratostylis  155 Ceratotheca  562–3 Ceratozamia  75 Cerbera  527 Cerberiopsis  527 Cercestis  119 Cercidiphyllum  235–6 Cercidoideae 249–50 Cercis  236, 249–50, 256, 259 Cercocarpus  263

Cerdia  447 cereal  16, 178, 200, 208, 210–11, 220, 451, 554, 584 Cereus  469 Cerinthe  531 Ceriops  305 Ceriscoides  517 Cerochlamys  457 Ceropegia  527 Cerosora  52 Ceroxyloideae 177 Ceroxylon  177 Ceruana  604 Cervantesia  427 Cervantesieae 427 Cervaria  633 Cespedesia  307 Cestrum  536 Ceterach  58 Ceuthocarpus  517 Ceuthostoma  281 Cevallia  476 Chabrea  633 Chacoa  604 Chadsia  251 Chaenactis  604 Chaenomeles  263 Chaenorhinum  553 Chaenostoma  555 Chaerophyllopsis  633 Chaerophyllum  633–4, 637 Chaetachme  272 Chaetadelpha  604 Chaetanthera  603 Chaetanthus  204 Chaetium  208 Chaetobromus  208 Chaetocalyx  251 Chaetocarpus  331 Chaetolepis  355 Chaetolimon  433 Chaetonychia  447 Chaetopappa  604 Chaetopoa  208 Chaetopogon  208 Chaetoseris  604 Chaetostachydium  517 Chaetostoma  355 Chaetymenia  604 chain-of-hearts 528 Chaiturus  576 Chalarothyrsus  564 Chalcanthus  416 Chalepophyllum  517 Chamabaina  275 Chamaeanthus  155 Chamaebatia  263 Chamaebatiaria  263 Chamaechaenactis  604 Chamaeclitandra  527 Chamaecostus  191 Chamaecrista  251 Chamaecyparis  84 Chamaedaphne  509 Chamaedorea  177 Chamaegastrodia  159 Chamaegeron  604 Chamaegigas  558 Chamaele  633 Chamaelirium  143 Chamaemeles  263 Chamaemelum  604, 611 Chamaepentas  517 Chamaepus  604

INDEX Chamaeraphis  208 Chamaerhodos  263 Chamaerops  177 Chamaesaracha  536 Chamaesciadium  633 Chamaescilla  168 Chamaesium  633 Chamaesphacos  576 Chamaexeros  174 Chamarea  633 chambala 539 Chambeyronia  177 Chamelaucium  351 Chamelophyton  155 Chameranthemum  564 chamfuta 258 Chamguava  352 Chamira  416 Chamissoa  450 chamomile 611 Chamorchis  159 Champereia  424 Championia  551 Chandrasekharania  208 Changium  633 Changnienia  155 changshan 474 chaparro 318 Chapelieria  517 Chapmannia  251 Chaptalia  603 Charadranaetes  604 Charadrophila  560 chard  518, 610–11 Chardinia  604 Charianthus  355 charoli nut  371 Charpentiera  450 Chartoloma  416 Chascanum  573 Chascotheca  341 Chasmanthe  167 Chasmanthera  218 Chasmantium  208 Chasmatophyllum  457 Chasmopodium  208 Chassalia  517 chaste tree  578 Chaubardia  155 Chaubardiella  155 Chauliodon  155 chaulmoogra oil  324 Chaunanthus  416 Chaunochiton  422 Chayamaritia  551 chayote 288 cheeseberry 266 cheesewood 628–9 Cheilanthes  53 Cheilanthoideae  50, 51, 53–4 Cheiloclinium  294 Cheilocostus  191 Cheilophyllum  553 Cheilosa  334 Cheilosoideae 334 Cheilosoria  53 Cheilotheca  509 Cheiradenia  155 Cheiranthera  628–9 Cheiridopsis  457 Cheirodendron  630 Cheirolaena  390 Cheirolophus  604 Cheiropleuria  33 Cheirostylis  159

Chelidonium  216 Chelonanthus  522 Chelone  553 Chelonistele  155 Chelonopsis  576 Chelyocarpus  177 chempedak 274 Cheniopanax  630 Chenopodiastrum  451 Chenopodioideae 450 Chenopodiopsis  555 Chenopodium  451 cherimoya 106 Cherleria  447 cherry plum  264 Chersodoma  604 chervil 634 Chesneya  251, 633 chestnut 635 Chevalierella  208 Chevreulia  604 Cheyniana  352 Chiangiodendron  324 Chiastophyllum  241 chica 567 chicasaw plum  265 Chichicaste  476 chickling vetch  255 chickpeas 254–5 chickrassy 381 chickweed 448 chicory 610 chicory 610 Chidlowia  253 chikanda 152 Chikusichloa  206 childlife tree  315 Chilean glory flower  568 Chilean hazel  226 Chilean holly  617 Chilean murta  352 Chilean palo verde  255 Chileranthemum  564 chili pepper  538 Chiliadenus  604 Chiliocephalum  604 Chiliophyllum  604 Chiliotrichiopsis  604 Chiliotrichum  604 Chilocardamum  417 Chilocarpus  527 Chiloglottis  159 Chilopsis  567–8 Chiloschista  155 Chimaerochloa  208 Chimantaea  603 Chimaphila  509 Chimarrhis  517 Chimonanthus  107 Chimonobambusa  208 Chimonocalamus  208 china doll  568 China rose  393 Chinese artichoke  573 Chinese ash  546 Chinese aster  613 Chinese cardamom  193 Chinese celery  636 Chinese foxglove  584 Chinese gooseberry  504 Chinese lantern  538, 541 Chinese olive  368 Chinese sweet plum  271 Chinese tallow tree  368 chinquapin 278 Chiococca  517

Chionachne  208 Chionanthus  546 Chione  517 Chionocharis  531 Chionochloa  208 Chionodoxa  173 Chionographis  143 Chionolaena  604 Chionopappus  604 Chionophila  553 Chionothrix  450 Chiranthodendron  390 Chironia  522 Chiropetalum  334 Chisocheton  381 ×Chitalpa  568 chives 171 Chlaenandra  218 Chlaenosciadium  633 Chlamydacanthus  564 Chlamydocardia  564 Chlamydocarya  513 Chlamydojatropha  335 Chlamydophora  604 Chlamydophytum  425 Chlamydostachya  564 Chlidanthus  171 Chloanthes  576 Chloracantha  604 Chloraea  159 Chloranthales 10–11, 114, 212 Chloranthus  114 Chloridoideae 206 Chloris  208 Chlorocalymma  208 Chlorocarpa  324 Chlorocrambe  417 Chlorogalum  174 Chloroleucon  253 Chlorophytum  173–4 Chloropyron  583 Chlorospatha  119 Chloroxylon  375 chocolate  1, 178, 255, 303, 325, 335, 391, 509, 578 chocolate cosmea  614 chocolate mint  578 Choerospondias  370 Choisya  375 chokeberry 266 chokecherry 265 cholla gum  470 Chomelia  517 Chondrilla  604 Chondrodendron  218 Chondropyxis  604 Chondrorhyncha  155 Chondroscaphe  155 Chondrostylis  334 Chondrosum  208 Chonemorpha  527 Chonocentrum  341 Chonopetalum  373 Chordifex  204 Choretrum  427 Choriantha  531 Choricarpia  352 Choriceras  539 Chorigyne  139 Chorilaena  375 Choriptera  451 Chorisandrachne  341 Chorisepalum  522 Chorisia  389

Chorisochora  564 Chorispora  417 Choritaenia  633 Chorizandra  202 Chorizanthe  434 Chorizema  251, 259 Chortolirion  168 Chouxia  373 Chresta  604 Christensenia  27 Christia  251 Christiana  389 Christianella  318 Christisonia  583 Christmas bush  629 Christmas tree  80, 430 Christolea  417 Christopheria  551 christophine 288 Chroestes  564 Chromolaena  604 Chromolepis  604 Chromolucuma  490 Chroniochilus  155 Chronopappus  604 Chrozophora  334 Chrysactinia  604 Chrysactinium  604 Chrysanthellum  604 Chrysanthemoides  602, 604 Chrysanthemum  603–4, 612–13 chrysanthemum 610, 612–13 chrysanthemum greens 610 Chrysanthoglossum  604 Chrysitrix  202 Chrysobalanus  322 Chrysocephalum  604, 613 Chrysochamela  417 Chrysochlamys  309 Chrysochloa  208 Chrysocoma  604 Chrysoglossum  155 Chrysogonum  604, 613 Chrysolaena  604 Chrysolepis  278 Chrysoma  604 Chrysophthalmum  604 Chrysophylloideae 490 Chrysophyllum  490 Chrysopogon  208, 211 Chrysopsis  604, 613 Chrysoscias  251 Chrysosplenium  239 Chrysothamnus  604 Chrysothemis  551 Chthonocephalus  604 Chuanminshen  633 Chucoa  603 Chukrasia  381 Chunia  235 Chuniophoenix  177 chuño 145 chupa chupa  391 chupadilla 371 Chuquiraga  603 churnwood 586 Chusquea  220 chutney  255, 370, 510, 538 Chydenanthus  486 Chylismia  349

Chylismiella  349 Chymocarpus  403 Chymsydia  633 Chysis  155 Chytranthus  373 Chytroglossa  155 Cibirhiza  527 Cibotioideae 42 Cibotium  39, 42 Cicendia  522 Cicer  251, 255 Cicerbita  603 Ciceronia  604 Cichorioideae 603 Cichorium  604, 610 Cicuta  633, 636 Cienfuegosia  390 cigar flower  347 cigar-tree 568 cilantro 636 Ciliosemina  517 Cimicifuga  208 Cinchona  517 Cinchonopsis  517 Cincinnobotrys  355 Cindasia  564 Cineraria  604 Cinna  208 Cinnadenia  112 Cinnamodendron  95 Cinnamomum  112 cinnamon  107, 112, 117, 152 cinnamon fern  30 Cinnamosma  95 Cionosicyos  287 Cipadessa  381 Cipocereus  469 Cipoia  311 Cipura  167 Cipuropsis  195 Circaea  349 Circandra  457 Circeastaer  216–17 Cirrhaea  155 Cirsium  604, 610, 613 ciruela 318 Cischweinfia  155 Cissampelopsis  604 Cissampelos  218 Cissus  245 Cistanche  582–4 Cistanthe  464 Cistus  386, 398 Cithareloma  417 Citharexylum  572–3 citrange 378 citrangequat 376 citron  375, 376, 378 Citronella  586 citronella grass  211 Citrophyllum  375 Citropsis  375 Citrullus  287 Citrus  375 citrus  188, 374–6, 378, 635 Cladanthus  605 Claderia  155 Cladium  202 Cladoceras  517 Cladochaeta  605 Cladocolea  430 Cladogelonium  335 Cladogynos  334 Cladopus  311

Cladoraphis  208 Cladostemon  413 Cladrastis  251, 259 Claoxylon  334 Clappertonia  389 Clappia  605 Clara  174 Clarisia  274 Clarkella  517 Clarkia  349 clarkia 349 Clastopus  417 Clathrotropis  251 Clausena  375 Clausia  417 Clausospicula  208 Clavatipollenites  92 Clavija  494 Claytonia  464 Cleghornia  527 Cleidiocarpon  334 Cleidion  334 Cleisocentron  155 Cleisomeria  155 Cleisostoma  155 Cleistachne  208 Cleistanthus  341 Cleistes  154 Cleistesiopsis  154 Cleistocactus  469 Cleistochlamys  106 Cleistochloa  208 Cleistogenes  208 Cleistopholis  106 Clematepistephium  154 Clematis  220, 222 Clematocissus  245 Clematoclethra  504 clementine 378 Cleobulia  251 Cleome  413–14 Cleomella  414 Cleonia  576 Cleoserrata  414 Cleretum  457 Clermontia  592 Clerodendrum  575–6, 578 Clethra  505 Cleyera  489 Clianthus  251, 259 Clibadium  605 Clidemia  355 Cliffortia  263 Cliftonia  506 climbing hydrangea  474 climbing lignum  435 Clinacanthus  564 Clinanthus  171 Clinopodium  575–7 Clinosperma  177 Clinostigma  177 Clintonia  150 Cliococca  337 Clistax  564 Clitandra  527 Clitoria  251, 259 Clitoriopsis  251 Clivia  171 clockvine 565 Cloezia  352 cloudberry 266 clove  112, 353, 568 clove pink  448 clover 257 Clowesia  155

Plants of the World

763

INDEX clubmoss  1, 18–19, 21 Clusia  309 Clusiella  310 cluster lily  174 Clutia  331, 395 Clymenia  375 Clypeola  417 Cnemidaria  43 Cneoridium  375 Cneoroideae 375 Cneorum  375 Cnesmocarpon  373 Cnesmone  334 Cnestidium  296 Cnestis  296 Cnicothamnus  603 Cnicus  611 Cnidiocarpa  633 Cnidoscolus  335 Coatesia  375 Coaxana  633 Cobaea  485 Cobana  167 Cobananthus  551 coca  306, 528 Coca Cola  306 cocaine 306 Coccineorchis  159 Coccinia  287 Coccochondra  517 Coccocypselum  517 Coccoloba  434 Cocconerion  335 Coccothrinax  177 Cocculus  218 cochineal 470 Cochleanthes  155 Cochlearia  417 Cochlianthus  251 Cochliostema  180 Cochlospermum  386, 397 cock’s-eggs 541 cockscomb 452 cocky apple  487 cocoa  102, 335, 390–1 coco-de-mer  116, 178–9, 254 cocona 539 coconut palm  179 cocoplums  322, 513 Cocos  177 cocozzelle 288 Codariocalyx  251, 259 Coddia  517 Codia  299 Codiaeum  335 Codiocarpus  585 Codon  531 Codonacanthus  564 Codonanthe  551 Codonanthopsis  551 Codonoboea  551 Codonocarpus  411 Codonopsis  592–3 Codonorchis  159 Coelachne  208 Coelachyrum  208 Coelanthum  462 Coelia  155 Coeliopsis  155 Coelocarpum  573 Coelocaryon  102 Coelogyne  155 Coeloneurum  536 Coelopyrena  517 Coelorachis  208

764

Coelospermum  517 Coelostegia  389 Coffea  517 coffee  1, 193, 255, 258, 278, 519, 610, 636 coffee plum  330 Cogniauxia  287 cohoba 258 Coilochilus  159 Coincya  417 Coix  208, 210–11 Cojoba  253 Cola  390 Colanthelia  208 colchicine 146 Colchicum  146 Coldenia  531 Colea  567 Coleactina  517 Coleanthera  509 Coleanthus  208 Colebrookea  576 Coleocarya  204 Coleocephalocereus  469 Coleochaete  3 Coleochloa  202 Coleocoma  605 Coleogyne  263 Coleonema  375 Coleostachys  318 Coleostephus  605 Coleotrype  180 coleus 578 Colicodendron  413 Colignonia  461 Collabium  155 Collaea  251 Colletia  271 Colletoecema  517 Colletogyne  119 Colliguaja  335 Collinsia  553 Collinsonia  576 Collomia  485 Collospermum  161 Colobanthera  605 Colobanthus  206, 447 Colocasia  119–20 Cologania  251 Cololobus  605 Colombian mahogany 487 Colona  389 Colophospermum  250 Colpias  555 Colpodium  208 Colpoon  427 Colpothrinax  177 Colquhounia  576, 578 coltsfoot 612 Colubrina  271 Columbia  389 Columbiadoria  605 columbine 222 Columellia  616–17 Columnea  551 Colutea  251 Coluteocarpus  417 Colvillea  253 Comaclinium  605 Comandra  427 Comandreae 427 Comanthera  198 Comanthosphace  576 Comarostaphylis  509 Comastoma  522

Christenhusz, Fay & Chase

Combera  536 Comborhiza  605 Combretocarpus  284 Combretum  346 Comesperma  262 Cometes  447 comfrey 532 Comiphyton  305 Commelina  180–1 Commelinales 10–11, 150, 179, 204 Commelinanae  175 Commelinoideae 180 Commersonia  389 Commidendrum  605, 614 Commiphora  368 common bean  7, 254 common reed  211 Comocladia  370 Comolia  355 Comoliopsis  355 Comoranthus  546 Comospermum  174 Comparettia  155 Compsoneura  102 Comptonella  375 Comptonia  279 Conandrium  494 Conandron  551 Conanthera  159, 165 Conceveiba  334 Conchidium  155 Conchocarpus  375 Conchopetalum  373 Condalia  271 Condaminea  517 Condea  576 Condylidium  605 Condylocarpon  527 Condylopodium  605 cone flower  611, 614 cone pepper  538 Confederate rose  393 Congea  576, 578 Congolanthus  522 congonha 586 Conicosia  457 conifer  71–3, 75, 77, 79, 81 Conimitella  239 Coniogramme  51 Conioselinum  633, 635 Conium  633, 636 Connarus  296 Connellia  195 Connorochloa  208 Conobea  553 Conocalyx  564 Conocarpus  346 Conocliniopsis  605 Conoclinium  605 Conomitra  527 Conopholis  582–3 Conophytum  457 Conopodium  633, 635 Conospermum  226 Conostegia  355 Conostephium  509 Conostomium  517 Conostylidoideae 184 Conostylis  184 Conothamnus  352 Conradina  576, 578 Conringia  417 Consolida  220 Constancea  605

Constantia  155 Convallaria  174 Convolvulus  534 Conyza  605 Conzattia  253 coohoy nut  226 Coombea  375 Coopernookia  600 Copaifera  250, 256 copal 256 Copernicia  177 Copiapoa  469 copihue 147 copperwood 368 Coprosma  517 Coptidium  220 Coptidoideae 220 Coptis  220 Coptocheile  551 Coptophyllum  517 Coptosapelta  517 Coptosperma  517 coral drops  174 coral vine  421, 435 coralbells 239 coralberry 494 Corallocarpus  287 Corallodiscus  551 Corallorhiza  155 Corbichonia  454 Corchoropsis  390 Corchorus  389 Cordeauxia  253, 255, 257 Cordia  531 Cordiera  517 Cordillera  322 Cordisepalum  534 Cordobia  318 Cordyla  251 Cordylanthus  583 Cordyline  173–4 Cordyloblaste  498 Cordylocarpus  417 Cordylostigma  517 Corema  509 Coreocarpus  605 Coreopsis  605, 612, 614 Corethamnium  605 Corethrodendron  251 Corethrogyne  605 coriander  436, 636 Coriandrum  633, 636 Coriaria  286 Corinthian capital  565–6 Coris  494 Corispermum  451 Coristospermum  633 cork oak  278 corkscrew willow  330 corkwood  276, 380 corn  200, 210 corn marigold  610 cornales  311, 471–3, 475, 477, 479, 514–5, 533, 620, 629 cornelian cherry  479 cornflower  613 Cornucopiae  208 Cornukaempferia  192 Cornulaca  451 Cornus  479 Cornutia  576, 578 Corokia  598 Coronanthera  551 Coronidium  605, 613 Coronilla  251, 259

Corpuscularia  457 Correa  375 Corrigiola  447 Corryocactus  469 Corsia  142 Corsiopsis  142 Cortaderia  208, 211 Cortia  633 Cortiella  633 Coryanthes  155 Corybas  159 Corycium  168 Corydalis  216 Coryloideae 282 Corylopsis  235 Corylus  282 Corymbia  352 Corymbioideae 603 Corymbium  605 Corymborkis  155 Corynabutilon  390 Corynaea  425 Corynanthe  517 Corynanthera  352 Corynephorus  208 Corynocarpus  285 Corynotheca  159 Corypha  177 Coryphantha  469 Coryphoideae 177 Coryphothamnus  517 Corythophora  486 Corytoplectus  551 Coscinium  218 Cosentinia  52 cosmea  548, 613 Cosmelia  509 Cosmianthemum  564 Cosmibuena  517 Cosmocalyx  517 Cosmos  605, 613–14 Cosmostigma  527 Cossinia  372–3 Costera  509 Costularia  202 Costus  191 Cota  605, 613–14 Cotinus  370 Cotoneaster  263 cotoneaster 266 Cotopaxia  633 Cottea  208 Cottendorfia  195 cotton  4, 42, 257, 337, 385, 392, 497, 612 cotton thistle  610, 612 Cottonia  155 cottonseed oil  392 Cottsia  318 Cotula  605, 614 Cotylanthera  522 Cotyledon  241 Cotylelobium  400 Cotylolabium  159 Couepia  322 Coula  422 Couloideae 422 Coulterella  605 Coulteria  253 Coulterophytum  633 Couma  527 Couratari  486 courgette 288–9 Couroupita  486 Coursetia  251 Cousinia  605

Cousiniopsis  605 cousroots 635 Coussapoa  275 Coussarea  517 Coutaportia  517 Coutarea  517 Coutoubea  522 cowbane 636 Cowiea  517 coyote mint  578 crab apple  265 Crabbea  564 Cracosna  522 Craibia  251 Craibiodendron  509 Craigia  389 Crambe  417 Crambella  417 cranberry 509–10 cranberry bush  620 crane’s bill  343 Cranichis  159 Craniolaria  560–1 Craniospermum  531 Craniotome  576 Cranocarpus  251 Craspedia  605, 614 Craspedolobium  251 Craspedorhachis  208 Craspidospermum  527 Crassocephalum  605 Crassothonna  605 Crassula  241 Crassuloideae 241 Crataegus  263 Crateranthus  486 Craterispermum  517 Craterocapsa  592 Craterosiphon  396 Craterostigma  558 Crateva  413 Cratoxylum  313 Cratylia  251 Cratystylis  605 Crawfurdia  522 creeping foxglove  565 creeping indigo  257 Cremanthodium  605 Cremaspora  517 Cremastra  155 Crematosperma  106 Cremersia  551 Cremnophyton  451 Cremnothamnus  605 Cremocarpon  517 Cremolobus  417 Cremosperma  551 Cremospermopsis  551 Crenea  346 Crenidium  536 Creochiton  355 crepe ginger  191 crepe myrtle  347 Crepidiastrum  605 Crepidifolium  605 Crepidium  155 Crepidorhopalon  558 Crepidospermum  368 Crepis  605 Crescentia  566–8 cress 419 Cressa  534 Cribbia  155 Crinipes  208 Crinitaria  605 Crinodendron  300

INDEX Crinum  171 Crioceras  527 Criscia  603 Criscianthus  605 Crispiloba  597 Cristaria  390 Cristatella  414 Cristonia  251 Crithmum  633, 636 Crithopsis  208 Critonia  605 Critoniadelphus  605 Critoniella  605 Critoniopsis  605 Croatiella  119 Crobylanthe  517 Crocanthemum  398 Crocidium  605 Crocoideae 167 Crocosmia  167 Crocus  167, 192, 612 Croizatia  341 Cromidon  555 Croninia  509 Cronquist, Arthur  475 Cronquistia  605 Cronquistianthus  605 crookneck squash  288 Croomia  138 Croptilon  605 crosnes 578 Crossandra  564–5 Crossandrella  564 crossberry 393 Crosslandia  202 Crossoglossa  155 Crossoliparis  155 Crossopetalum  294 Crossopteryx  517 Crossosoma  363 Crossosomatales 358 Crossosperma  375 Crossostemma  327 Crossostephium  605 Crossostylis  305 Crossothamnus  605 crossvine 568 Crossyne  171 Crotalaria  250–1, 258 Croton  15, 355 croton 336 Crotonogyne  335 Crotonogynopsis  334 Crotonoideae 335 crowberry 509 Crowea  375 crown daisy  613 crown of thorns  266, 271, 326 crown-beard 614 crowsfoot elm  392 Crucianella  517 Cruciata  517 Crucifera  416 Crucihimalaya  417 Cruckshanksia  517 Cruddasia  251 Crudia  250 Crumenaria  271 crummock 634 Crupina  605 Crusea  517 Cryosophila  177 Crypsis  208 Crypt ra  271 Cryptantha  531

Cryptanthus  195 Cryptarrhena  155 Crypteronia  356 Cryptocapnos  216 Cryptocarpus  461 Cryptocarya  112 Cryptochilus  155 Cryptochloa  208 Cryptocodon  592 Cryptocoryne  119–20 Cryptogramma  51 Cryptogrammoideae 50 Cryptolepis  527 Cryptomeria  84 Cryptopus  155 Cryptopylos  155 Cryptosepalum  250 Cryptospora  417 Cryptostegia  527 Cryptostephanus  171 Cryptostylis  159 Cryptotaenia  633, 636 Ctenardisia  494 Ctenitis  66–7 Ctenium  208 Ctenolepis  287 Ctenolophon  304 Cuatrecasanthus  605 Cuatrecasasiella  605 Cuatresia  536 Cubanola  517 cubebs 99 Cubilia  373 Cubitanthus  558 cucumber 287–90 Cucumis  287 Cucurbita  287 Cucurbitales 283 Cucurbitella  287 Cuervea  294 Cuitlauzina  155 Culcasia  119 Culcita  39, 41–2, 44 Culcitium  605 Culcitoideae 41 Cullen  251 Cullenia  389 Cullumia  605 cumin  141, 636 Cuminia  576 Cuminum  633, 636 Cunila  576–7 Cunninghamia  84 Cunonia  299 Cunuria  335 cup-and-saucer plant  578 Cupania  373 Cupaniopsis  373 Cuphea  346 Cuphonotus  417 Cupid’s dart  613 Cuprella  417 Cupressales  10–11, 83 Cupressus  84 cupuaçu 391 curare 218 Curarea  218 Curatella  231 Curculigo  162–3 Curcuma  191–3 Curio  605, 614 Curitiba  352 currant tree  341 currants  238, 245, 355, 435 curry plant  612, 614

curry tree  378 Curtia  522 Curtisia dentata  477 Curupira  422 Cuscatlania  461 Cuscuta  534 Cusickiella  417 Cuspidaria  567 Cuspidia  605 Cussetia  311 Cussonia  630 custard apple  106 Cutandia  208 Cuttsia  589 Cuviera  517 Cyamopsis  251, 256 Cyanaeorchis  155 Cyananthus  592 Cyanastrum  164–5 Cyanea  592–3 Cyanella  165 Cyanicula  159 Cyanixia  167 Cyanocephalus  576 Cyanoneuron  517 Cyanostegia  576 Cyanothamnus  375 Cyanotis  180 Cyanthillium  605 Cyathea  39–44 Cyatheales  10–11, 36, 39 Cyatheoideae  39–40, 43 Cyathobasis  451 Cyathocalyx  106 Cyathochaeta  202 Cyathocline  605 Cyathocoma  202 Cyathodes  509 Cyathomone  605 Cyathophylla  447 Cyathopsis  509 Cyathopus  208 Cyathoselinum  633 Cyathostegia  251 Cyathostemon  352 Cyathula  450 Cybebus  159 Cybianthus  494 Cybistax  567 cycad  71, 73–4 Cycadales  10–11, 73 Cycadeoidales 72 Cycas  73–4 Cyclacanthus  564 Cycladenia  527 Cyclamen  494 Cyclanthera  287 Cyclantheropsis  287 Cyclanthus  139 Cyclea  218 Cyclocarpa  251 Cyclocarya  280 Cyclocheilon  583 Cyclocodon  592 Cyclocotyla  527 Cyclodium  66 Cyclolepis  603 Cyclolobium  251 Cycloloma  451 Cyclopeltis  66–7 Cyclophyllum  517 Cyclopia  251, 256 Cyclopogon  159 Cyclorhiza  633 Cyclosorus  59 Cyclospermum  633

Cyclostachya  208 Cyclotrichium  576 Cycnium  583 Cycnoches  155 Cycnogeton  127 Cydonia  263 Cylicodiscus  253 Cylicomorpha  405 Cylindrocarpa  592 Cylindrocline  605 Cylindrophyllum  457 Cylindropsis  527 Cymaria  576 Cymatocarpus  417 Cymbalaria  553 Cymbaria  583 Cymbidiella  155 Cymbidium  152, 155 Cymbocarpa  134 Cymbolaena  605 Cymbonotus  605 Cymbopappus  605 Cymbopetalum  106 Cymbopogon  208, 211 Cymbosema  251 Cymodocea  131 Cymophora  605 Cymophyllus  202 Cymopterus  633 Cynanchum  527 Cynapium  633 Cynara  605, 610 Cynarospermum  564 Cyne  430 Cynodon  208, 211 Cynoglossopsis  531 Cynoglossum  531 Cynoglottis  531 Cynometra  250 Cynomorium  244, 425 Cynophalla  413 Cynorhiza  633 Cynorkis  159 Cynosciadium  633 Cynosurus  208 Cypella  167 Cyperochloa  208 Cyperoideae  200, 202 Cyperus  202 Cyphacanthus  564 Cyphanthera  536 Cyphia  592 Cyphocardamum  417 Cyphocarpus  592 Cyphochlaena  208 Cyphokentia  177 Cypholophus  275 Cypholoron  155 Cyphonanthus  208 Cyphophoenix  177 Cyphosperma  177 Cyphostemma  245 Cyphostigma  191 Cyphostyla  355 Cyphotheca  355 cypre 531 cypress vine  535 Cypringlea  202 Cypripedium  154 Cypselea  457 Cypselocarpus  411 Cyrilla  506 Cyrillopsis 338 Cyrtandra  551 Cyrtandromoea  551 Cyrtanthus  171

Cyrtocarpa  370 Cyrtochiloides  155 Cyrtochilum  155 Cyrtochloa  208 Cyrtococcum  208 Cyrtocymura  605 Cyrtogonone  335 Cyrtomidictyum  66 Cyrtopodium  155 Cyrtorchis  155 Cyrtosia  154 Cyrtosperma  119 Cyrtostachys  177 Cyrtostylis  159 Cysticapnos  216 Cystoathyrium  56 Cystodium  47 Cystopteridoideae 55–6, 60 Cystopteris  55–6, 60 Cystorchis  159 Cystostemon  531 Cytinus  386 Cytisophyllum  251 Cytisopsis  251 Cytisus  251, 256, 258–9 Cytogonidium  204 Cyttaranthus  334   dabai 368 Daboecia  509 Dacrycarpus  82 Dacrydium  82 Dacryodes  368 Dacryotrichia  605 Dactyladenia  322 Dactylaena  414 Dactylanthus  425 Dactylicapnos  216 Dactyliophora  430 Dactylira  287 Dactylis  208 Dactylocardamum  417 Dactylocladus  356 Dactyloctenium  208 Dactylopetalum  305 Dactylopsis  457 Dactylorhiza  159 Dactylostalix  155 Daemonorops  177 Daenikera  427 daffodil 171 Dahlia  605, 611, 613–15 dahlia  611, 613 Dahliaphyllum  633 Dahlstedtia  251 dahoon 589 daikon 419 Daiotyla  155 Dais  396 Daiswa  143 daisy bush  613 Dalbergia  251, 258 Dalbergiella  251 Dalea  251, 259 Dalechampia  334 Dalembertia  335 Dalenia  355 Dalhousiea  251 Dallachya  271 Dallwatsonia  208 Dalrympelea  361 Dalzellia  311 Damasonium  122 dame’s violet  419 Damnacanthus  517

Damnamenia  605 Damnxanthodium  605 Dampiera  600 Damrongia  551 damson 264 damson-plum 490 Danaë  173–4 Danaea  27 Danaeoideae 27 Danais  517 dandelion 610 Dandya  174 danewort 620 Danguya  564 Danhatchia  159 Daniellia  250, 256 Dansiea  346 Danthonia  208 Danthoniastrum  208 Danthonidium  208 Danthonioideae 206 Danthoniopsis  208 Danxiaorchis  155 dao 370–1 Dapania  297 Daphnandra  109 Daphne  396 Daphniphyllum  236 Daphnopsis  396 Dapsilanthus  204 Darcya  553 Darcyanthus  536 Darlingia  226 Darlingtonia  501 Darmera  239 darnel 211 Darwin, Charles 1, 152, 239, 257, 541, 614–5 Darwinia  352 Darwiniothamnus  605 Dasispermum  633 Dasistoma  583 Dasyandantha  605 Dasyanthina  605 Dasychloa  208 Dasycondylus  605 Dasylepis  324 Dasylirion  173–4 Dasymaschalon  106 Dasynotus  531 Dasyphyllum  603 Dasypogon  175 Dasypyrum  208 Dasysphaera  450 Dasytropis  564 datach 256 date palm  178–9 date plum  492 Datisca  291 Datura  536 Daubenya  174 Daucus  633–4 Dauphinea  576 Dauresia  605 Daustinia  534 Davallia  39, 68 Davallioideae  63, 67–68, 70 Davallodes  68 Daveaua  605 Davenportia  534 Davidia  472 Davidsea  208 Davidson’s plum  299 Davidsonia  299 Daviesia  251

Plants of the World

765

INDEX Davilla  231 daylilies 168 de Candolle, Augustin Pyrame 257 de Jussieu, Antoine Laurent 349 dead nettles  578 deadly nightshade  540 Debia  517 Debregeasia  275 Decachaeta  605 Decagonocarpus  375 Decaisnea  217 Decaisnina  430 Decalepidanthus  531 Decalepis  527 Decalobanthus  534 Decarya  465 Decarydendron  111 Decaryella  208 Decaryochloa  208 Decaschistia  389 Decaspermum  352 Decastylocarpus  605 Decatoca  509 Decatropsis  375 Decazesia  605 Decazyx  375 Deccania  517 Deceptor  155 Deckenia  177 Declieuxia  517 Decodon  346 Decorsea  251 Decorsella  325 Decumaria  473 Dedeckera  434 Deeringia  450 Degeneria  104 Degenia  417 Degranvillea  159 Deguelia  251 Dehaasia  112 Deherainia  494 Deianira  522 Deidamia  327 Deinacanthon  195 Deinandra  605 Deinanthe  474 Deinbollia  373 Deinostema  553 Deinostigma  551 Deiregyne  159 Delairea  605, 614 Delamerea  605 Delarbrea  629, 631 Delavaya  373 Delilia  605 Delissea  592 Delonix  253, 255, 259 Delosperma  457 Delostoma  567 Delphinium  220, 222 Delpinophytum  417 Delpydora  490 Deltaria  396 Demavendia  633 Demosthenesia  509 Dendroarabis  417 Dendrobangia  514 Dendrobium  15, 152, 155 Dendrocacalia  605 Dendrocalamus  208, 211 Dendrochilum  155 Dendrocnide  275 Dendroconche  70

766

Dendrokingstonia  106 Dendrolobium  251 Dendromecon  215–16 Dendromyza  427 Dendropanax  630–1 Dendropemon  430 Dendrophorbium  605 Dendrophthoe  430 Dendrophthora  427 Dendrophylax  155 Dendrosenecio  605 Dendroseris  605 Dendrosicyos  287 Dendrosida  390 Dendrosipanea  517 Dendrosma  375 Dendrotrophe  427 Denekia  605 Denhamia  294 Denmoza  469 Denscantia  517 Dentella  517 Depanthus  551 Deparia  60 depgul 579 Deplanchea  567 Deppea  517 Deprea  536 Dermatobotrys  555 Derris  251, 259 Deschampsia  206, 208 Descurainia  417 desert date  247, 371 desert kumquat  376 desert rose  393, 528 desert spoon  174 desert willow  568 Desfontainia  523, 616–17 Desmanthodium  605 Desmanthus  253 Desmaria  430 Desmazeria  208 Desmidorchis  527 Desmocladus  204 Desmodiastrum  251 Desmodium  251 Desmogymnosiphon  134 Desmoncus  177 Desmophlebium  58 Desmopsis  106 Desmos  106 Desmoscelis  355 Desmostachya  208 Desmostachys  513 Desmotes  375 Desplatsia  389 Detarioideae 249–50 Detarium  250, 256 Dethawia  633 Deuterocohnia  195 Deutzia  473 Deutzianthus  335 Deverra  633 Devia  167 devil’s claw  563 devil’s hand tree  393 Devogelia  155 dewberry 266 Dewevrea  251 Dewevrella  527 Dewildemania  605 dhani 636 Dhofaria  413 Diabelia  622 diabetic sugar  611 Diacidia  318

Christenhusz, Fay & Chase

Diacranthera  605 Diacrodon  517 Dialioideae  249, 251 Dialium  251, 255 Dialyceras  394 Dialypetalanthus  517 Dialypetalum  592 Dialytheca  218 Diamantina  311 Diamena  174 Diandrolyra  208 Dianella  168, 170 Dianthoseris  605 Dianthoveus  139 Dianthus  447 Diapensia  499 Diaphananthe  155 Diaphanoptera  447 Diaphractanthus  605 Diarrhena  208 Diarthron  396 Diascia  555 Diaspasis  600 Diaspis  318 Diastatea  592 Diastella  226 Diastema  551 Diatenopteryx  373 Dibrachionostylus  517 Dicarpellum  294 Dicarpidium  389 Dicella  318 Dicellostyles  390 Dicellra  355 Dicentra  216 Dicerandra  576 Diceratella  417 Diceratostele  155 Diceratotheca  564 Dicercoclados  605 Dicerocaryum  562 Dichaea  155 Dichaetanthera  355 Dichaetaria  208 Dichaetophora  605 Dichanthium  208 Dichapetalum  320 Dichasianthus  417 Dichazothece  564 Dichelachne  208 Dichelostemma  174 Dicheranthus  447 Dichilanthe  517 Dichilus  251 Dichocarpum  220 Dichondra  534 Dichopogon  174 Dichorisandra  180 Dichosciadium  633 Dichostemma  335 Dichotomanthes  263 Dichroa  473 Dichrocephala  605 Dichromanthus  159 Dichromochlamys  605 Dichrostachys  253 Dickasonia  155 Dickinsia  633 Dicksonia  39, 44 Dicksonioideae 44 Dicladanthera  564 Diclidanthera  262 Diclinanona  106 Dicliptera  564 Diclis  555 Dicoelia  341

Dicoma  603 Dicomopsis  605 Dicoria  605 Dicorynia  251 Dicoryphe  235, 386 Dicraeanthus  311 Dicraeopetalum  251 Dicranocarpus  605 Dicranolepis  396 Dicranopteris  32 Dicranopygium  139 Dicranostegia  583 Dicranostigma  216 Dicranostyles  534 Dicraspidia  387 Dicrastylis  576 Dicrocaulon  457 Dictamnus  375 Dictyandra  517 Dictymia  70 Dictyocaryum  177 Dictyolimon  433 Dictyoloma  375 Dictyoneura  373 Dictyophleba  527 Dictyophragmus  417 Dictyosperma  177 Dictyospermum  180 Dictyostega  134 Dicyclophora  633 Dicymbe  250 Dicypellium  112 Didelotia  250 Didelta  605 Didesmandra  231 Didesmus  417 Didierea  465 Didiscus  633 Didissandra  551 Didonica  509 Didymaea  517 Didymanthus  451 Didymaotus  457 Didymeles  228 Didymocarpoideae 551 Didymocarpus  551 Didymochlaena  63–4, 66 Didymochlaenoideae  63–4 Didymochlamys  517 Didymocistus  340 Didymodoxa  275 Didymogonyx  208 Didymophysa  417 Didymoplexiella  155 Didymoplexis  155 Didymopogon  517 Didymosalpinx  517 Didymostigma  551 Dieffenbachia  119–20 Diegodendron  394, 397 Dielitzia  605 Diellia  58 Dielsantha  592 Dielsia  204 Dielsiocharis  417 Dielsiochloa  208 Dielsiodoxa  509 Dielsiothamnus  106 Dierama  167 Diervilla  622, 625 Diervilloideae 622 Dieteria  605 Dieterlea  287 Dietes  167 Digera  450

Digitacalia  605 Digitalis  553 Digitaria  208, 210 Dignathia  208 Digomphia  567 Digoniopterys  318 Diheteropogon  208 dika bread  303 Dilatris  184 Dilkea  327 dill 636 Dillandia  605 Dillenia  231 Dilleniales 231 Dillwynia  251 Dilobeia  226 Dilodendron  373 Dilopha  417 Dilophotriche  208 Dimeresia  605 Dimeria  208 Dimerocostus  191 Dimerostemma  605 Dimetra  546 Dimocarpus  373 Dimorphandra  253, 257–8 Dimorphanthera  509 Dimorphocalyx  335 Dimorphocarpa  417 Dimorphocoma  605 Dimorphosciadium  633 Dimorphotheca  605, 613 Dinebra  208 Dinema  155 Dinemagonum  318 Dinemandra  318 Dinetus  534 Dinizia  253 Dinklageella  155 Dinklageodoxa  567 Dinochloa  208 Dinophora  355 Dinosperma  375 Dintera  553, 558 Dinteranthus  457 Dioclea  251 Diodella  517 Diodia  517 Diodonopsis  155 Diodontium  605 Dioecrescis  517 Diogenesia  509 Diogoa  422 Dioicodendron  517 Dionaea  437 Dioncophyllum  441 Dionycha  355 Dionychastrum  355 Dioon  75 Diora  174 Dioscorea  134–5, 534, 586 Dioscoreales  10–11, 132, 142, 147, 150 Dioscoreophyllum  218 Diosma  375 Diospyros  492 Diostea  573 Dipcadi  174 Dipelta  622 Dipentodon  384 Diphasia  375 Diphasiopsis  375 Diphyllarium  251 Diphylleia  219

Diphysa  251 Diplachne  208 Diplacrum  202 Diplacus  580 Diplarche  509 Diplarpea  355 Diplarrena  165, 167 Diplasia  202 Diplaspis  633 Diplatia  430 Diplaziopsidoideae 55–8 Diplaziopsis  57, 60 Diplazium  55, 57–8, 60 Diplectria  355 Diplocentrum  155 Diploclisia  218 Diplocyclos  287 Diplodiscus  389 Diploglottis  373 Diplokeleba  373 Diploknema  490 Diplolaena  375 Diplolepis  527 Diplolophium  633 Diplomeris  159 Diploon  490 Diplopanax  472, 629 Diplopeltis  373 Diplophractum  389 Diplopogon  208 Diploprora  155 Diplopterygium  32 Diplopterys  318 Diplorhynchus  527 Diplosoma  457 Diplospora  517 Diplostephium  605 Diplotaenia  633 Diplotaxis  417 Diplotropis  251 Diplusodon  346 Diplycosia  509 Dipodium  155 Dipogon  251 Dipoma  417 Diposis  633 Dipsacales 619 Dipsacoideae 623 Dipsacus  623, 625 Dipteris  33 dipterocarp 400 Dipterocarpoideae 400 Dipterocarpus  400 Dipterocome  605 Dipterocypsela  605 Dipteronia  372–3 Dipteropeltis  534 Dipterygium  414 Dipteryx  251, 256 Diptychandra  253 Diptychocarpus  417 Dipyrena  573 Dirca  396 Dirhamphis  390 Dirichletia  517 Disa  152, 159 Disakisperma  208 Disanthoideae 235 Disanthus  235 Disaster  389 Discaria  271 Dischidia  525, 527 Dischisma  555 Dischistocalyx  564 Disciphania  218 Discocalyx  494

INDEX Discocapnos  216 Discocarpus  341 Discoclaoxylon  334 Discocleidion  334 Discocnide  275 Discoglypremna  334 Discolobium  251 Discophora  585 Discopodium  536 Discospermum  517 Discretitheca  576 Discyphus  159 Diselma  84 Disepalum  106 Disocactus  469 Disparago  605 Disperis  159 Disphyma  457 Disporopsis  146, 174 Disporum  145–6, 150 Dissanthelium  208 Dissiliaria  539 Dissocarpus  451 Dissochaeta  355 Dissochondrus  208 Dissomeria  329 Dissothrix  605 Dissotis  355 Disteganthus  195 Distemonanthus  251 Distephanus  605 Disterigma  509 Distichia  200 Distichirhops  340 Distichlis  208 Distichochlamys  192 Distichoselinum  633 Distichostemon  373 Distimake  534 Distrianthes  430 Distyliopsis  235 Distylium  235 Distylodon  155 Disynaphia  605 Disynstemon  251 dita bark  528 Ditassa  527 Ditaxis  334 Ditepalanthus  425 Dithyrea  417 Dithyrostegia  605 Ditrysinia  335 Ditta  335 dittander 419 dittany 577–8 Dittrichia  605, 612 Diuranthera  174 Diuris  159 divi-divi 257 divine sage  578 Diyaminauclea  517 Dizygostemon  553 Djaloniella  522 Djinga  311 Dobera  408 Dobinea  370 Dobinea  370 dock  253, 435 Docynia  263 Dodartia  579 Dodecadenia  112 Dodecahema  434 Dodecastigma  335 Dodonaea  373 Dodonaeoideae 373 Doellia  605

Doellingeria  605 Doerpfeldia  271 dog rose  267 dog’s tooth violet  150 dog-fennel 613 dogwood 479 Dolianthus  517 Dolichandra  567–8 Dolichandrone  567 Dolichlasium  603 Dolichodelphys  517 Dolichoglottis  605 Dolicholobium  517 Dolicholoma  551 Dolichometra  517 Dolichopentas  517 Dolichopetalum  527 Dolichopsis  251 Dolichorrhiza  605 Dolichos  251 Dolichostachys  564 Dolichothrix  605 Dolichoura  355 Doliocarpus  231 doll’s daisy  613 Dolomiaea  605 Dolpojestella  633 Dombeya  390 Dombeyoideae 390 Domeykoa  633 Domingoa  155 Donatia  595–6 Donax  190 dondurma 152 Doniophyton  603 donkey-tea 344 Donnellsmithia  633 Donnelyanthus  517 Dontostemon  417 Doodia  61 Dopatrium  553 Doratoxylon  373 Dorcoceras  551 Dorema  633, 636 Doric columns  635 Dorobaea  605 Doronicum  605, 614 Dorotheanthus  457 Dorrigo pepper  96 Dorstenia  274 Doryanthes  164 Dorycnium  251, 259 Dorycnopsis  251 Doryopteris  53 Doryphora  109 Dorystoechas  576 Doryxylon  334 Dossinia  159 Douepea  417 doum palm  178 Douradoa  422 dove tree  472 Dovyalis  329 Downingia  592 Doyerea  287 Doyleanthus  102 Draba  417 Drabastrum  417 Drabella  417 Dracaena  173–4 Dracocephalum  576, 578 Dracoglossum  66–7 Dracomonticola  159 Draconanthes  155 Draconopteris  67 Dracontioides  119

Dracontium  119 Dracontomelon  370 Dracophilus  457 Dracophyllum  509 Dracosciadium  633 Dracoscirpoides  202 Dracula  155 Dracunculus  119 dragon fruit  470 dragon tree  174 dragon’s head  578 dragonhead 578 dragonmouth 578 Drakaea  152, 159 Dransfieldia  177 Draperia  531 Drapetes  396 dream herb  612 Dregea  527, 633 Dregeochloa  208 Drepananthus  106 Drepanocaryum  576 Drepanostachyum  208 Dressiantha  404 Dresslerella  155 Dressleria  155 Dresslerothamnus  605 Driessenia  355 Drimia  173–4 Drimiopsis  174 Drimycarpus  370 Drimys  96 Droceloncia  334 Droguetia  275 Droogmansia  251 Drosanthemum  457 Drosera  437, 596 Drosophyllum  440 Drummondita  375 Drusa  633 Dryadella  155 Dryadodaphne  109 Dryadoideae 263 Dryadorchis  155 Dryandra  226 Dryas  263 Drymaria  447 Drymoanthus  155 Drymonia  551 Drymophila  144–5 Drymophloeus  177 Drynaria  70 Drynarieae  70 Dryobalanops  400 Dryopetalon  417 Dryopoa  208 Dryopolystichum  66 Dryopsis  67 Dryopterideae  66 Dryopteridoideae 63–4, 66–7 Dryopteris  63, 66 Drypetes  315 Drypis  447 Duabanga  346 Dubautia  605 Duboisia  536 Duboscia  389 Dubouzetia  300 Dubyaea  605 Duckea  197 Duckeanthus  106 Duckeella  154 Duckeodendron  536 Duckesia  323 duckweed 118–20

Ducrosia  633 dudaim  288, 541 Dudleya  240–1 Dufrenoya  427 Dugesia  605 Duguetia  106 Duhaldea  605 Duidaea  603 Duidania  517 duke cherries  264 Dulacia  422 Dulichium  202 Duma  435 Dumasia  251 Dunalia  536 Dunbaria  251 Dunnia  517 Dunstervillea  155 Duosperma  564 Duparquetia  250 Duparquetioideae 249–50 Duperrea  517 Duperreya  534 Duplipetala  522 Dupontia  208 Dupontiopsis  208 durango root  291 Duranta  572–3 durian 391 Durio  389 Duroia  517 Durringtonia  517 durum wheat  209–10 Duseniella  603 Dussia  251 Dutailliopsis  375 Dutaillyea  375 Dutch elm disease  272 Duthiastrum  167 Duthiea  208 Duvalia  527 Duvaliandra  527 Duvernoia  564 duzhong 514 dwarf cabbage tree  593 dwarf ebony  393 Dyakia  155 Dyckia  195 dyer’s broom  257 dyer’s rocket  412 Dyera  527 Dyerophytum  433 Dymondia  605 Dypsis  177 Dysaster  605 Dyschoriste  564 Dyscritothamnus  605 Dysodiopsis  605 Dysolobium  251 Dysopsis  334 Dysosma  219 Dysoxylum  381 Dysphania  451 Dyssochroma  536 Dyssodia  605 Dystaenia  633 Dystovomita  309   Earina  155 earth almond  202 earthnut 635 Eastwoodia  605 Eatonella  605 Ebenopsis  253 Ebenus  251 Eberhardtia  490

Eberlanzia  457 ebony  258, 393, 491–2 Ebracteola  457 Ecballium  287 Ecbolium  564 Ecclinusa  490 Eccoptocarpha  208 Eccremocarpus  567–8 Ecdeiocolea  206 Echeandia  174 Echeveria  240–1 Echidnopsis  527 Echinacanthus  564 Echinacea  605, 611, 614 Echinaria  208 Echinocactus  469 Echinocereus  469 Echinochloa  208, 210 Echinocitrus  375 Echinocodon  592 Echinocoryne  605 Echinocystis  287 Echinodorus  122 Echinolaena  208 Echinopepon  287 Echinophora  633 Echinopogon  208 Echinops  605, 614 Echinopsis  469 Echinopterys  318 Echinorhyncha  155 Echinosepala  155 Echinospartum  251 Echinostephia  218 Echiochilon  531 Echiostachys  531 Echites  527 Echium  531 Eclecticus  155 Eclipta  605, 612 Ectadium  527 Ectopopteris  318 Ectrosia  208 Ecua  527 Ecuadendron  250 edelweiss  612, 614 Edgeworthia  396 Edithcolea  527 Edmondia  605 Edmundoa  195 Edraianthus  592–3 Eduandrea  195 eelgrass 127–8 Efulensia  327 Egeria  125 Eggelingia  155 eggplant  see aubergine eglantine 267 Egletes  605 egusi seed  289 Egyptian cotton  392 Ehrendorferia  216 Ehretia  531 Ehrharta  206 Ehrhartoideae 206 Eichhornia  182–3 Eidothea  226 Eigia  417 einkorn wheat  209 Einomeia  101 Eirmocephala  605 Eitenia  605 Eithea  171 Eizia  517 Ekebergia  381 Ekima  633

Ekmania  605 Ekmanianthe  567 Ekmaniopappus  605 Ekmanochloa  208 Elachanthemum  605 Elachanthus  605 Elacholoma  580 Elachyptera  294 Elaeagia  517 Elaeagnus  269 Elaeis  177 Elaeocarpus  300 Elaeodendron  294 Elaeoluma  490 Elaeoselinum  633 Elaeosticta  633 Elaphandra  605 Elaphoglosseae  66 Elaphoglossum  63, 66–7 Elasis  180 Elateriospermum  335 Elatine  316 Elatostema  275 Elattostachys  373 Elburzia  417 elder 619–21 elecampane 611 electric buttons  610 Elegia  204 Eleiodoxa  177 Eleiotis  251 Elekmania  605 Eleocharis  202 Eleorchis  155 elephant ear  119–20 elephant foot  135 elephant garlic  171 elephant grass  211 elephant’s eye  174 elephant-ears 239 Elephantomene  218 Elephantopus  605 Elephantorrhiza  253, 257 Elettaria  191, 193 Elettariopsis  191 Eleusine  208, 210 Eleutharrhena  218 Eleutherandra  324 Eleutheranthera  605 Eleutherine  167 Eleutherococcus  630–1 Eleutherospermum  633 Eleutherostylis  389 Eleuthranthes  517 Eliea  313 Eligmocarpus  251 Elingamita  494 Elionurus  208 Elizabetha  250 Elleanthus  155 Ellenbergia  605 Elliottia  509 Ellipanthus  296 Ellisia  531 Ellisiophyllum  553 Ellisochloa  208 elm  271–2, 393 Elmera  239 Elmerillia  103 Elodea  125 Eloyella  155 Elsholtzia  576, 578 Eltroplectris  159 Elymandra  209 Elymus  208 Elythranthera  159

Plants of the World

767

INDEX Elytranthe  430 Elytraria  564 Elytropappus  605 Elytrophorus  209 Elytropus  527 Elytrostachys  208 Emarhendia  551 Embadium  531 Embelia  494 Emblemantha  494 Emblingia  409, 521 Embolanthera  235 Embothrium  226 Embreea  155 Embryopsida  3 Emelianthe  430 emerald tree  568 Emex  435 Emicocarpus  527 Emilia  605 Emiliella  605 Eminia  251 Eminium  119 Emmenanthe  531 Emmenopterys  517 Emmenosperma  271 Emmeorhiza  517 emmer wheat  209 Emmotum  514 Emorya  555 emperor tree  487 Empetrum  509 emping 77 Emplectanthus  527 Empleuridium  294 Empleurum  375 Empodisma  204 Empodium  163 Empogona  517 empress tree  581 emu bush  557 Enantiophylla  633 Enarganthe  457 Enarthrocarpus  417 Enaulophyton  355 Encelia  605 Enceliopsis  605 Encephalartos  75 Encephalosphaera  564 Encholirium  195 Enchylaena  451 Encopella  553, 558 Encyclia  155 Endertia  250 Endiandra  112–3 endive 610 Endlicheria  112 Endocaulos  311 Endocellion  605 Endocomia  102 Endodesmia  310 Endonema  358 Endopappus  605 Endopleura  323 Endosamara  251 Endospermum  335 Endostemon  576 Endotheca  101 Enekbatus  352 Engelhardtia  280 Engelmannia  605 Engler, Adolf  429 Engleria  605 englerianus  304 Englerina  430 Englerocharis  417

768

Englerodendron  250 Englerophytum  490 English daisy  613 English oak  278 Engomegoma  422 Enhalus  125 Enicosanthum  106 Enicostema  522 Enkala 112 Enkianthoideae 509 Enkianthus  507, 509 Enkleia  396 Ennealophus  167 Enneapogon  209 Enriquebeltrania  334 Ensete  187 Entada  253, 258–9 Entandrophagma  381 Entelea  389 Enterolobium  253 Enteropogon  209 Entolasia  209 Entomophobia  155 Entoplocamia  209 Enydra  605 Eocaltha  220 Eomecon  216 Eosanthe  517 Epacris  509 Epaltes  605 epazote 452 Eperua  250, 256 Ephedra  78 Ephedrales 10–11, 77–8 Ephedranthus  106 ephedrine  78, 392 Ephippiandra  111 Ephippianthus  155 Epiblastus  155 Epiblema  159 Epiclastopelma  564 Epidendroideae 154–5, 158 Epidendrum  15, 152, 155, 158 Epidryos  197 Epifagis  582 Epifagus  583 Epigaea  509 Epigynum  527 Epilasia  605 Epilobium  349 Epilyna  155 Epimedium  219–20 Epipactis  155 Epipetrum  134 Epiphyllum  469 Epipogium  155 Epipremnum  119–20 Epiprinus  334 Epirixanthes  262 Epischoenus  202 Episcia  551 Epistemma  527 Epistephium  154 Epithelantha  469 Epithema  551 Epitriche  605 Epixiphium  553 Eplingiella  576 Equisetales  8, 10–11, 24 Equisetanae  8 Equisetidae 8 Equisetideae 8 Equisetinae 8

Christenhusz, Fay & Chase

Equisetineae 8 Equisetoideae 8 Equisetophyta 8 Equisetophytina 8 Equisetopsida  3, 8 Equisetum  3, 8, 24–5, 72, 78 Eragrostiella  209 Eragrostis  209–11 Eranthemum  564–5 Eranthis  220, 222 Erasanthe  155 Erato  605 Erblichia  327 Ercilla  459 Erechtites  605 Eremaea  352 Eremalche  390 Eremanthus  605, 612 Eremiolirion  165 Eremitilla  583 Eremitis  208 Eremobium  417 Eremoblastus  417 Eremocaulon  208 Eremocharis  633 Eremochloa  209, 211 Eremocitrus  375 Eremocrinum  174 Eremodaucus  633 Eremogeton  555 Eremogone  448 Eremolaena  399 Eremolimon  433 Eremomastax  564 Eremophea  451 Eremophila  555 Eremophyton  417 Eremopoa  208 Eremopyrum  208 Eremosparton  251 Eremospatha  177 Eremosyne  615–16 Eremothamnus  605 Eremothera  349 Eremurus  168, 170 Erepsia  457 Ergocarpon  633 Eria  155 Eriachaenium  603 Eriachne  209 Erianthecium  208 Erianthemum  430 Eriastrum  485 Eriaxis  154 Erica  509 Ericales 479 Ericameria  605 Erichsenia  251 Ericksonella  159 Ericoideae 509 Erigenia  633 Erigeron  605, 613 Erinacea  251 Erinocarpus  389 Erinus  553 Eriobotrya  263 Eriocapitella  220, 222 Eriocaulon  198 Eriocephalus  605 Eriochilus  159 Eriochlamys  605 Eriochloa  209 Eriochrysis  209 Eriocnema  355 Eriocoelum  373

Eriodes  155 Eriodictyon  531 Eriogonoideae 434 Eriogonum  434 Eriogynia  263 Eriolaena  390 Eriolarynx  536 Eriolobus  263 Erioneuron  209 Eriope  576 Eriophorum  202 Eriophyllum  605 Eriophyton  576 Eriopidion  576 Eriopsis  155 Erioscirpus  202 Eriosema  251 Eriosemopsis  517 Eriosolena  396 Eriosorus  52 Eriospermum  173–4 Eriostemon  375 Eriostylos  450 Eriosyce  469 Eriosynaphe  633 Eriotheca  390 Eriothymus  576 Eriotrix  605 Erira  262 Erisma  350 Erismadelphus  350 Erismanthus  334 Erithalis  517 Eritrichium  531 Erlangea  605 Ernestia  355 Ernodea  517 Erodiophyllum  605 Erodium  342 Erophaca  251 Errazurizia  251 Ertela  375 Eruca  417 Erucaria  417 Erucastrum  417 Erycibe  534 Erycina  155 Erymophyllum  605 Eryngiophyllum  605 Eryngium  633, 636–7 eryngo root  635 Erysimum  417 Erythradenia  605 Erythranthe  580 Erythrina  251, 258, 259 Erythrocephalum  603 Erythrochiton  375 Erythrococca  334 Erythrodes  159 Erythronium  150 Erythropalum  422 Erythropaloideae 422 Erythrophleum  253, 259 Erythrophysa  373 Erythrophysopsis  373 Erythrorchis  154 Erythroselinum  633 Erythroseris  605 Erythrospermum  324 Erythrostemon  253, 259 Erythroxylum 306 Escallonia  615–16 Escalloniales 615 escarole 610 Eschscholzia  216 Eschweilera  486

Escobedia  583 Escontria  469 Esenbeckia  375 Espadaea  536 Espejoa  605 Espeletia  605 Espostoa  469 Espostoopsis  469 essia 487 Esterhazya  583 Esterhuysenia  457 Etaballia  251 Eteriscius  517 Ethulia  605 Etlingera  191, 193 etrog 376 Euadenia  413 Eubotrys  509 Eubrachion  427 Eucalyptopsis  352 Eucalyptus  259, 352, 429 eucalyptus 351–3 Eucarpha  226 Eucephalus  605 Euceraea  330 Euchaetis  375 Eucharis  171 Euchilopsis  251 Euchiton  605 Euchorium  373 Euchresta  251 Euclasta  209 Euclea  492 Euclidium  417 Euclinia  517 Eucnide  476 Eucodonia  551 Eucomis  174 Eucommia  514–5 Eucorymbia  527 Eucrosia  171 Eucryphia  299 Eucrypta  531 Eudema  417 eudicots  10–11, 213 Eugeissona  177 Eugenia  15, 351–3 Euglypha  101 Eulalia  209 Eulaliopsis  209 Euleria  370 Eulobus  349 Eulophia  155 Eulophiella  155 Eulychnia  469 Eumorphia  605 Euodia  375 Euonymopsis  294 Euonymus  294 Eupatoriastrum  605 Eupatorina  605 Eupatoriopsis  605 Eupatorium  603, 605, 614 Euphorbia  15, 101, 333–6, 532 Euphorbioideae 335 Euphorianthus  373 Euphrasia  582–4 Euphronia  321 Euphrosyne  605 Euplassa  226 Eupodium  27 eupolypod 45 eupolyplods  23, 49, 55–6, 60, 63 Eupomatia  105

Euptelea  214 Eureira  287 Euroschinus  370 Eurya  489 Euryale  91 Eurybia  605 Euryblema  155 Eurycentrum  159 Eurychone  155 Eurychorda  204 Eurycoma  380 Eurycorymbus  373 Eurydendron  489 Euryomyrtus  352 Euryops  605 Eurypetalum  250 Eurysolen  576 Eurystyles  159 Eurytaenia  633 Eusideroxylon  112 Eusiphon  564 Eustachys  209 Eustegia  527 Eustephia  171 Eustigma  235 Eustoma  522 Eustrephus  147, 174 Eutaxia  251 Euterpe  177 Eutetras  605 Euthamia  605 Euthystachys  560 Eutrema  417 Euxylophora  375 Evacidium  605 Evandra  202 evening primrose  349 Everardia  202 Everistia  517 everlastings 613 Eversmannia  252 Evodianthus  139 Evodiella  375 Evolvulus  534 Evotella  159 Ewartia  605 Ewartiothamnus  605 Exaculum  522 Exacum  522 Exallage  517 Exarata  570–1 Exbucklandia  235 Exbucklandioideae 235 Excentradenia  318 Excoecaria  335 Excremis  168 Exellia  106 Exellodendron  322 Exhalimolobos  417 Exoacantha  633 Exocarpos  427 Exocarya  202 Exochaenium  522 Exochorda  263 Exodeconus  536 Exomiocarpon  605 Exomis  451 Exorhopala  425 Exostema  517 Exostigma  605 Exostyles  252 Exothea  373 Exotheca  209 eyebright 584 Eysenhardtia  252 Ezosciadium  633  

INDEX Faba  259 Fabales 248 Fabiana  536 Faboideae  249, 251 Facchinia  448 Facelis  605 Facheiroa  469 Fadenia  451 Fadogia  517 Fadogiella  517 Fagales 276 Fagaropsis  375 Fagonia  247 Fagopyrum  435 Fagraea  522–3 Faguetia  370 Fagus  278 Faidherbia  253 fairy potatoes  464 fairy wing  220 Falcaria  633 Falcataria  253 Falcatifolium  82 Falconeria  335 Falkia  534 Fallopia  435 Fallugia  263 false aralia  629 false black pepper  494 false buck’s-beard  239 false cardamom  193 false cilantro  636 false flax  419 false yam  513 fameflower  466–7 Faradaya  577 Faramea  517 Farfugium  605, 614 Fargesia  208 Farmeria  311 Faroa  522 Farquharia  527 Farrago  209 Farsetia  417 Fascicularia  195 Fatoua  274 ×Fatshedera  631 Fatsia  630–1 Faucaria  457 Faucherea  490 Faujasia  605 Faujasiopsis  605 Faurea  226 fava bean  254 Favratia  592 Faxonia  605 Feddea  605 Fedia  623 Feeria  592 Fegimanra  370 feijoa 352 Feldstonia  605 Felicia  605, 613 Feliciadamia  355 Fendlera  473 Fendlerella  473 Fenerivia  106 Fenestraria  457 Fenestratarum  119 Fenixia  605 fennel  133, 610, 635–7 fenugreek 256 Ferdinandusa  517 Feretia  517 Fergania  633 Fergusonia  517

Fernaldia  527 Fernandezia  155 Fernandoa  567 Fernelia  517 ferns  1–3, 9–12, 14, 22 Fernseea  195 ferntree 374 Ferocactus  469 Feroniella  375 Ferraria  167 Ferreyranthus  605 Ferreyrella  605 Ferrocalamus  208 Ferula  633, 636–7 Ferulago  633 Ferulopsis  633 Fessia  173–4 Festuca  208, 211 feverbark 589 feverbush 515 feverfew 612 Fevillea  287 Fezia  417 Fibigia  417 Fibraurea  220 Ficalhoa  487 Ficaria  218 Ficinia  202 Ficus  274, 386 fiddleheads  30, 49, 55 fiddlewood  573, 578 fidèle, bois  573, 578 Fiebrigiella  252 field eryngo  635 field horsetail  24 field mint  577 Fieldia  551 fig  274, 458 figwort  555, 557 Filago  605 Filarum  119 filbert  282 Filetia  564 Filgueirasia  208 Filicium  373 Filifolium  605 Filipendula  263 Fillaeopsis  253 Fimbristylis  202 finger lime  376 Fingerhuthia  209 fingerroot  193 Finlaysonia  527 Finschia  226 Fioria  389 firecracker flower  565 firecracker plant  555 firevine  535 fireweed  349 Firmiana  390 Fischeria  527 fish poison tree  487 Fissicalyx  252 Fissistigma  106 Fitchia  605, 615 Fittingia  494 Fittonia  564 Fitzalania  106 Fitzroya  84 Fitzwillia  605 Flabellaria  318 Flabellariopsis  318 Flacourtia  330, 383 Flagellaria  204 Flagenium  517 flageolet beans  254

flamboyant  255, 257 flame lily  146 flame vine  568 flannel flower  637 Flaveria  605 flax  170, 337, 338, 611 fleabane  612–13 Fleischmannia  605 Fleischmanniopsis  605 Flemingia  252 Flexanthera  517 Flindersia  375 flixweed  419 Floerkea  406 Florestina  605 florist’s cinerarias  613 florist’s mimosa  259 Floscaldasia  605 Floscopa  180 Flosmutisia  605 Flourensia  605 flour-sack tree  404 flowering cherries  267 Floydia  226 Flueggea  341 fluted pumpkin  289 Flyriella  605 Fockea  527 fodder beet  451 Foeniculum  633, 635, 637 Foetidia  486 Fokienia  84 Foleyola  417 fonio 210 Fontainia  335 Fontanesia  546 fool’s parsley  637 Forchhammeria  411–3 Forcipella  564 Fordia  252 Fordiophyton  355 Forestiera  546 Forgesia  616 forget-me-not  529, 532 Forsskaolea  275 Forstera  595–6 Forsteronia  527 Forsythia  546 Fortunearia  235 Fortunella  375 Fortuynia  417 Fosbergia  517 Fosterella  195 Fothergilla  235 Fouquieria  484 four o’clock flower  461 Foveolina  605 fox and cubs  614 foxglove  554–5, 584 Fragaria  15, 263, 265 Frailea  469 Franciscodendron  390 Francoa  344 frangipani  528, 629 Frankenia  431 frankincense 367–8 Franklandia  226 Franklin’s shrub  497 Franklinia  496 Frantzia  287 Frasera  522 Fraunhofera  294 Fraxinus  546 Freesia  167 Fremontodendron  390 French marigold  613

French rhubarb  635 Freycinetia  140 Freylinia  555 Freziera  489 Fridericia  567 Friedrichkarlmeyeria  417 Friesodielsia  106 fringe tree  547 fringecups 239 Frithia  457 Fritillaria  149–50 Fritzschia  355 Froelichia  450 Froelichiella  450 Froesia  307 Froesiochloa  208 Froesiodendron  106 frogbit  123, 125 frogfruit 573 Frommia  633 Frondaria  155 Froriepia  633 frosted mint  578 fruit of wisdom  188 Fryxellia  390 Fuchsia  349 fuchsia 348–9 Fuernrohria  633 Fuerstia  577 Fuertesia  476 Fuertesiella  159 Fuertesimalva  390 Fuirena  202 fuki 610 fukien tea  532 Fulcaldea  603, 612 Fumana  398 Fumaria  216 Fumarioideae 216 Fumariola  216 fumbwa 77 Funastrum  527 Funifera  396 Funkiella  159 Funtumia  527 Furcraea  174 Furtadoa  119 Fusaea  106 Fuscospora  277 Fusifilum  174 Fusispermum  325 Futabanthus  106 fynbos (family)  202   Gabon chocolate  303 Gabonius  250 Gadoria  553 Gaertnera  517 Gaga  53 Gagea  150 Gagnebina  253 Gagnepainia  192 Gahnia  202 Gaiadendron  430 Gaillardia  605, 614 Gaimardia  204 Galactia  252 Galactites  605 Galactophora  527 Galantharum  119 Galanthus  170–1 Galatella  605 galatsida 610 Galax  499 Galbulimima  104 Galeana  605

Galeandra  155 Galearia   302 Galearis  159 Galega  252, 259 Galenia  457 Galeoglossum  159 Galeola  154 Galeomma  605 Galeopsis  577 Galeottia  155 Galeottiella  159 Galianthe  517 Galiniera  517 Galinsoga  605, 610 galip nut  368 Galipea  375 Galitzkya  417 Galium  517 gallant soldiers  610 Gallardoa  318 Gallesia  460 Gallienia  517 Galopina  517 Galphimia  318 Galpinia  346 Galvezia  553 Gamanthus  451 Gambelia  554 gambier 520 Gamblea  630 gamboge 310 Gamocarpha  601 Gamochaeta  605 Gamochaetopsis  605 gampi 396 Ganguelia  517 Ganophyllum  373 Gaoligongshania  208 Garberia  605 Garcia  335 Garciadelia  334 Garcibarrigoa  605 Garcinia  309 garden abelia  625 garden asparagus  174 garden bacopa  557 garden balsam  480 garden nasturtium  403 garden pea  254 garden phlox  485 garden pyrethrum  613 garden rue  378 garden verbena  573 Gardenia  517 Gardeniopsis  517 Gardneria  523 Gardnerina  605 Garhadiolus  605 garlic 171 garlic chives  171 garlic mustard  419 garlic vine  568 Garnieria  226 Garnotia  209 Garrettia  575, 577 Garrya  515 Garryales 514 Garuga  368 Garuleum  605 Gasteranthus  551 Gasteria  168, 170 Gastonia  629 Gastoniella  52 Gastridium  208 Gastrochilus  155

Gastrocotyle  531 Gastrodia  152, 155 Gastrolepis  585 Gastrolobium  252 Gastrorchis  155 Gaudichaudia  318 Gaudinia  208 Gaulettia  322 Gaultheria  509 gaura 349 Gaussia  177 Gavilea  159 Gaya  390 gayfeather 614 Gaylussacia  509 Gayophytum  349 Gazania  605, 613 Gearum  119 Geesinkorchis  155 geiger tree  532 Geigeria  605 Geijera  375 Geissanthus  494 Geissaspis  252 Geissois  299 Geissolepis  605 Geissomeria  564 Geissorhiza  168 Geissospermum  527 Geitonoplesium  147, 167–8 Gelasine  167 Geleznowia  375 Gelidocalamus  208 Gelsemium  523–4 gemsbok cucumber  288 Geniostemon  522 Geniostoma  523 genip 520 Genipa  517 Genista  252, 256–8 Genistidium  252 Genlisea  569–70 Gennaria  159 Genoplesium  159 gentian  520, 522 Gentiana  522 Gentianales 516 Gentianella  522 Gentianopsis  522 Gentianothamnus  522 Gentingia  517 Geobalanus  322 Geocarpon  448 Geocaryum  633 Geocaulon  427 Geocharis  192 Geochloa  209 Geococcus  417 Geodorum  155 Geoffroea  252, 255 Geogenanthus  180 Geomitra  134 Geonoma  177 Geophila  517 Georgeantha  206 Geosiridoideae 167 Geosiris  165, 167 Geostachys  192 Geraea  605 Geraniales 342 Geranium  342–3 geranium oil  343 Gerardiina  583 Gerbera  603, 613 Gereaua  373

Plants of the World

769

INDEX Germainia  209 germander 578 Geropogon  605 Gerrardanthus  287 Gerrardina  383 Gerritea  209 Gesneria  551 Gesnerioideae 551 Gesnouinia  275 Gethyllis  170–1 Geum  263 Gevuina  225–6 Ghaznianthus  433 gherkins 287 Ghikaea  583 giant ash  353 giant balsam  480 giant granadilla  328 giant hogweeds  637 giant hyssop  578 giant lily  150 giant nato  259 giant scabious  625 giant taro  119 Gibasis  180 Gibasoides  180 Gibbaeum  457 Gibbaria  605 Gibbsia  276 Gibsoniothamnus  571 Gigantochloa  208 Gigasiphon  250 Gilberta  605 Gilbertiella  106 Gilbertiodendron  251 Gilgiochloa  209 Gilia  485 Giliastrum  485 Gillbeea  299 Gillenia  263 Gillespiea  517 Gilletiodendron  251 Gilliesia  171 Gilliesieae  171 Gilmania  434 Gilruthia  605 gin  84, 167, 193, 265, 378, 519, 636 Ginaloa  427 ginger  101, 191–3, 378, 610 Gingidia  633 Ginkgo  71, 73, 75 Ginkgoales  10–11, 75 Ginkgoidae 75 Ginoria  346 ginseng 630 Girardinia  276 girasole 611 Girgensohnia  451 Gironniera  273 Gisekia  446, 456, 459 Githopsis  592 Givotia  335 Gjellerupia  424 Glabrella  551 Gladiolimon  433 Gladiolus  167 Gladiopappus  603 Glandonia  318 Glandora  531 Glandularia  573 Glastaria  417 Glaucidioideae 220 Glaucidium  220 Glaucium  216

770

Glaucosciadium  633 Glaziophyton  208 Gleadovia  583 Gleasonia  517 Glebionis  605, 610, 613 Glechoma  577–8 Glechon  577 Gleditsia  253, 256, 259 Glehnia  633 Gleichenella  32 Gleichenia  32 Gleicheniales 10–11, 31–2 Glekia  555 Glenniea  373 Glia  633 Glinus  462 Glionnetia  517 Gliricidia  252, 259 Glischrocaryon  243 Glischrocolla  358 Globba  191–3 globe amaranth  452 globe artichoke  610 globe daisy  555 globe mallow  392 globe thistles  614 globeflower  222 Globimetula  430 Globularia  554 Globulariopsis  555 Globulostylis  517 Glochidotheca  633 Gloeocarpus  373 Gloeospermum  325 Glomera  155 Glomeropitcairnia  195 Gloriosa  146 glorybower 578 Glossarion  603 Glossocalyx  108 Glossocardia  605 Glossocarya  577 Glossochilus  564 Glossodia  159 Glossonema  527 Glossopappus  605 Glossopetalon  363 Glossostemon  389 Glossostigma  580 Glossostipula  517 Glottiphyllum  457 Gloveria  294 Gloxinia  551 gloxinias 551 Gluema  490 Glumicalyx  555 Gluta  370 Glyceria  208 Glycine  252, 254, 258 Glycosmis  375 Glycydendron  335 Glycyrrhiza  252, 256 Glyphaea  389 Glyphochloa  209 Glyptopetalum  294 Glyptopleura  605 Glyptostrobus  84 Gmelina  576, 578 Gnaphaliothamnus  605 Gnaphalium  605 Gnephosis  605 Gnetales  10–11, 77 Gnetidae 71 gnetoids  71–2, 75, 79 Gnetum  77

Christenhusz, Fay & Chase

Gnidia  395–6 Gnomophalium  605 Goa bean  255 gobō  610 Gochnatia  603 Gochnatioideae 602 godetia 349 Godmania  567 Goeppertia  190 Goethalsia  389 Goethea  389 Goetzea  536 goji berry  539 gold apple  492 Goldbachia  417 golden alexanders  637 golden chamomile  612 golden currant  238 golden dewdrop  573 golden drop  532 golden grass  198 golden leastdaisy  612 golden ray  614 golden samphire  609 golden saxifrage  239 golden trumpet vine  528 goldenbush 614 goldenrain tree  374 goldenrod 614 goldenseal 222 goldenstar 613 goldfields  611 Goldmanella  605 goldshower 318 goldstaff 614 Gomesa  152, 155 Gomphandra  585 Gomphichis  159 Gomphocalyx  517 Gomphogyne  287 Gompholobium  252 Gomphostemma  577–8 Gomphostigma  555 Gomphrena  450 Gonatogyne  341 Gonatopus  119 Gonatostylis  159 Gongora  155 Gongrodiscus  373 Gongronema  527 Gongrospermum  373 Gongrostylus  605 Gongylocarpus  349 Gongylolepis  603 Gongylosciadium  633 Gongylotaxis  633 Gonialoë  168 Goniocaulon  605 Goniodiscus  294 Goniolimon  433 Gonioma  527 Goniophlebium  70 Goniorrhachis  251 Goniothalamus  106 gonnabos 396 Gonocalyx  509 Gonocarpus  243 Gonocaryum  586 Gonocytisus  252 Gonolobus  527 Gonospermum  605 Gonostegia  276 Gontscharovia  577 Gonystylus  396 Gonzalagunia  517 Gonzalezia  605

good King Henry  452 Goodallia  396 Goodenia  599–600 Goodia  252 Goodmania  434 Goodyera  159 Goodyerinae  151 gooseberry  237–8, 341, 470, 504, 538 gooseberry tree  371 goosefoot 452 gooseweed 542 Gorceixia  605 Gordonia  496 Gorgonidium  119 Gorteria  605 Gosela  555 Gossia  352 Gossweilerodendron  251, 256 Gossypioides  390 Gossypium  42, 385, 390, 392 gotu kola  636 Gouania  271 Gouinia  209 goumi 269 Goupia  326 goutweed 636 Govenia  155 Goyazia  551 Goyazianthus  605 Graderia  583 Graellsia  417 Graffenrieda  355 Grahamia  468 grains of paradise  193 Grajalesia  461 Grammangis  155 Grammatophyllum  155 Grammatotheca  592 Grammitis  63, 70 Grammosciadium  633 Grammosolen  536 granadilla  327, 328 Grandidiera  324 Grandiphyllum  155 Grangea  605 Grangeopsis  605 Grangeria  322 granite gooseberry  238 Granitites  271 grape  220, 244–5, 341, 435 grape hyacinth  174 grapefruit 376–7 Graphandra  564 Graphephorum  208 Graphistemma  527 Graphistylis  605 Graphorkis  155 Graptopetalum  240–1 Graptophyllum  564–5 grass trees  170 grasses  87, 206 Gratiola  554 Gratwickia  605 Grauanthus  605 Gravesia  355 Grayia  451 Grazielia  605 Grazielodendron  252 great pignut  635 greater celandine  216 greater galangal  193 Greek basil  578

green algae  3 green cardamom  193 green poison-berry  540 greencurrant 238 Greenea  518 greengage 264 Greeniopsis  518 Greenmaniella  605 Greenwayodendron  106 Greenwoodiella  159 Gregbrownia  195 Greigia  195 Grenacheria  494 grenadine 347 Greslania  208 Grevea  542 Grevillea  226 Grevilleoideae 226 Grewia  389 Grewioideae 389 Greyia  344 Grias  486 Grielum  365, 388 Griffinia  171 Griffithella  311 Griffithsochloa  209 Griffonia  250, 257 Grimmeodendron  335 Grindelia  605, 612 Grisebachianthus  605 Griselina  628 Grisollea  585 Grobya  155 Groenlandia  129 Gronovia  476 groseille 238 Grosourdya  155 Grossera  335 Grossularia  238 Grosvenoria  605 ground elder  636 ground ivy  577–8 groundnut 254–5 grumichama 352 Guacamaya  197 guacimilla 273 Guadalupe palm  178 Guadua  208 Guaduella  206 Guaiacum  247 Guamatela  361 guanábana 106 Guanchezia  155 Guapira  461 guar gum  256 guaraná 374 Guardiola  605 Guarea  381 Guarianthe  155 guarrie bark  492 Guatteria  106 guava  352, 520 guaya 373 guayacán 257 guayamochil 255 Guayania  605 guayato 527 guayule 612 guayusa 589 Guazuma  386, 389 Gueldenstaedtia  252 Guelder rose  620 Guettarda  518 Guevaria  605 Guiana chestnut  393 Guianodendron  252

Guibourtia  251, 256 Guichenotia  389 Guiera  346 Guihaia  177 Guihaiothamnus  518 Guilandina  253, 258 Guilfoylia  260 Guilleminea  450 Guillonea  633 Guindilia  373 Guinea henwood  460 Guinea millet  210 Guinea peach  520 Guinetia  253 Guioa  373 Guiraoa  417 Guizotia  605, 611 gulf greytwig  428 gum ammoniacum  636 gum arabic  256 gum copal  256 gum galbanum  636 gum opopanax  636 gum tragacanth  256 Gumillea  364 gumweed 612 gumwood 614 Gundelia  605 Gundlachia  605 Gunillaea  592 Gunnarella  155 Gunnera  230 Gunnerales 229 Gunnessia  527 Gunniopsis  457 Gurania  287 gurjun oil  400 Gustavia  486 Gutenbergia  605 Guthriea  324 Gutierrezia  605 Guynesomia  605 Guyonia  355 Guzmania  195 Gyminda  294 Gymnacranthera  102 Gymnadenia  159 Gymnanthemum  605 Gymnanthera  527 Gymnanthes  335 Gymnarrhena  606 Gymnarrhenoideae 602 Gymneia  577 Gymnocalycium  469 Gymnocarpium  55–6 Gymnocarpos  448 Gymnocladus  253, 255–6, 259 Gymnocondylus  606 Gymnocoronis  606 Gymnodiscus  606 Gymnogrammitis  68 Gymnolaena  606 Gymnomyosotis  531 Gymnopentzia  606 Gymnophyton  633 Gymnopodium  435 Gymnopogon  209 Gymnoschoenus  202 Gymnosiphon  134 gymnosperm  1–3, 10–12, 14, 22, 71 Gymnosperma  606 Gymnospermium  219 Gymnospora  262 Gymnosporia  294

INDEX Gymnostachyoideae  118–19 Gymnostachys  118–19 Gymnostachyum  564 Gymnostemon  380 Gymnostephium  606 Gymnosteris  485 Gymnostoma  281 Gymnotheca  98 Gynandra  522 Gynandriris  167 Gynandropsis  414 Gynatrix  390 Gynerium  209 Gynocardia  324 Gynochthodes  518 Gynocraterium  564 Gynoglottis  155 Gynostemma  287 Gynotroches  305 Gynoxys  606 Gynura  606, 610, 614 Gypothamnium  603 Gypsacanthus  564 Gypsophila  448 Gyptidium  606 Gyptis  606 Gyranthera  390 Gyrinops  396 Gyrocarpoideae 110 Gyrocarpus  110, 386 Gyrocaryum  531 Gyrocheilos  551 Gyrodoma  606 Gyrogyne  551 Gyrostemon  411 Gyrostipula  518 Gyrotaenia  276   H eliodendron  373 Haageocereus  469 Haastia  606 Habenaria  152, 159 Haberlea  551 Hablitzia  451 Habranthus  171 Habrochloa  209 Habroneuron  518 Habropetalum  441 Habrosia  448 Hachettea  425 Hackelia  531 Hackelochloa  209 Hacquetia  633, 637 Haeckeria  606 Haegiela  606 Haemanthus  171 Haematocarpus  218 Haematodendron  102 Haematostaphis  370 Haematostemon  334 Haematoxylum  253, 257, 386 Haemodoroideae 184 Haemodorum  184 Haenianthus  546 Hagenbachia  174 Hagenia  263 Hagsatera  155 Hainania  389 Hainardia  208 hairy chervil  637 Haitia  346 Hakea  226 Hakonechloa  209 Halacsya  531

Halanthium  451 Halarchon  451 Halenia  522 Halerpestes  220 Halesia  500 Halfordia  375 Halgania  531 Halimium  398 Halimocnemis  451 Halimodendron  252 Halimolobos  417 Halleorchis  159 Halleria  560 Hallianthus  457 hallucinogen 102 Halocarpus  82 Halocharis  451 Halocnemum  451 Halodule  129, 131 Halogeton  451 Halopegia  190 Halopeplis  451 Halophila  125 Halopyrum  209 Haloragis  243 Haloragodendron  243 Halosarcia  451 Halosciastrum  633 Haloselinum  633 Halosicyos  287 Halostachys  451 Halothamnus  451 Haloxylon  451 Hamadryas  220 Hamamelidoideae 235 Hamamelis  235 Hamburg parsley  636 Hamelia  518 Hamilcoa  335 Hammarbya  155 Hammatolobium  252 hammer orchid  152 Hampea  390 Hanabusaya  592 Hanburia  287 Hancea  334 Hanceola  577 Hancockia  155 Hancornia  527 Handelia  606 handkerchief tree  472 Handroanthus  567–8 Hanguana  179–80, 204 Haniffia  192 Hannafordia  389 Hannoa  380 Hannonia  171 Hansenia  633 Hanseniella  311 Hanslia  252 Hapaline  119 Hapalorchis  159 Haplanthodes  564 Haplocarpha  606 Haplochorema  192 Haploclathra  310 Haplocoelopsis  373 Haplocoelum  373 Haploësthes  606 Haplolobus  368 Haplopappus  606 Haplophyllum  375 Haplophyton  527 Haplopteris  54 Haplorhus  370 Haplormosia  252

Haplosciadium  633 Haplosphaera  633 Haplospondias  370 Haplostachys  577 Haplostichanthus  106 Haplothismia  134 happy tree  472 Haptanthus  227–8 Haptocarpum  414 Haptotrichion  606 Harbouria  633 Hardenbergia  252, 259 Hardwickia  251 hardy gloxinia  568 hare’s ears  637 Harfordia  435 haricots 254 haricots verts  254 Harleya  606 Harleyodendron  252 harmal 366 Harmandia  422 Harmogia  352 Harmonia  606 Harmsia  390 Harmsiodoxa  417 Harmsiopanax  630 Harnackia  606 Harpachne  209 Harpagonella  531 Harpagophytum  562–3 Harpalyce  252 Harpephyllum  370 Harperia  204 Harperocallis  121 Harpochilus  564 Harpochloa  209 Harpullia  373 Harrimanella  509 Harrisia  469 Harrisonia  375 Harrysmithia  633 harsinger tree  547 Hartleya  585 Hartliella  558 Hartogiopsis  294 hartwort 636 Hartwrightia  606 Harungana  313 Harveya  583 Hasseltia  330 Hasseltiopsis  330 Hasteola  606 Hastingsia  174 Hatiora  469 Hatschbachiella  606 hau 393 Haumania  190 Haumaniastrum  577 hausa potato  578 Haussknechtia  633 Hauya  349 Havardia  253 Hawaiian goosefoot  452 Hawaiian palm  593 Hawaiian silverswords 615 Haworthia  168, 170 Haworthiopsis  168 hawthorn 266 Haya  448 Haydenoxylon  294 Hazardia  606 hazel  226, 235, 282 hazelnuts  280, 282 Hazomalania  110

he cabbage  614 headache tree  578 heather  347, 506, 509 Hebanthe  450 Hebecarpa  262 Hebeclinium  606 Hebenstretia  555 Hebepetalum  337 Heberdenia  494 Hebestigma  252 Hecastocleis  603 Hecatostemon  330 Hechtia  195 Hecistopteris  54 Heckeldora  381 Hedbergia  583 Hedeoma  577 Hedera  629–31 Hederorkis  155 hedge hyssop  555 Hedosyne  606 Hedraianthera  294 Hedstromia  518 Hedycarya  111 Hedychium  192–3 Hedyosmum  114 Hedyotis  518 Hedypnois  606 Hedysarum  252 Hedyscepe  177 Hedythyrsus  518 Heeria  370 Hegnera  252 Heimia  346 Heinsenia  518 Heinsia  518 Heisteria  422 Hekistocarpa  518 Hekkingia  325 Heladena  318 Helanthium  122 Heldreichia  417 Helenium  606, 613–14 Helia  522 Heliamphora  501 Heliantheae  602 Helianthella  606, 614 Helianthemum  398 Helianthostylis  274 Helianthus  606, 611, 613–15 Helicanthes  430 Helichrysopsis  606 Helichrysum  386, 606, 612–14 Helicia  226 Helicilla  450 Heliciopsis  226 Helicodiceros  119 Heliconia  185–6 Helicostylis  274 Helicteres  389 Helicteroideae 389 Helicteropsis  390 Helictonema  294 Helictotrichon  208 Helietta  375 Helinus  271 Heliocarpus  389 Heliocauta  606 Heliomeris  606 Heliophila  417 Heliopsis  606, 614 heliotrope 614 Heliotropium  531 Helixanthera  430

hellebore 222 Helleborus  220, 222 Hellenia  191 Hellenocarum  633 Helleriella  155 Hellmuthia  202 Helmholtzia  182 Helminthostachys  25 Helminthotheca  606 Helmiopsiella  390 Helmiopsis  390 Helmontia  287 Helogyne  606 Helonias  143 Heloniopsis  143 Helonoma  159 Helosis  425 Helwingia  585, 587–8 Hemarthria  209 Hemerocallidoideae 168 Hemerocallis  8, 160, 168 Hemiandra  576 Hemiarrhena  558 Hemiboea  551 Hemichaena  580 Hemichroa  450 Hemicrambe  417 Hemidesmus  527 Hemidictyum  57–8 Hemigenia  576 Hemigraphis  564–5 Hemilophia  417 Hemimeris  555 Hemionitis  53 Hemiorchis  192 Hemiphora  576 Hemiphragma  554 Hemiphylacus  173–4 Hemipilia  159 Hemipogon  527 Hemiptelea  272 Hemiscola  414 Hemiscolopia  330 Hemisorghum  209 Hemistylus  276 Hemithrinax  177 Hemitomes  509 Hemizonella  606 Hemizonia  606 hemlock 636–7 hemlock-parsley 635 hemlock water-dropwort 636 hemp  188, 258, 272–3, 291, 528 hemp-agrimony 614 hempvine 614 Hemradenia  296 Hemsleya  287 henbane 540 Henckelia  551 henequen 174 Henleophytum  318 henna 347 Hennecartia  111 Henonia  450 Henoonia  536 Henophyton  417 Henradia  208 Henricksonia  606 Henriettea  355 Henriquezia  518 Henrya  564 Hensmania  168 Hepatica  220, 222 Heppiella  551

Heptacodium  622–23, 625 Heptanthus  606 Heptaptera  633 Heracleum  633, 637 herb of grace  378 herb robert  343 Herbertia  167 Herbstia  450 Herderia  606 Hereroa  457 Herissantia  390 Heritiera  390 Hermannia  389 Hermas  623, 633 Hermbstaedtia  450 Herminium  159 Hermodactylus  167 Hernandia  110 Hernandioideae 110 Herniaria  448 Herodotia  606 Herpetacanthus  564 Herpetospermum  287 Herpolirion  168 Herpysma  159 Herpyza  252 Herrania  389 Herreranthus  606 Herreria  174 Herreriopsis  174 Hertia  606 Hesperalbizia  253 Hesperaloë  174 Hesperantha  167 Hesperelaea  546 Hesperevax  606 Hesperidanthus  417 Hesperis  417 Hesperocallis  174 Hesperochiron  531 Hesperocnide  276 Hesperocyparis  84 Hesperolaburnum  252 Hesperolinon  337 Hesperomannia  606 Hesperomecon  216 Hesperomeles  263 Hesperothamnus  252 Hesperoxiphion  167 Hesperoyucca  174 Hesperozygis  577 Hessea  171 Hestia  119 Hetaeria  159 Heterachne  209 Heteracia  606 Heteradelphia  564 Heteranthelium  208 Heteranthemis  606 Heteranthera  183 Heteranthia  536 Heteranthoecia  209 Heterocentron  355 Heterochaenia  592–4 Heterocodon  592 Heterocoma  606 Heterocondylus  606 Heterocypsela  606 Heteroderis  606 Heterolepis  606 Heteromeles  263 Heteromera  606 Heteromma  606 Heteromorpha  632, 633 Heteropanax  630

Plants of the World

771

INDEX Heteropholis  209 Heterophragma  567 Heterophyllaea  518 Heteroplexis  606 Heteropogon  209 Heteropolygonatum  174 Heteropsis  119 Heteropterys  318 Heteropyxis  351 Heterorhachis  606 Heterosamara  262 Heterosavia  341 Heterosmilax  148 Heterospathe  177 Heterosperma  606 heterosporous ferns  36–8 Heterostachys  451 Heterostemma  527 Heterostemon  251 Heterothalamus  606 Heterotheca  606 Heterotoma  592 Heterotrichum  355 Heuchera  239 heuningbos 256 Hevea  335 Hewittia  534 Hexacyrtis  146 Hexalectris  155 Hexalobus  106 Hexapterella  134 Hexasepalum  518 Hexaspora  294 Hexatheca  551 Heynea  381 Heynella  527 Heywoodia  341 Hibbertia  231 Hibiscadelphus  390 Hibiscus  390 Hickelia  208 hickory 280 Hicksbeachia  226 Hidalgoa  606 Hieracium  603, 606 Hieris  567 Hiernia  583 Hierochloe  208 Hieronyma  340 Hieronymiella  170–1 Hieronymusia  239 Hilaria  209 Hildaea  209 Hildebrandtia  534 Hildegardia  390 Hilleria  460 Hillia  518 Hilliardia  606 Himalaiella  606 Himalayacalamus  208 Himalayan balsam  480 Himalocodon  592 Himalrandia  518 Himantoglossum  159 Himatanthus  527 Hindsia  518 Hinterhubera  606 Hintonella  155 Hintonia  518 Hionanthera  346 Hippeastreae  171 Hippeastrum  171 Hippeophyllum  155 Hippia  606 Hippobroma  592–3 Hippobromus  373

772

Hippocastanoideae 373 Hippocratea  294 Hippocrepis  252, 259 Hippolytia  606 Hippomane  335 Hippophaë  269 Hippotis  518 Hippuris  554 Hiptage  318 Hiraea  318 Hirania  373 Hirpicium  606 Hirschfeldia  417 Hirtella  322 Hispanic thyme  573 Hispaniolanthus  413 Hispidella  606 Histiopteris  49 Hitchcockella  208 Hitchenia  191–2 Hladnikia  633 hoary basil  578 hoary cress  419 hoary thyme  578 Hochreutinera  390 Hockinia  522 Hodgkinsonia  518 Hodgsonia  287 Hodgsoniola  168 Hoehnea  577 Hoehneella  155 Hoehneophytum  606 Hoffmannanthus  606 Hoffmannia  518 Hoffmanniella  606 Hoffmannseggia  253, 256 Hofmeisterella  155 Hofmeisteria  606 hog potato  256 hogbrake 614 hogweed 637 Hohenackeria  633 Hohenbergia  195 Hohenbergiopsis  195 Hoheria  390 Hoita  252 Holacantha  380 Holandrea  633 Holarrhena  527 Holcoglossum  155 Holcolemma  209 Holcus  208 Holigarna  370 Hollandaea  226 Hollermayera  417 Hollisteria  435 holly  318, 336, 588–9, 617 holly grape  220 hollyhock 393 Holmbergia  451 Holmgrenanthe  554 Holmgrenia  349 Holmskioldia  577–8 Holocalyx  252 Holocarpha  606 Holocheila  577 Holocheilus  603 Holochlamys  119 Holodiscus  263 Holographis  564 Hololachna  432 Hololeion  606 Hololepis  606 Holoregmia  561 Holoschkuhria  606 Holostemma  527

Christenhusz, Fay & Chase

Holosteum  448 Holostylis  101 Holothrix  159 Holozonia  606 Holstianthus  518 Holttumochloa  208 Holubia  562 holy flax  611 holy thistle  610 Holzneria  554 Homalanthus  335 Homalium  330 Homalocalyx  352 Homalocarpus  633 Homalocladium  435 Homalodiscus  412 Homalomena  119 Homalopetalum  155 Homalosciadium  629, 632–3 Homalosorus  57 Homalospermum  352 Homeria  167 Homocodon  592 Homogyne  606 Homolepis  209 Homollea  518 Homolliella  518 Homonoia  334 Homopholis  209 Homoranthus  352 Homozeugos  209 Honckenya  448 Hondurodendron  422 honesty 419 honey locust  256 honeybells 557 honeyberry 623 honeybush 256 honeyherb 573 honeysuckle  560, 568, 621, 623, 625 honeywort 532 Hoodia  527 Hooglandia  299 hopbush 374 Hopea  400 Hopkinsia  204 Hoplestigma  531 Hoplophyllum  606 Hoppea  522 hops 273 hop-tree 374 Horaninovia  451 horchata de chufa  202 Hordeleymus  208 Hordeum  208, 210 horehound 578 Horichia  155 Hormathophylla  417 Horminum  577–8 hornbeam 282 Hornea  373 horned melon  288 horn-of-plenty 623 Hornschuchia  106 Hornstedtia  192–3 Hornungia  417 hornwort  3, 212 horny goat weed  220 Horovitzia  405 horse chestnut  374 horse mango  371 horse mint  577 horse-heal 611 horseradish 419

horseradish tree  404 horsetails  24, 202 Horsfieldia  102 Horsfordia  390 Horstrissea  633 hortensia 473–4 Hortia  375 Hortonia  111 Horvatia  155 Horwoodia  417 Hosackia  252 Hosea  577 Hosiea  513 Hosta  174 Hottea  352 hottentot-fig  458 Hottonia  494 Houlletia  155 hound’s tongue  532 houseleek 241 Houssayanthus  373 Houstonia  518 Houttuynia  98 Hovea  252, 259 Hovenia  271 Hoverdenia  564 Howardia  101 Howea  177 Howellanthus  531 Howellia  592 Howelliella  554 Howittia  390 Hoya  525, 527 Hsenhsua  159 Hua gabonii  296 Huanaca  633 huanita 532 Huarpea  603 huauzontle 452 Hubbardia  209 Hubbardochloa  209 Huberantha  106 Huberia  355 Huberodendron  390 Huberopappus  606 Hubertia  606 huckleberry  509, 539 Hudsonia  398 Huernia  527 Huertea  361, 384 Huerteales 382 Hughesia  606 Hugonia  337 Hugonioideae 337 Huidobria  476 Hulemacanthus  564 Hullettia  274 Hulsea  606 Hulteniella  606 Humbertacalia  606 Humbertia  534 Humbertianthus  390 Humbertiella  390 Humbertiodendron  319 Humbertioturraea  381 Humbertochloa  206 Humblotiodendron  375 Humboldtia  251 Humeocline  606 Humiria  323 Humiriastrum  323 Humularia  252 Humulus  273 Hunga  322 hungry rice  210 Hunnemannia  215–16

Hunteria  527 Huntleya  155 Hunzikeria  536 Huodendron  500 Huperzia  19 Hura  335 hutchinsia 419 Hutchinsonia  518 Huttonaea  159 hyacinth  7, 171, 174 Hyacinthella  173–4 Hyacinthoides  173–4 Hyacinthus  7–8, 173–4 Hyaenanche  539 Hyalis  603 Hyalocalyx  327 Hyalochlamys  606 Hyalocystis  534 Hyalolaena  633 Hyaloseris  603 Hyalosperma  606 Hybanthopsis  325 Hybanthus  325 Hybridella  606 Hydatella  14, 89 Hydnocarpus  324 Hydnophytum  518 Hydnora  100–1 Hydnoroideae 100–1 Hydrangea  473 Hydrangeoideae 473 Hydrastidoideae 220 Hydrastis  220, 222 Hydriastele  177 Hydrilla  125 Hydrobryum  311 Hydrocharis  125 Hydrocharitites  124 Hydrochorea  253 Hydrocleys  122–3 Hydrocotyle  629–31 Hydrocotyloideae 629–30 Hydrodiscus  311 Hydrogaster  389 Hydroidea  606 Hydrolea 543 Hydromystria  124 Hydropectis  606 Hydrophilus  204 Hydrophylax  518 Hydrophyllum  531 Hydropteridales 37 Hydrothauma  209 Hydrothrix  182–3 Hydrotriche  554 Hygea  551 Hygrochloa  206 Hygrophila  564–5 Hygroryza  206 Hylaea  527 Hylaeanthe  190 Hylandia  335 Hylebates  209 Hylenaea  294 Hylocarpa  323 Hylocereus  469 Hylodendron  251 Hylodesmum  252 Hylomecon  216 Hylophila  159 Hylotelephium  240–1 Hymenachne  209 Hymenaea  249, 251 Hymenandra  494 Hymenasplenium  58 Hymenidium  633

Hymenocallis  170–1 Hymenocardia  340 Hymenocarpos  252 Hymenocoleus  518 Hymenocrater  577 Hymenodictyon  518 Hymenogyne  457 Hymenolaena  633 Hymenolepis  606 Hymenolobium  252 Hymenonema  606 Hymenopappus  606 Hymenophyllales  10, 30 Hymenophyllopsis  39, 43 Hymenophyllum  31 Hymenopus  322 Hymenopyramis  577 Hymenorchis  155 Hymenosporum  629 Hymenostegia  251 Hymenostemma  606 Hymenostephium  606 Hymenothrix  606 Hymenoxys  606 Hyobanche  582–3 Hyophorbe  177 Hyoscyamus  536 Hyoseris  606 Hyospathe  177 Hypacanthium  606 Hypagophytum  241 Hyparrhenia  209 Hypecoum  215–16 Hypelate  373 Hypenia  577 Hyperacanthus  518 Hyperbaena  218 Hypericophyllum  606 Hypericum  313 Hypertelis  455, 462 Hyperthelia  209 Hyphaene  177 Hypobathrum  518 Hypocalymma  352 Hypocalyptus  252 Hypochaeris  606 Hypodaphnis  112 Hypodematioideae 23, 63–4, 68 Hypodematium  64 Hypoderris  67 Hypodiscus  204 Hypoestes  564–5 Hypogomphia  577 Hypolaena  204 Hypolobus  527 Hypolytrum  202 Hypopitys  509 Hypoxidia  163 Hypoxis  163 Hypsela  593 Hypselodelphys  190 Hypseloderma  372 Hypseocharis  342 Hypseochloa  208 Hypserpa  218 Hypsophila  294 Hyptianthera  518 Hyptidendron  577 Hyptis  577–8 Hyssopus  577 Hysterionica  606 Hystrichophora  606 hyuganatsu 376   Ianhedgea  417

INDEX Ianthopappus  603 Iberis  417 Ibervillea  287 Ibicella  561 iboga 528 ibogaine 528 Icacina  513 Icacinales 511 icacos 322 icecream bean  255 Ichang lemon  376 Ichang papeda  376 Ichnanthus  209 Ichnocarpus  527 Ichthyothere  606 Ichtyoselmis  216 Ichtyostoma  564 Icuria  251 Idahoa  417 Idesia  330 Idiopappus  606 Idiospermum  107 Idiothamnus  606 Ifloga  606 Iguanura  177 Ihsanalshehbazia  417 Ikonnikovia  433 Ileostylus  430 Ilex  589 Iliamna  390 Illecebrum  448 Illiciales 94 Illicium  94–5 Illigera  110 illipe nuts  491 Iltisia  606 Imeria  606 Imerinaea  155 Impatiens  15, 479, 480 Imperata  209, 211 Imperatoria  633 Incarum  119 Incarvillea  567–8 incense  112, 256, 368, 378, 396, 398, 427, 528, 636 inchi tree  335 Indian almond  346 Indian bean tree  568 Indian boxwood  315 Indian gooseberry  341 Indian hemp  528 Indian nard  623 Indian paint  532 Indian pink  522, 524 Indian plum  330 Indian rhubarb  239 Indian swampweed  565 Indian tonic  519 Indianthus  190 Indigastrum  252 indigo  257, 419, 436, 528 Indigofera  250, 252, 257 Indobanalia  450 Indocalamus  208 Indocypraea  606 Indodalzellia  311 Indofevillea  287 Indomelothria  287 Indopiptadenia  253 Indopoa  209 Indorouchera  337 Indosasa  208 Indoschultzia  633 Indotristicha  311 Inezia  606

Inga  253, 255, 258 Inhambanella  490 inkberry 540 Inocarpus  252, 255 inoi nut  284 Insitiocarpus  223 interrupted fern  30 Intsia  251 Inula  606, 611–12, 614 Inulanthera  606 inulin 611 Inuloides  606 Inulopsis  606 Inversodicraea  311 Involucrella  518 Io  606 Iocenes  606 Iochroma  536 Iodanthus  417 Iodes  513 iodine bush  452 Iodocephalopsis  606 Iodocephalus  606 Iogeton  606 Ionacanthus  564 Ionactis  606 Ionopsidium  417 Ionopsis  155 Iostephane  606 Iotasperma  606 ipê 568 ipecacuanha plant  519 Ipheion  171 Iphigenia  146 Iphiona  606 Iphionopsis  606 ipoh 274 Ipomoea  534 Ipomopsis  485 ipos 218 Ipsea  155 Iranecio  606 Irania  417 Irenella  450 Irenepharsus  417 Iresine  450 Iriartea  177 Iriartella  177 Iridoideae 167 Iridosma  380 Iris  167, 184 Irlbachia  522 iroko 274 iron weed  614 ironwood  311, 339, 546 ironwort 578 Irvingbaileya  585 Irvingia  303 Irwinia  606 Iryanthera  102 Isabelia  155 Isachne  209 isano oil  423 Isatis  257, 417, 419 Ischaemum  209 Ischnea  606 Ischnogyne  155 Ischnolepis  527 Ischnosiphon  190 Ischyrolepis  204 Iseia  534 Iseilema  209 Isertia  518 Isidodendron  319 Isidorea  518 Iskandera  417

island cotton  392 Ismelia  606 Ismene  171 Isoberlinia  251 Isocarpha  606 Isochilus  155 Isocoma  606, 614 Isodendrion  325 Isodon  577 Isoëtales  11, 20–1 Isoëtes  18, 20 Isoëtites  20 Isoëtopsis  606 Isoglossa  564 Isolepis  202 Isoleucas  577 Isolona  106 Isomeris  414 Isonandra  490 Isonema  527 Isophysidoideae 167 Isophysis  165–7 Isopogon  226 Isopyrum  220 Isostigma  606 Isotheca  564 Isotoma  592 Isotrema  101 Isotria  154 Isotropis  252 Itasina  633 Itaya  177 Itea  237 Itoa  330 Itzaea  534 Iva  606 Ivania  417 Ivanjohnstonia  531 Ivodea  375 ivy  371, 565, 577–8, 614, 429, 631 ivy gourd  288 Ixanthus  522 Ixchelia  325 Ixerba  360 Ixeridium  606 Ixeris  606 Ixia  167 Ixianthes  560 Ixiochlamys  606 Ixiolaena  606 Ixiolirion  164, 168 Ixodia  606 Ixodonerium  527 Ixonanthes 338 Ixophorus  209 Ixora  518 Ixyophora  155   Jablonskia  340 Jaborosa  536 Jacaranda  567–8 Jacaratia  405 jack bean  255 jackal food  101 jackalberry 492 jacket plum  373 jackfruit 274 Jackiopsis  518 Jacksonia  252 Jacmaia  606 Jacob’s ladder  485 Jacobaea  614 Jacobsenia  457 Jacquemontia  534 Jacqueshuberia  253

Jacquinia  494 Jacquiniella  155 jade plant  241 Jadunia  564 Jaegeria  606 Jaeschkea  522 Jaffrea  271 Jagera  373 Jailoloa  177 Jaimehintonia  174 jak 274 Jalcophila  606 Jaliscoa  606 Jaltomata  536, 594 jam tree  387 Jamaican gooseberry tree 341 jambolan plum  352 Jamesbrittenia  555 Jamesia  473 Jamesianthus  606 Jamesioideae 473 Jamesonia  52 jammy mouth  565 janatsi 276 Janotia  518 Jansenella  209 Janusia  318 Japanese anemone  222 Japanese fibre banana  188 Japanese knotweed  436 Japanese laurel  515 Japanese maple  374 Japanese parsley  636 Japanese pepper  278 Japanese umbrella plant 614 Jaramilloa  606 Jarandersonia  389 Jarilla  405 Jasarum  119 Jasione  592–3 jasmine  494, 528, 546–7, 568 Jasminocereus  469 Jasminum  546 Jasonia  606 Jateorhiza  218 Jatropha  335 Jaumea  606 Java apple  352 Java indigo  528 Jedda  396 Jefea  606 Jeffersonia  219 Jeffreya  606 Jeffreycia  606 jeheb nut  257 Jejewoodia  155 jellico 635 jelly palms  178 jellyfish tree  308 Jenmaniella  311 Jensenobotrya  457 Jensia  606 Jepsonia  239 Jerdonia  551 Jerusalem artichoke  611 Jerusalem sage  578 Jessea  606 jicama 256 jigal tree  257 Joannesia  335 Job’s tears  210–11 Jobinia  527 jocote 371

Jodina  427 Johanneshowellia  435 Johannesteijsmannia  177 Johnsonia  164 Johrenia  633 Johreniopsis  633 Joinvillea  240–5 jojoba oil  443 Jollydora  296 Joosia  518 Jordaaniella  457 Joseanthus  606 Josephinia  562 jostaberry 238 Jouvea  209 Jovellana  551 Jovetia  518 joyweed 452 Juania  177 Juanulloa  536 Jubaea  177 Jubaeopsis  177 Jubelina  318 Juglans  280 jujube 271 Julbernardia  251 Julostylis  390 jumby bead  258 jumby pepper  446 Jumellea  155 Jumelleanthus  390 jumping cholla  470 Juncago  127 Juncus  127, 200 juneberry 266 Junellia  573 Jungia  603 juniper 84 Juniperus  84 Juno  167 Jurausia  564 Jurinea  606 Justicia  564–5 jute 392 Juttadinteria  457   kabosu 736 Kabulia  448 Kabuyea  164–5 Kadenia  633 Kadsura  94 Kadua  518 Kaempferia  191–3 kaeng 567 kaffir lime  376 kaffir plum  371 Kafirnigania  633 Kageneckia  248, 263 kaikomako 625 Kailarsenia  518 Kailashia  633 Kairoa  111 Kairothamnus  539 Kaisupeea  551 Kajewskiella  518 kaki 492 kakoti 578 Kalaharia  577 Kalakia  633 Kalanchoë  241 Kalanchoideae 241 Kalappia  251 Kalbreyeriella  564 kale 418 Kalidium  451 Kalimeris  606, 614

Kaliphora  542 kaljeera 222 Kallstroemia  247 Kalmia  509 Kalmiopsis  509 kalonji 222 Kalopanax  630–1 kamala tree  336 Kamelinia  633 Kamettia  527 Kampochloa  209 Kanahia  527 Kanakomyrtus  352 Kanaloa  253 Kandaharia  633 Kandelia  305 Kania  352 kankong 534 Kaokochloa  209 kapok 392 kapok bush  452 Kappia  527 karakara 504 karanda nut  300 karanga 527 Karatavia  633 Karatophyllum  195 Kardomia  352 Karelinia  606 Karima  335 Karina  522 Karinia  202 Karnataka  633 karo 629 Karomia  577 Karvandarina  606 Karwinskia  271 Kaschgaria  606 Kashmiria  554 katsura 236 kauila 271 Kaunia  606 kawa pepper  99 Kawa-kawa 99 kaya 86 Kayea  310 keaki 272 Kearnemalvastrum  390 Keayodendron  341 Kebirita  252 Keckiella  554 Kedarnatha  633 Kedhalia  192 Kedrostis  287 Keenania  518 Keetia  518 Kefersteinia  155 Kegeliella  155 kei apple  330 Keiskea  577 Keithia  413 Kelleria  396 Kelleronia  247 Kellochloa  209 Kelloggia  518 Kelseya  263 keluak 324 Kelussia  633 kemiri nut  335 Kemulariella  606 kenaf 392 kencur 193 Kendrickia  355 kendyr fibre  528 keneep 373 kenguel seed  610

Plants of the World

773

INDEX Kennedia  252, 259 Kentiopsis  177 Kentrothamnus  271 Kentucky coffee tree  255 kepayang 324 Keraudrenia  389 Keraunea  534 Keraymonia  633 Kerianthera  518 Kermadecia  226 Kernera  417 kerosine bush  614 kerrawang 393 Kerria  263 Kerriochloa  209 Kerriodoxa  177 Kerriothyrsus  355 Kersting’s groundnut  255 keruing 400 kesom oil  436 ketchup 536 Keteleeria  80 keule 108–9 Kewa  455 key lime  377–8 Keysseria  606 khadi 458 Khadia  457 Khaosokia  202 Khasiaclunea  518 Khaya  381 Khmeriosicyos  287 Kibara  111 Kibatalia  527 Kickxia  554 kidney beans  254 kiekie 140 Kielmeyera  310 Kigelia  567–8 Kiggelaria  324 Kihansia  136 Killickia  577 Killipia  355 Kinabaluchloa  208 Kingdonia  216–17 kingfisher daisy  613 Kinghamia  606 Kingia  175 Kingianthus  606 Kingiodendron  251, 256 kingwood 522 Kinugasa  143 Kionophyton  159 Kipandiorchis  159 Kippistia  606 Kirengeshoma  473 Kirilowia  451 Kirkbridea  355 Kirkia  366, 382 Kissenia  476 kissing comfits  636 Kitagawia  633 Kitaibela  390 kite tree  560 kiwano 288 kiwi rose  226 kiwifruit 503–4 Kjellbergiodendron  352 Klackenbergia  522 Klaineanthus  335 Klainedoxa  303 Klaprothia  476 Klarobelia  106 Klasea  606 Klattia  165, 167 Kleinhovia  389

774

Kleinia  606, 614 Klossia  518 Klotzschia  633 Kmeria  103 knapweed 613 Knautia  623, 625 Knema  102 Knightia  226 Kniphofia  160, 168, 170 Knorringia  434 knotgrass 436 knotweed  434, 436 Knowltonia  220 Knoxia  518 Koanophyllon  606 Kobresia  202 kochia 452 Kochummenia  518 Kodalyodendron  375 Koeberlinia  407, 413 Koehneola  606 Koehneria  346 Koeleria  208 Koellensteinia  155 Koellikeria  551 Koelpinia  606 Koelreuteria  373 Koenigia  434 Koernickanthe  190 Kogelbergia  560 Kohautia  518 Kohleria  551 kohlrabi 418 kohuhu 629 Koilodepas  334 kokam 310 Kokia  390 Kokoona  294 koku-koku 322 kokuwa 504 kola nuts  391 Kolkwitzia  622 Kolobopetalum  218 Komarovia  633 komatsuma 418 konnyaku 120 Koompassia  251, 259 Koordersiochloa  208 Koordersiodendron  370 Kopsia  527 Kopsiopsis  583 korakaha 355 korarima 193 Korean aster  614 koribo 567 korintji cinnamon  112 Korshinskia  633 Korthalsella  427 Korthalsia  177 Korupodendron  350 Kosmosiphon  564 Kosopoljanskia  633 Kosteletzkya  389 Koster’s curse  355 Kostermansia  389 Kostermanthus  322 Kotschya  252 Kotschyella  417 kou 531 Koyamaea  220 koyamaki 83 Koyamasia  606 Kozlovia  633 kozo 274 Kraenzlinella  155 Krameria  246

Christenhusz, Fay & Chase

Krapfia  202 Krapovickasia  390 Krascheninnikovia  451 Krauseola  448 Kraussia  518 Kremeriella  417 Kreodanthus  159 kriek 264 Krigia  606 Krubera  633 Krugiodendron  271 kuansu 140 Kubitzkia  112 kudingcha 589 Kudoacanthus  564 kudzu 256 Kuepferia  522 Kuhlhasseltia  159 Kuhlmanniodendron  324 kulilawan 112 Kulinia  204 Kumara  220 kumbaru 255 Kumlienia  168 Kummerowia  252 kumquat 376 Kundmannia  633 Kungia  240–1 Kunhardtia  197 Kunstleria  252 Kuntheria  145–6 Kunzea  352 Kupea  136 Kuramosciadium  633 kurdee 612 kurrajong 393 Kurzamra  577 kusum tree  374 Kutchubaea  518 kuwini 371 Kydia  390 kyetpaung 527–8 Kyhosia  606 Kyllinga  220 Kyphocarpa  450 Kyrsteniopsis  606   labdanum 398 Labichea  251 Labisia  494 Lablab  252, 255, 259 Labourdonnaisia  490 Labrador tea  509 Labramia  490 Laburnum  252, 259 Lacaena  155 Lacaitaea  531 Lacandonia  89, 136 Laccodiscus  373 Laccopetalum  202 Laccospadix  177 Laccosperma  177 lacebark 396 Lachanodes  606, 614 Lachenalia  174 Lachnaea  396 Lachnagrostis  208 Lachnanthes  184 Lachnocapsa  417 Lachnocaulon  198 Lachnoloma  417 Lachnophyllum  606 Lachnorhiza  606 Lachnospermum  606 Lachnostachys  576 Lachnostylis  341

Lacistema  328 Lackeya  252 Lacmella  527 Lacomucinaea  427 lacquer  170, 256, 371, 611 lacquer ferns  56 Lactoridoideae 100–1 Lactoris  100–2 Lactuca  603, 606, 609, 611 Lactucella  603 Lacunaria  307 Ladakiella  417 ladan 398 Ladeania  252 Ladenbergia  518 lady fern  59–60 lady’s fingers  391 lady’s slipper vine  565 Ladyginia  633 lady-of-the-night 540 Laelia  155 Laennecia  606 Laestadia  606 Laetia  330 Lafoensia  346 Lafuentea  554 Lagarosiphon  125 Lagarostrobos  82 Lagascea  606 Lagedium  603 Lagenandra  119 Lagenantha  451 Lagenanthus  522 Lagenaria  287 Lagenias  522 Lagenocarpus  202 Lagenocypsela  606 Lagenophora  606 Lagerstroemia  346 Lagetta  396 Laggera  606 Lagochilus  577 Lagoecia  633 Lagophylla  606 Lagopsis  577 Lagorosolen  551 Lagoseriopsis  606 Lagotis  554 Lagrezia  450 Lagunaria  390 Laguncularia  346 Lagurus  208 Lalldhwojia  633 Lallemantia  577–8 Lamanonia  299 Lamarchea  352 Lamarckia  208 lamb’s ears  578, 610 lamb’s lettuce  623 Lambertia  226 Lamiales 543 Lamiodendron  567 Lamioideae 575–6 Lamium  577, 578 Lamourouxia  582–3 Lampadaria  551 Lampas  430 Lampayo  573 Lampranthus  457 Lamprocapnos  216 Lamprocephalus  606 Lamprolobium  252 Lampropappus  606 Lamprothamnus  518 Lamyropappus  606

Lamyropsis  606 Lanaria  162 Lancea  579 Landiopsis  518 Landolphia  527 lang du  396 Langebergia  606 Langloisia  485 langsat 381 Langsdorffia  425 Lankesterella  159 Lankesteria  564 Lannea  370 Lanonia  177 Lansium  381 Lantana  573 Lantanopsis  606 lantern lily  146 laos (spice)  193 Laosanthus  191 Lapageria  147 Lapanthus  195 Lapeirousia  167 Lapidaria  457 Lapidia  606 Lapiedra  171 Laplacea  496 Laportea  276 Lappula  531 Lapsana  606 Lapsanastrum  606 lapsi 371 Lardizabala  217 Laretia  633 large-flowered waterweed 125 Larix  80 larkspur 222 Larnax  536 Larrea  247 Larreoideae 247 Larryleachia  527 Larsenaikia  518 Larsenianthus  192 Laser  633 Laserpitium  633 Lasia  119 Lasiacis  209 Lasiadenia  396 Lasianthaea  606 Lasianthera  585 Lasianthus  518 Lasiarrhenum  531 Lasimorpha  119 Lasiobema  250 Lasiocarpus  318 Lasiocaryum  531 Lasiocephalus  606 Lasiochlamys  330 Lasiocladus  565 Lasiococca  334 Lasiocroton  334 Lasiodiscus  271 Lasiolaena  606 Lasiopetalum  389 Lasiopogon  606 Lasiosiphon  396 Lasiospermum  606 Lasiurus  209 Lastarriaea  435 Lasthenia  606, 611 Lastreopsis  66–7 Latace  171 Latania  177 Lateropora  509 Lathraea  583–4

Lathraeocarpa  518 Lathrophytum  425 Lathyrus  252, 255–6, 258–9 Latouchea  522 Latrobea  252 Latua  536 Laubertia  527 Laumoniera  380 Launaea  606 Laurales  10–11, 88, 92–3, 107 laurel  396, 515, 593 laurel cherries  267 Laurelia  109 Laureliopsis  109 Laurembergia  243 Lauridia  294 laurifolia  487 Laurophyllus  370 Laurus  112–13 Lauterbachia  111 lavandin 578 Lavandula  577–8 Lavatera  389 lavender 578 lavender cotton  611 Lavigeria  513 Lavoisiera  355 Lavrania  527 lawn 211 Lawrencella  606 Lawrencia  390 Lawsonia  346 lawyer vines  178 Laxmannia  174 Laxoplumeria  527 Layia  606, 613 leadwort 433 leafless orchid  470 Leandriella  565 Leavenworthia  417 Lebanon cedar  80 Lebeckia  252 Lebetanthus  509 lebombo ironwood  339 Lebronnecia  390 Lebrunia  310 Lebruniodendron  251 Lecananthus  518 Lecaniodiscus  373 Lecanophora  390 Lecanopteris  70 Lecanorchis  154 Lecanthus  276 Lecariocalyx  518 leche caspi  528 Lechea  398 Lechenaultia 600 Lecocarpus  606 Lecointea  252 Lecokia  633 Lecomtedoxa  490 Lecomtella  209 Lecostemon  443 Lecythis  486 Ledebouria  174 Ledebouriella  633 Ledenbergia  460 Ledermanniella  311 Ledothamnus  509 Ledum  509 Leea  244–5 leech lime  376 Leefructus  220 leek 171

INDEX Leeoideae 245 Leersia  206 Leeuwenbergia  335 Lefebvrea  633 Legazpia  558 Legenere  592 Legnephora  218 Legousia  592–3 Legrandia  352 Lehmanniella  522 Leibnitzia  603 Leidesia  334 Leiocarpaea  417 Leionema  375 Leiospora  417 Leiothrix  198 Leiothylax  311 Leiotulus  633 Leipoldtia  457 Leitneria  379–80 Lelya  518 Lembocarpus  551 Lembotropis  252 Lemmaphyllum  70 Lemna  119 Lemnoideae 118–20 lemon  376, 378 lemon balm  577 lemon basil  578 lemon thyme  578 lemon verbena  573 lemonade 378 lemongrass 211 lemon-scented oil  353 lemonwood 111 Lemooria  606 Lemurella  155 Lemurodendron  253 Lemurophoenix  177 Lemuropisum  253, 258 Lemurorchis  155 Lemurosicyos  287 Lemyrea  518 Lenbrassia  551 Lennea  252 Lennoa  531 Lenophyllum  240–1 Lens  252, 255 Lentibularia  570 lentils 254–5 Lenwebbia  352 Lenzia  464 Leocereus  469 Leochilus  155 Leocus  577 Leonardoxa  251 Leonia  325 Leonis  606 Leonotis  577–8 Leontice  219 Leontochir  144 Leontodon  606 Leontopodium  606, 612, 614 Leonurus  577–8 leopard plant  614 leopard’s bane  614 Leopoldia  173–4 Leopoldinia  177 Lepanthes  15, 152, 155 Lepanthopsis  155 Lepechinia  577–8 Lepechiniella  531 Lepeostegeres  430 Lepianthes  99 Lepidagathis  565

Lepidaploa  606 Lepidaria  430 Lepiderema  373 Lepidesmia  606 Lepidium  417 Lepidobolus  204 Lepidocaryum  177 Lepidoceras  427 Lepidocordia  531 Lepidodendrales 20 Lepidogyne  159 Lepidolopha  606 Lepidolopsis  606 Lepidomicrosorium  70 Lepidonia  606 Lepidopetalum  373 Lepidophorum  606 Lepidophyllum  606 Lepidorrhachis  177 Lepidospartum  606 Lepidosperma  202 Lepidostemon  417 Lepidostephium  606 Lepidostoma  518 Lepidothamnus  82 Lepidotrichilia  381 Lepidotrichum  417 Lepidozamia  75 Lepilaena  129 Lepinia  527 Lepiniopsis  527 Lepionurus  424 Lepironia  202 Lepisanthes  373 Lepismium  469 Lepisorus  70 Lepistemon  534 Lepistemonopsis  534 Leporella  159 Leptactina  518 Leptadenia  527 Leptagrostis  209 Leptaleum  417 Leptarrhena  239 Leptaspis  206 Leptaulus  585–6 Leptecophylla  509 Leptinella  606, 614 Leptobalanus  322 Leptoboea  551 Leptocarpha  606 Leptocarpoideae 203–4 Leptocarpus  204 Leptocarydion  209 Leptoceras  159 Leptocereus  469 Leptochilus  70 Leptochiton  171 Leptochloa  209 Leptoclinium  606 Leptocoryphium  209 Leptodermis  518 Leptoderris  252 Leptodesmia  252 Leptoglossis  536 Leptogonum  435 Leptohyptis  577 Leptolaena  399 Leptolobium  252 Leptomeria  427 Leptomischus  518 Leptonema  340 Leptonychia  389 Leptopetalum  518 Leptoplax  417 Leptopteris  29–30

Leptopus  341 Leptopyrum  220 Leptorhabdos  583 Leptorhynchos  606 Leptoscela  518 Leptosema  252 Leptosiphonium  565 Leptosolena  192 Leptospermum  352 leptosporangiate ferns  12, 22, 29–30 Leptostelma  606 Leptostigma  518 Leptoterantha  218 Leptotes  155 Leptothrium  209 Leptothyrsa  375 Leptotriche  606 Lepturidium  209 Lepturopetium  209 Lepturus  209 Lepuropetalon  294 Lepyrodia  204 Lepyrodiclis  448 Lera  355 Lerchea  518 Lereschia  633 Leretia  513 Lescaillea  606 Lesia  551 Lespedeza  252, 259 lesquerella 419 Lesqueria  103 lesser galangal  193 Lessertia  252 Lessingia  606 Lessingianthus  606 Letestua  490 Letestudoxa  106 Letestuella  311 Lethedon  396 Lethia  167 letterwood 274 Lettowia  606 Lettowianthus  106 lettuce  1, 464, 609, 611, 623 lettuce tree  461 Leucactinia  606 Leucadendron  226 Leucaena  253 Leucanthemella  606, 614 Leucanthemopsis  606 Leucanthemum  606, 614 Leucas  575, 577 Leucaster  461 Leucheria  603 Leuchtenbergia  469 Leucoblepharis  606 Leucocarpus  580 Leucocasia  119 Leucochloron  253 Leucochrysum  606 Leucocodon  518 Leucocoryne  170–1 Leucocrinum  173, 174 Leucocroton  334 Leucocyclus  606 Leucogenes  606 Leucojum  170–1 Leucolophus  518 Leucomphalos  252 Leuconotis  527 Leucophyllum  555 Leucophysalis  536 Leucophyta  606

Leucopogon  509 Leucoptera  606 Leucosalpa  583 Leucosceptrum  577 Leucosidea  263 Leucospermum  226 Leucosphaera  450 Leucospora  554 Leucostegane  251 Leucostegia  64, 68 Leucosyke  276 Leucothoë  509 Leucothrinax  177 Leunisia  603 Leutea  633 Levant storax  234 Levenhookia  595–6 Levieria  111 Levisticum  633, 635 Lewisia  464 Leycesteria  621, 623, 625 Leymus  208, 211 Leysera  606 Liabum  606 Lianthus  313 Liatris  606, 613–14 Libertia  167 Libidibia  253, 257 Libinhania  606 Libocedrus  84 Librevillea  251 Licania  322 Licaria  112 Lichtensteinia  633 Licuala  177 Lidbeckia  606 Liebigia  551 Lifago  606 Ligaria  430 Ligeophila  159 lignum rhodium  535 lignum vitae  247 Ligularia  606, 614 Ligulariopsis  606 Ligusticopsis  633 Ligusticum  633, 635 Ligustrum  546 Lijndenia  355 lilac  381, 393, 547 Lilaeopsis  633 Liliales  11, 141, 344 Liliopsida  97 Lilium  150 lily  87, 149–50 lily-of-the-valley 174 lily-of-the-valley tree  504 lilyturf 174 lima bean  254 Limacia  218 Limaciopsis  218 Limadendron  252 Limahlania  522 Limbarda  606, 609 lime caviar  376 limelo 378 limequat 376 limes  376, 392 Limeum  445, 453, 455 Limnanthes  406 Limnas  208 Limnobiophyllum  119 Limnobium  125 Limnocarpus  130 Limnocharis  122–3 Limnocitrus  375 Limnodea  208

Limnophila  554 Limnophyton  122 Limnopoa  209 Limnosciadium  633 Limnosipanea  518 Limodorum  155 limoncillo 611 Limonia  375 Limoniastrum  433 Limoniopsis  433 Limonium  433 Limosella  555 Linanthus  485 Linaria  554 Linariantha  565 Linariopsis  562 Linconia  618 Lindackeria  324 Lindbergella  208 Lindelophia  531 linden 392–3 Lindenbergia  582–4 Lindera  112 Lindernia  558 Linderniella  558 Lindheimera  606, 614 Lindleya  263 Lindmania  195 Lindsaea  45–7, 49 Lindsayomyrtus  352 linen  4, 337 Lingelsheimia  341 lingon berry  509 Linnaea  622–3, 625 Linnaeoideae 622–3 Linnaeosicyos  287 Linnaeus, Carolus  622 Linodendron  396 Linoideae 337 linoleum 337 Linospadix  177 Linostoma  396 linseed oil  337 Lintonia  209 Linum  337 Linzia  606 Lipandra  451 Liparia  252 Liparis  152, 155 Liparophyllum  598 Lipoblepharis  606 Lipocarpha  202 Lipochaeta  606 Lippia  573 lippia 573 Lipskya  633 lipstick tree  397 Liquidambar  234 liquorice 256 liquorice plant  614 Liriodendron  103 Liriope  174 Lisaea  633 Lisianthius  522 lisianthus 522 Lissanthe  509 Lissocarpa  492 Listrostachys  155 Litchi  373 Lithachne  208 Lithobium  355 Lithocarpus  278 Lithodora  531 Lithodraba  417 Lithomyrtus  352 Lithophila  450

Lithophragma  239 Lithops  457 Lithospermum  531 Lithraea  370 Litobrochia  52 Litogyne  606 Litothamnus  606 Litrisa  606 Litsea  112 Littledalea  208 Littonia  146 Litwinowia  417 liverleaf 222 liverwort 3 living stones  458 Livistona  177 Llagunoa  373 llareta 637 Llavea  51 Llerasia  606 Lloydia  150 Loasa  476 Lobanilia  334 Lobelia  592–4 Lobelioideae 591–2 Lobostemon  531 Lobularia  417 Lockhartia  155 locoto pepper  539 locust beans  256 locustberry 318 Lodoicea  177, 259 Loefgrenianthus  155 Loeflingia  448 Loeselia  485 Loesenera  251 Loeseneriella  294 Loewia  327 loganberry 266 Logania  523 Logfia  606 logwood 257 Loheria  494 Loiseleuria  509 Loliolum  208 Lolium  208, 211 lollipop plant  565 Lomagramma  66–7 Lomandra  174 Lomandroideae 173–4 Lomaria  67 Lomariopsidoideae 63, 66–7 Lomariopsis  66–7 Lomatia  226 Lomatium  633, 635 Lomatocarpa  633 Lomatogonium  522 Lomatozona  606 Lonas  606 Lonchitis  46 Lonchocarpus  252 Londonpride 239 long pepper  99 longan 273 Longetia  539 Lonicera  623, 625 lontar palm  178 loofah 289 Lopezia  349 Lophachme  209 Lophanthera  318 Lophanthus  577 Lophatherum  209 Lophiocarpus  446, 454, 459

Plants of the World

775

INDEX Lophiola  133, 162 Lophocarpinia  253 Lophogyne  311 Lopholaena  606 Lopholepis  209 Lophomyrtus  352 Lophopappus  603 Lophopetalum  294 Lophophora  469 Lophophytum  425 Lophopogon  209 Lophopterys  318 Lophopyxis  314 lophos 555 Lophosoria  39, 44–5 Lophospermum  554 Lophostemon  352 Lophostigma  373 Lophostoma  396 Lophozonia  277 Lopriorea  450 loquat 266 Loranthus  430 Lordhowea  606, 615 Lorentzianthus  606 Lorenzia  119 Loreya  355 Loricalepis  355 Loricaria  606 Loropetalum  235 Lorostemon  309 Lotononis  252, 257 Lotus  252, 257, 259 lotus  87, 223–4, 271 lotus fruit  271 Loudetia  209 Loudetiopsis  209 Louisiella  209 Lourteigia  606 Lourtella  346 Louteridium  565 lovage  635, 636 love-in-a-mist 222 love-in-a-puff 274 love-plant  468, 541 Lovoa  381 Lowia  185 Lowryanthus  606 Loxanthera  430 Loxocalyx  577 Loxocarya  204 Loxococcus  177 Loxodera  209 Loxodiscus  373 Loxogramme  69–70 Loxogrammeae  70 Loxoma 40 Loxonia  551 Loxopterygium  370 Loxoscaphe  58 Loxostigma  551 Loxostylis  370 Loxothysanus  606 Loxsoma  39–40 Loxsomatoideae 40 Lozanella  273 Lozania  328 Lubaria  375 lucerne 257 Lucilia  606 Luciliocline  606 lucky bean tree  258 Luculia  518 lúcuma 490 Lucya  518 Ludekia  518

776

Ludia  330 Ludisia  159 Ludovia  139 Ludwigia  349 Lueckelia  155 Lueddemannia  155 Luehea  389 Lueheopsis  389 Luetzelburgia  252 Luffa  287 Luina  606 Luisia  155 Lulia  603 Luma  352 Lumnitzera  346 Lunania  330 Lunaria  417 Lunasia  375 Lundellianthus  606 Lundia  567 Lundinia  606 lungwort 532 lupine bean  255 Lupinus  252, 255, 257, 259 Luronium  122 Lutzia  417 Luvunga  375 Luxemburgia  308 Luziola  206 Luzonia  252 Luzula  200 Luzuriaga  144–5, 147 Lyallia  464 Lycapsus  606 Lycaste  155 lychee 373 Lychniothyrsus  565 Lychnodiscus  373 Lychnophora  606 Lychnophoriopsis  606 Lycianthes  536 Lycium  536 Lycochloa  208 Lycomormium  155 Lycopodiales 10–11, 18–19 Lycopodiella  19 Lycopodium  18–19 lycopodium powder  19 lycopods  3, 10–11, 14, 18 Lycopus  577 Lycoris  171 Lycoseris  603 Lycurus  209 Lydenburgia  294 Lygeum  208 Lyginia  204 Lygisma  527 Lygodesmia  606 Lygodioideae 36 Lygodium  35–6 Lymania  195 lyme grass  211 Lyonia  509 Lyonothamnus  263 Lyperanthus  159 Lyperia  555 Lyrocarpa  417 Lyroglossa  159 Lysiana  430 Lysicarpus  352 Lysichiton  119 Lysidice  251 Lysiloma  253 Lysimachia  494 Lysinema  509

Christenhusz, Fay & Chase

Lysionotus  551 Lysiosepalum  389 Lysiostyles  534 Lysiphyllum  250, 257 Lysipomia  592 Lythrum  346 Lytocaryum  177   ma yuen  211 Maackia  252, 259 Maasia  106 Mabea  335 Mabrya  554 Maburea  422 maca 419 Macadamia  226 macadamia nuts  226 Macairea  355 Macaranga  334 macaranga gum  336 Macarenia  311 Macarisia  305 Macarthuria  445, 455 macaw bush  541 Macbridea  577 Macbrideina  518 mace 102 Macgregoria  295 Machaeranthera  606 Machaerina  202 Machaerium  252 Machairophyllum  457 Machaonia  518 mache 623 Mackaya  565 Mackinlaya  633 Mackinlayoideae 633 Maclaudia  527 Macleania  509 Macleaya  215–16 Maclura  274 Maclurochloa  208 Maclurodendron  375 Maclurolyra  208 Macodes  159 Macoubea  527 Macowania  606 Macphersonia  373 macqui 300 Macrachaenium  603 Macradenia  158 Macranthera  583 Macrocarpaea  522 Macrocentrum  355 Macroclinidium  603 Macroclinium  158 Macrocnemum  518 Macrococculus  218 Macrohasseltia  330 Macrolenes  355 Macrolobium  251 Macromeles  263 Macromeria  531 Macropanax  630 Macropeplus  111 Macropharynx  527 Macropidia  184 Macropiper  99 Macropodanthus  158 Macropodiella  311 Macropodina  606 Macropodium  417 Macropsychanthus  252 Macropteranthes  346 Macroptilium  252 Macrosamanea  253

Macroscepis  527 Macroselinum  633 Macrosolen  430 Macrosphyra  518 Macrostelia  389 Macrostylis  375 Macrosyringion  583 Macrothelypteris  59 Macrothumia  330 Macrotorus  111 Macrotyloma  252, 255 Macrozamia  75 Macvaughiella  606 Madagascar cloves  112 Madagascar palm  528 Madagascar periwinkle 528 Madagasikaria  318 Madagaster  606 madder 520 Madeira vine  466 Madhuca  490 Madia  606, 611, 615 Madlabium  577 madwort  419, 532 Maerua  413 Maesa  494 Maesobotrya  340 Maesoideae 494 Maesopsis  271 Maeviella  554 Maferria  311 magabeira rubber  528 Magadania  632 Magallana  403 Magdalenaea  583 magenta lilly pilly  352 Magnistipula  322 Magnolia  10–11, 94, 102–6, 114, 514 Magnoliales  10–11, 94, 102, 114, 514 Magnoliidae 72 Magnoliids  10–11, 88, 95, 213 Magnoliopsida  97 Magodendron  490 Magonia  373 Magoniella  435 Maguireanthus  355 Maguireocharis  518 Maguireothamnus  518 Magydaris  633 Maharanga  531 Mahawoa  527 mahoe 393 mahogany  380–1, 487 maholtine 392 Mahonia  219 mahua 490 Mahurea  310 mai yang  400 Maianthemophyllum  173 Maianthemum  173–4 maidenhair fern  50 Maieta  355 Maihuenia  469 Maingaya  235 Maireana  451 Mairetis  531 Mairia  606 maize  1, 116, 210, 335, 410 Majidea  373 Malabar melastome  355 Malabar spinach  465–6

Malachra  390 Malacocarpus  366 Malacocera  451 Malacothamnus  390 Malacothrix  606 Malagasia  226 Malanea  518 Malania  422 Malaxis  158 Malay apple  352 Malcolmia  417 Malephora  458 Malesherbia  327 Malesherbioideae 327 Malleastrum  381 Malleostemon  352 Mallophyton  355 Mallotus  334 mallow  388, 392–3 mallowwort 393 Malmea  106 Malmeanthus  606 Malmeoideae 106 Malope  390 Malosma  370 Malouetia  527 Malouetiella  527 Malperia  606 Malpighia  318 Malpighiales 302 Malpighiodes  318 Malpighioideae 318 Maltebrunia  206 Maltese mushroom  244 Malus  263 Malva  390 Malvales 385 Malvastrum  390 Malvaviscus  389 Malvella  390 Malvoideae 390 mamey sapote  490 Mammea  310 mammey apples  310 Mammillaria  469 Mammilloydia  469 mamoncillo 373 Manaosella  567 manbarklak 487 Mancoa  417 mandarin 376–8 Mandenovia  633 Mandevilla  527 Mandragora  536 mandrake 540–1 man-eating plant  483 Manekia  99 Manettia  518 Manfreda  173 mange-tout 254 Mangifera  370 Manglieta  103 mango 370–1 mangold 451 Mangonia  119 mangosteen 309 mangrove  304–5, 345–6, 348, 433 mangrove ferns  51 mangrove palm  178 mangrove trumpet tree 567 Manicaria  177 Manihot  335 manil 310 Manila hemp  188

Manilkara  490 Maniltoa  251 manioc 335 Manisuris  209 manjack 531 Manjekia  177 Mankyua  25 manna 256 Mannagettaea  583 Manniella  159 Manniophyton  335 Manoao  82 Manochlamys  450 Manostachya  518 Manotes  296 Mansoa  567–8 Mansonia  389 Mantalania  518 Mantisalca  606 Mantisia  191 Manulea  555 Manuleopsis  555 Manyonia  606 Maoutia  276 Mapania  202 Mapanioideae 202 maples  372, 374 Mappia  513 Mappianthus  513 Maprounea  335 Maquira  274 Maracaibo boxwood  330 maracuja 327–8 Marah  287 Marahuacaea  197 marama bean  254 Maraniona  252 Maranta  190 Maranthes  322 maranti wood  400 Marantochloa  190 Marasmodes  606 Marathrum  311 Marattia  10–11, 22, 27 Marattiales  10–11, 27–8, 43 Marattioideae 27 Marcania  565 Marcelliopsis  450 Marcetia  355 Marcgravia  481 Marcgraviastrum  481 marestail 24 Mareya  334 Mareyopsis  334 Margaranthus  536 Margaritaria  341 Margaritolobium  252 Margaritopsis  518 Margelliantha  158 Margotia  633 marguerite 614 Margyricarpus  263 marigold  610, 612–13 Marila  310 Marina  252 Maripa  534 mariposa lily  150 Markea  536 Markhamia  567–8 Marlierea  352 Marlothiella  633 marmalade  109, 266, 289, 376, 378, 381, 527 Marmaroxylon  253 marmelo 266

INDEX Marmoritis  577 Marojejya  177 Marquesia  400 marram 211 marrow 288 Marrubium  577–8 Marsdenia  527 Marshallia  606, 614 Marshalljohnstonia  606 marshmallow 392–3 Marsilea  36–7 Marsippospermum  200 Marsypianthes  577 Marsypopetalum  106 Martellidendron  140 Martensianthus  518 Marthella  134 Martianthus  577 Marticorenia  603 Martinella  567 Martiodendron  251 Martretia  340 Martynia  561 marula fruit  370 Mascagnia  318 Mascarenhasia  527 Maschalocephalus  197 Maschalodesme  518 Masdevallia  152, 158 mashua 403 mask flower  557 Masmenia  417 Masoala  177 Massonia  174 Massularia  518 Mastersia  252 Mastersiella  204 masterwort 637 mastic 370 Mastigosciadium  633 Mastigostyla  167 Mastixia  472 Mastixiodendron  518 matasano 378 Matayba  373 Matelea  527 Mathewsia  417 Mathiasella  633, 637 Mathieua  171 Mathurina  327 matilija 216 Matisia  390 Matonia  34 Matricaria  606, 611 mat-rush 174 Matsumurella  577 Matteuccia  61 Mattfeldanthus  606 Mattfeldia  606 Matthaea  111 Matthiola  417 Matudaea  235 Matudanthus  180 Matudina  606 mauca 461 Mauloutchia  102 maulsari 491 Maundia  127 Maurandella  554 Maurandya  554 Mauranthemum  606 Mauria  370 Mauritia  177 Mauritiella  177 Maurocenia  295 Mausolea  606

Maxburretia  177 Maxillaria  152, 158 Maxonia  66 Maxwellia  389 may lily  174 Mayaca  199 Mayanaea  325 Mayna  324 Mayodendron  567 maypop 328 Maytenus  295 Mazaea  518 Mazus  579–80 mboga 458 Mcneillia  448 Mcvaughia  318 meadow rue  222 meadow saffron  146 meadowsweet 266 Mecardonia  554 Mechowia  450 Mecomischus  606 Meconella  216 Meconopsis  216 Mecopus  252 Mecranium  355 mecrussé 339 Medemia  177 Medeola  150 Mediasia  633 Medicago  252, 255, 257 Medicosma  375 Medinilla  355 Mediocalcar  158 Mediusella  399 medlar 266 medronho 509 Medusagyne  307, 618 Medusagynoideae 307 Medusandra  231–2 Medusantha  577 Medusanthera  585 Meeboldia  633 Meeboldina  204 Meehan’s mint  578 Meehania  577–8 Megacarpaea  417 Megacodon  522 Megacorax  349 Megadenia  417 Megahertzia  226 Megalachne  220 Megalastrum  66 Megaleranthus  208 Megalochlamys  565 Megaloprotachne  209 Megalorchis  159 Megalostoma  565 Megaphrynium  190 Megaphyllaea  381 Megaskepasma  565 Megastachya  209 Megastigma  375 Megastylis  159 Megatritheca  389 Megistostegium  390 Megistostigma  334 Meineckia  341 Meiocarpidium  106 Meiogyne  106 Meionectes  243 Meiostemon  346 Meiracyllium  158 Meizotropis  252 Melaleuca  352 Melampodium  606

Melampyrum  583–4 Melananthus  536 Melanocenchris  209 Melanochyla  370 Melanococca  370 Melanodendron  606, 614 Melanolepis  334 Melanophylla  627 Melanopsidium  518 Melanortocarya  531 Melanosciadium  633 Melanoselinum  633 Melanospermum  557 Melanostachya  204 Melanoxylon  253, 257 Melanthera  606 Melanthioideae 143 Melanthium  143 Melasma  584 Melasphaerula  167 Melastoma  355 Melastomastrum  355 Melastomatoideae 355 melastome 355 Melhania  390 Melia  381 Melianthus  344 Melica  208, 211 Melichrus  509 Melicoccus  373 Melicope  375 Melicytus  325 Melidiscus  414 Melientha  424 Melilotus  252 Melinis  209 Melioideae 381 Meliosma  223 Melissa  577 Melittacanthus  565 Melittis  577–8 Mellera  565 Melliniella  252 Melliodendron  500, 513 Mellissia  536 Melocactus  469 Melocalamus  208 Melocanna  208 Melochia  389 Melodinus  527 Melodorum  106 Melolobium  252 melon 288 Melosperma  554 Melothria  287 Melothrianthus  287 Memecylon  355 Memoremea  531 Menais  531 Mendoncia  563–4 Mendoravia  251 Menepetalum  295 mengkulang 393 Meniocus  417 Meniscium  59 Meniscogyne  276 Menispermum  218 Menkea  417 Menonvillea  417 Mentha  575, 577–8 Mentzelia  476 Menyanthes  598–9 Menziesia  509 Meomussaenda  518 Merciera  592 Mercurialis  334

Merendera  146 Meriandra  577 Meriania  355 Merianthera  355 Mericarpaea  518 Merinthopodium  536 Merope  375 Merostachys  208 Merremia  534 Merrillanthus  527 Merrillia  375 Merrilliodendron  513 Merrilliopanax  630 Merrittia  606 Mertensia  531 Merumea  518 Merwilla  173–4 Merxmuellera  209 Meryta  630–1 Mesadenella  159 Mesadenus  159 Mesanthemum  198 Mesanthophora  606 mescal 174 Mesechites  527 Mesembryanthemum  457–8 Mesocapparis  413 Mesogramma  606 Mesogyne  274 Mesomelaena  202 Mesosetum  209 Mesosphaerum  577 mesquite 255 Mestoklema  458 Mesua  310 Metagentiana  522 Metalasia  606 Metanemone  220 Metapanax  630 Metapetrocosmea  551 Metaplexis  527 Metaporana  534 Metarungia  565 Metasequoia  84 Metashangrilaia  417 Metastachydium  577 Metaxya  39, 45 Metaxyoideae 44–5 Metcalfia  208 Meteomyrtus  352 Meterostachys  240–1 Metharme  247 Metopium  370 Metrodorea  375 Metrosideros  352 Metroxylon  177 Metteniusa  514 Metteniusales 513 Metternichia  536 Meum  633, 637 Mexacanthus  565 Mexerion  606 Mexianthus  606 Mexican devil  614 Mexican flame vine  614 Mexican heather  347 Mexican oregano  573 Mexican violet  555 Meximalva  390 Mexipedium  154 Mexotis  518 Meyenia  564 meyer lemon  378 Meyerophytum  458 Meyna  518

mezereon  394, 396 Mezia  318 Meziella  243 Mezilaurus  113 Mezoneuron  253 Mezzettia  106 mfukufuku 295 mfungu 452 Mibora  208 mibuna 418 Michauxia  592–3 michay 220 Michelia  103 Michelsonia  251 Michigan banana  106 Michrachne  209 Mickelia  66 Micklethwaitia  251 Miconia  355 Micractis  606 Micraira  209 Micrairoideae 206 Micrandra  335 Micrandropsis  335 Micrantha  417 Micranthemum  558 Micranthes  239 Micrantheum  539 Micranthocereus  469 Micranthus  167 Micrargeria  584 Micrargeriella  584 Micrasepalum  518 Micricarpaea  580 Microberlinia  251 Microbiota  84 Microcachrys  82 Microcalamus  209 Microcarpaea  558 Microcaryum  531 Microcephala  606 Microcharis  252 Microchilus  159 Microchirita  551 Microchloa  209 Microcitrus  375 Microcnemum  451 Micrococca  334 Microcodon  592 Microcoelia  158 Microcorys  576 Microcos  389 Microcybe  375 Microcycas  75 Microdesmia  322 Microdesmis  302 Microdon  557 Microdracoides  202 Microepidendrum  158 Microglossa  606 Microgramma  70 Microgyne  606 Microgynoecium  451 Microlepidium  417 Microlepis  355 Microliabum  606 Microlicia  355 Microlobius  253 Microloma  527 Micromelum  375 Micromyrtus  352 Micronychia  370 Micropera  158 Micropholis  490 Microphyes  448 Microphysa  518

Micropleura  633 Microplumeria  527 Micropora  112 Micropsis  606 Micropus  606 Micropyropsis  208 Micropyrum  208 Microrphium  522 Microsaccus  158 Microsciadium  633 Microseris  606, 611 Microsoreae  70 Microsorum  70 Microspermum  606 Microstachys  335 Microstegium  209 Microsteira  318 Microstigma  417 Microtea  446, 454, 459 Microthlaspi  417 Microtis  159 Microtoena  577–8 Microtropis  295 Microula  531 Mida  427 Middletonia  551 midnight horror  567 midwinterhoorn 621 Miersia  171 Miersiella  134 mignonette 411–12 Mikania  606, 614 Mikaniopsis  606 Mila  469 Mildbraedia  335 Mildbraediodendron  252 milfoil 609 Milium  208 Miliusa  106 milk thistles  610 milkweed 528 Milla  174 Milleria  606 millet 210 Milletia  252 Millicia  274 Milligania  160–1 Millingtonia  567–8 Millotia  606 Miltonia  158 Miltoniopsis  158 Mimetanthe  580 Mimetes  226 Mimophytum  531 Mimosa  250, 253, 259 Mimosoideae 249–50 mimosoids 250 Mimozyganthus  253 Mimulopsis  565 Mimulus  579–80 Mimusops  490 minari 636 Minaria  527 Minasia  606 miner’s lettuce  464 Minquartia  422 mintbush 578 Minthostachys  577–8 Minuartia  447–8 Minuartiella  448 Minuria  606 Mionandra  318 Miquelia  513 mirabelles 264 Mirabilis  461 miracle berry  490

Plants of the World

777

INDEX Mirandea  565 Mirbelia  252 Miricacalia  606 mirto 353 Misbrookea  606 Miscanthus  209, 211 Mischarytera  373 Mischocarpus  373 Mischodon  539 Mischogyne  106 Misodendrum  428 Misopates  554 mistletoe  278, 427 mistol fruits  271 miswak 408 Mitchella  518 Mitella  239 Mitostemma  327 Mitracarpus  518 Mitragyna  518 Mitranthes  352 Mitrantia  352 Mitraria  551 Mitrasacme  523 Mitrasacmopsis  518 Mitrastemon  510 Mitrella  106 Mitreola  523 Mitrephora  106 Mitriostigma  518 Mitrophyllum  458 mitsuba 636 mitsumata 396 Miyakea  220 Miyamayomena  606 mizuna 418 Mkilua  106 Mnesithea  209 Mniochloa  208 Mniodes  606 Mobilabium  158 mock oranges  474 mockernut hickory  280 Mocquerysia  330 Modesciadium  633 Modiola  390 Modiolastrum  390 Moehringia  448 Moenchia  448 Mogoltavia  633 Mohavea  554 Mohria  36 Moldenhawera  253 Molinadendron  235 Molinaea  373 Molinaria  163 Molinia  209, 211 Mollia  389 Mollinedia  111 Mollugo  455, 462 Molongum  527 Molopanthera  518 Molopospermum  633, 637 Moltkia  531 Moltkiopsis  531 Moluccan cabbage  461 Moluccella  577–8 mombin 371 Mommsenia  355 Momordica  287 Monachather  209 Monachosorum  49 Monactis  606 Monanthes  241 Monanthocitrus  375 Monanthotaxis  106

778

Monarda  577–8 Monardella  577–8 Monarrhenus  606 Monarthrocarpus  252 Mondia  527 Monelytrum  209 Moneses  509 money maple  374 money plant  241 Moniera  375 Monilaria  458 Monilicarpa  413 Monimia  111 Monimopetalum  295 Monizia  633 monk’s pepper  578 monkey comb  568 monkey grass  174 monkey ladder  259 monkey-apple 284 monkey-puzzle tree  81 monkshood 222 Monniera  375 Monnina  262 Monocarpia  106 Monochaetum  355 Monochasma  584 Monochilus  577 Monochoria  183 Monococcus  460 Monocostus  191 monocots  114 Monoculus  606 Monocyclanthus  106 Monocymbium  209 Monodia  209 Monodora  106 Monogereion  606 Monogramma  54 Monolena  355 Monolepis  451 Monolluma  527 Monolopia  606 Mononeuria  448 Monoon  106 Monopera  554 Monophyllaea  551 Monophyllanthe  190 Monophyllorchis  158 Monoporus  494 Monopsis  592 Monopteryx  252 Monoptilon  606 Monopyle  551 Monosalpinx  518 Monostylis  311 Monotagma  190 Monotaxis  334 Monotes  400 Monothecium  565 Monotoca  509 Monotoideae 400 Monotrema  197 Monotremoideae 197 Monotropa  509 Monotropastrum  509 Monotropoideae 509 Monotropsis  509 Monsanima  527 Monsonia  342 Monstera  119–20 Montanoa  606, 612 Monteiroa  390 Montia  464 Monticalia  606 Montigena  252

Christenhusz, Fay & Chase

Montinia  542 Montiopsis  464 Montitega  509 Montrichardia  119 Montrouziera  309 Monttea  554 moonflower  535 Moonia  606, 615 moonwort 25 moosewood 396 Moquilea  322 Moquinia  606 Moquiniastrum  606 Moquiniella  430 Mora  259 Moraea  167 Morelia  518 morello 264 Morelotia  202 Moreton Bay chestnut 255 Morettia  417 Moricandia  417 moriche palm  178 Morierina  518 Morina  621–3, 625 morina  621–2, 625 Morinda  518 Morindopsis  518 Moringa  404 Morisia  417 Morisonia  413 Morithamnus  606 Moritzia  531 Morkilioideae 247 Morkillia  247 Mormodes  158 Mormon tea  78 morning glory  535 Moronobea  309 Morrenia  527 Morsacanthus  565 Mortonia  295 Mortoniella  527 Mortoniodendron  389 Morus  274 Mosannona  106 Moscharia  603 moschatel 621 Moschopsis  601 Mosdenia  209 Mosiera  352 Mosla  577 Mosquitoxylum  370 moss phlox  487 mosses  3, 10–11, 22 Mossia  458 Mostacillastrum  417 Mostuea  523–4 Motandra  527 moth bean  255 mother-in-law’s tongue 174 mother-of-thousands 239 Motherwellia  630 motherwort 578 Motleyia  518 Moullava  253 Moultonianthus  334 mountain ash  265 mountain phlox  485 mountain sorrel  436 mountain tail-leaf  612 mountain tea  578 mountain trumpet  485 Mourera  311

Mouretia  518 Mouriri  355 mousehole tree  557 Moussonia  551 Moutabea  262 Mozaffariania  633 Msuata  606 Mtonia  606 Mucoa  527 Mucronea  435 Mucuna  252 Muehlbergella  592 Muehlenbeckia  435 Muellera  252 Muelleranthus  252 Muellerargia  287 Muellerina  430 Muellerolimon  433 mufira  296 mugwort 611 Muhlenbergia  209 Muilla  174 Muiriantha  375 Mukdenia  239 mukunawanna 452 mulberry 273–4 mulberry fig  274 mule’s ears  614 Mulgedium  603 Mulguraea  573 muli 419 Mulinum  633 mullein 557 mulli 371 Multidentia  518 Mundulea  252 mung bean  255 Munnozia  606 Munroa  209 Munronia  381 Muntingia  387 Munzothamnus  606, 614 Muraltia  262 Murbeckiella  417 Murchisonia  174 Murdannia  180 Muricaria  417 murnong 611 Murraya  375 Musa  187–8 Musanga  276 muscadine 245 Muscari  173–4 Muschleria  606 Musella  187 Musgravea  226 Musineon  633 muskmelon 288 musk-rose 267 muskwood 479 Musopsis  187 Mussaenda  518 Mussaendopsis  518 Musschia  592–3 mustard  401, 419 mustard cress  419 mustard oils  315, 401, 405, 415 , 419 Mutellina  633 Mutisia  603 Mutisioideae  602, 603 Mwasumbia  106 Myagrum  417 Myanmaria  606 Mycaranthes  220 Mycelis  603

Mycerinus  509 Mycetia  518 Myodocarpus  629–31 myoga 193 Myopordon  606 Myoporum  557 Myoschilos  427 Myosotidium  531 Myosotis  531 Myosurus  158 Myoxanthus  158 Myracrodruon  370 Myracroduon  370 Myrceugenia  352 Myrcia  352 Myrcianthes  352 Myrciaria  352 Myriactis  606 Myrialepis  177 Myrianthus  276 Myriaspora  355 Myrica  279 Myricanthe  335 Myricaria  432 Myriocarpa  276 Myriocephalus  606 Myriocladus  208 Myriolimon  433 Myrioneuron  518 Myriophyllum  243 Myriopteron  527 Myriostachya  209 Myripnois  603 Myristica  95, 102 Myrmechis  159 Myrmecodia  518 Myrmeconauclea  518 Myrmecophila  158 Myrmephytum  518 Myrocarpus  252 Myrosma  190 Myrosmodes  159 Myrospermum  252, 256 Myrothamnus  229 Myroxylon  252, 256 myrrh 367–368 Myrrhidendron  633 Myrrhinium  352 Myrrhis  633, 636 Myrsine  494 Myrsinoideae 494 Myrtales 345 Myrtama  432 Myrtastrum  352 Myrtella  352 Myrteola  352 Myrtilaria  234–5 Myrtillocactus  469 myrtle  273, 347, 350–353 myrtle berry  352 myrtle-leaved orange  377 Myrtoideae 351 Myrtopsis  375 Myrtus  352 Mysanthus  252 Mystacidium  158 Mystropetalon  425 Mystroxylon  295 Mytilarioideae 235 Myxochlamys  192 Myxopappus  606 Myxopyrum  546   Nabaluia  158 Nabalus  606 Nageia  82

Najas  123–5, 129 naked ladies  145–6 Nama  531 Namacodon  592 Namaquanthus  458 Namaquanula  171 Namataea  373 Namibia  458 Namophila  174 Nananthea  606 Nananthus  458 nanche 318 Nandina  219–20 Nandinoideae 219 Nannoglottis  606 Nannorrhops  177 nannyberry 620 Nanobubon  633 Nanochilus  192 Nanocnide  276 Nanodea  426–7 Nanodeae 427 Nanophyton  451 Nanorrhinum  554 Nanothamnus  606 Nanuza  137 Napaea  390 Napeanthus  551 Napoleonaea  486 naranjilla 539 narcissi 171 Narcissus  170–1 nardoo 37 Nardophyllum  606 Nardostachys  623 Narduroides  208 Nardus  208 Naregamia  381 Nargedia  518 Naringi  375 narras 288 Narthecium  132–3 Narvalina  606 Nasa  476 nashi pear  265 Nashia  573 Nassauvia  603 Nastanthus  601 Nasturtiopsis  417 Nasturtium  417 nasturtium 402–3 Nastus  208 Natal plum  527 Nathaliella  557 Natsiatopsis  513 Natsiatum  513 Nauclea  518 Naucleopsis  274 Naudinia  375 Naufraga  633 Nautilocalyx  551 Nautonia  527 Navarretia  485 Navia  195 Nayariophyton  390 Nealchornea  335 Neanotis  518 Neatostema  531 Neblinaea  603 Neblinantha  522 Neblinanthera  355 Neblinathamnus  518 Necastocleidoideae 602 Necepsia  334 Nechamandra  125 Necranthus  584

INDEX Nectandra  112 nectarine 264 Nectaropetalum 306 Nectaroscordum  171 Nectouxia  536 Neea  461 Needhamiella  509 needle bush  256 neem 380–1 Neeopsis  461 Neesenbeckia  202 Neesia  389 Neesiochloa  209 Negria  551 negrita extract  392 Neillia  263 Nelia  458 Nelmesia  202 Nelsia  450 Nelson’s saxifrage  239 Nelsonia  564 Nelsonianthus  606 Nelsonioideae 564 Nelumbites  223 Nelumbo  223–4 Nemacaulis  435 Nemacladoideae 592 Nemacladus  592 Nemaconia  158 Nemastylis  167 Nematanthus  551 Nematolepis  375 Nematopoa  209 Nematostylis  518 Nemesia  557 nemesias 557 Nemopanthus  589 Nemophila  531 Nemosenecio  606 Nemuaron  109 Nemum  202 Nenax  518 Nenga  177 Neoalsomitra  287 Neoapaloxylon  251 Neoastelia  161 Neobaclea  390 Neobalanocarpus  400 Neobartsia  584 Neobassia  451 Neobathiea  158 Neobeguea  381 Neobertiera  518 Neoblakea  518 Neobolusia  159 Neobouteloua  209 Neoboutonia  335 Neobracea  527 Neobrachyactis  606 Neobrittonia  390 Neobuchia  390 Neobuxbaumia  469 Neobyrnesia  375 Neocabreria  606 Neocallitropsis  84 Neocalyptrocalyx  413 Neocarya  322 Neocentema  450 Neocheiropteris  70 Neochevalierodendron  251 Neocinnamomum  112 Neoclemensia  158 Neocogniauxia  158 Neocollettia  252 Neocouma  527

Neocuatrecasia  606 Neodillenia  231 Neodistemon  276 Neodregea  146 Neodriessenia  355 Neoeplingia  577 Neofabricia  352 Neogaerrhinum  554 Neogardneria  158 Neoglaziovia  195 Neogoezia  633 Neogontscharovia  433 Neoguillauminia  335 Neogyna  158 Neoharmsia  252 Neohemsleya  490 Neohenricia  458 Neoholmgrenia  349 Neohouzeaua  208 Neohymenopogon  518 Neojeffreya  606 Neojobertia  567 Neokochia  451 Neolamarckia  518 Neolemonniera  490 Neolitsea  112 Neolloydia  469 Neololeba  208 Neoluederitzia  247 Neomarica  167 Neomezia  494 Neomicrocalamus  208 Neomillspaughia  435 Neomirandea  606 Neomitranthes  352 Neomoorea  158 Neomortonia  551 Neomuretia  633 Neomyrtus  352 Neonauclea  518 Neonelsonia  633 Neonesomia  606 Neonicholsonia  178 Neonotonia  252 Neopallasia  606 Neopanax  630 Neoparrya  633 Neopicrorhiza  554 Neopringlea  330 Neoptychocarpus  330 Neoraimondia  469 Neoraputia  375 Neorautanenia  252 Neoregelia  195 Neoregnellia  389 Neorites  226 Neoroepera  539 Neorudolphia  252 Neoschmidea  375 Neoschumannia  527 Neosciadium  630–1, 633 Neoscirpus  202 Neoscortechinia  334 Neosepicaea  567 Neoshirakia  335 Neosparton  573 Neosprucea  330 Neostapfia  209 Neostapfiella  209 Neostearia  235 Neostenanthera  106 Neotatea  310 Neotessmannia  387 Neothorelia  412 Neotina  373 Neotinea  159

Neotorularia  417 Neottia  158 Neotysonia  606 Neo-uvaria  106 Neoveitchia  178 Neowerdermannia  469 Nepal cardamom  193 Nepali hog plum  371 Nepenthes  438 Nepeta  577–8 Nepethoideae 575 Nephelaphyllum  158 Nephelium  373 Nephelochloa  208 Nephradenia  527 Nephrangis  158 Nephrodesmus  252 Nephrolepis  63, 66–7 Nephrophyllum  534, 598 Nephrosperma  178 Nephrotheca  606 Nephthytis  119 Nepsera  355 Neptunia  253, 259 Neraudia  276 néré 255 Neriacanthus  565 nerica rice  210 Nerine  171 Nerisyrenia  417 Nerium  527 Nernstia  518 Nertera  518 nerve plant  565 Nervilia  158 Nesaea  346 Nesampelos  606 Nesiota  271 Neslia  417 Nesocaryum  531 Nesocodon  592–4 Nesogenes  584 Nesogordonia  390 Nesohedyotis  518 Nesolindsaea  49 Nesoluma  490 Nesomia  606 Nesphostylis  252 nest fern  55 Nestegis  546 Nestlera  606 Nestronia  427 nettle  275–6, 278 Neuburgia  523 Neuontobotrys  417 Neuracanthus  565 Neurachne  209 Neurada  388 Neuradopsis  388 Neurocallis  52 Neurocalyx  518 Neurolaena  606 Neurolakis  606 Neuropeltis  534 Neuropeltopsis  534 Neuropoa  208 Neurotheca  522 Neurotropis  417 Neuwiedia  154 Nevada  417 Nevillea  204 Neviusia  263 New England aster  614 New South Wales Christmas bush  299 New York aster  614

New Zealand flax  170 New Zealand spinach  458 Newbouldia  567–8 Newcastelia  576 Newmania  192 Newtonia  253 Neyraudia  209 niangon 393 niaouli 353 Nicandra  536 Nichallea  518 nicker bean  258 Nicobariodendron  295 Nicolasia  606 Nicolletia  606 Nicotiana  536 nicotine 539 Nidema  158 Nidorella  606 Nidularium  195 Niedenzuella  318 Niemeyera  490 Nierembergia  536 Nietneria  133 Nigella  220, 222 niger seed  611 Nigerian golden walnut 381 night phlox  557 night-scented stock  419 nightshade 536–40 Nihon  531 Nikitinia  606 Nile trumpet  568 ninde 578 Niphaea  551 Niphogeton  633 Nipponanthemum  606, 614 Nirarathamnos  633 nispero 266 Nissolia  252 Nitraria  247, 366 Nitrophila  450 Nivellea  606 Nivenia  165, 167 Nivenioideae 167 Noaea  451 Noahdendron  235 Noccaea  417 Noccidium  417 Nodocarpaea  518 Nodonema  551 Nogra  252 Nohawilliamsia  158 Noisettia  325 Nolana  536 Nolina  173–4 Nolinoideae 173–4 Nolletia  606 Noltea  271 Nomosa  531 Nonea  531 noni 519–20 nopal 470 Norantea  481 Nordenskioldia  227 Nordenstamia  606 Norfolk Island pine  81 Norlindhia  606 Normanbya  178 Normandia  518 Normandiodendron  251 Normantha  633 Noronhia  546 Norrisia  523

Northia  490 Nostoc  230 Nostolachma  518 Notanthera  430 Notechidnopsis  527 Notelaea  546 Nothaphoebe  112 Nothapodytes  513 Notheria  158 Nothoalsomitra  287 Nothobaccaurea  340 Nothobaccharis  606 Nothobartsia  584 Nothocestrum  536 Nothochelone  554 Nothochilus  584 Nothocissus  245 Nothocnide  276 Nothofagus  277 Notholaena  53 Notholirion  150 Notholithocarpus  278 Nothopegia  370 Nothosaerva  450 Nothoschkuhria  606 Nothoscordum  171 Nothosmyrnium  633 Nothospondias  380 Nothostele  159 Nothotalisia  364 Nothotsuga  80 Nothovernonia  606 Noticastrum  606 Notiosciadium  633 Notobasis  606 Notobubon  633 Notoceras  417 Notochloe  209 Notoleptopus  341 Notopleura  518 Notopora  509 Notopterygium  633 Notoseris  606 Notothixos  427 Notothlaspi  417 Nototriche  390 Nototrichium  450 Notylia  158 Notyliopsis  158 Nouelia  603 Noveloa  311 Novenia  606 Nowickea  459 Nucularia  451 Nuphar  91 nutmeg  102, 112 Nuttallanthus  554 Nuxia  560 Nuytsia  430 Nyctaginia  461 Nyctanthes  546 Nycticalanthus  375 Nyctocalos  567 Nymania  381 Nymphaea  91, 223 Nymphaeales  10, 88–9, 116, 203 Nymphaeanae 88 Nymphoides  598–9 Nypa  140, 177 Nypoideae 177 Nyssa  472 Nyssanthes  450   oak 277–8 oats 210

obedient plant  578 Oberholzeria  252 Oberonia  158 Oberonioides  158 Obetia  276 Obliia  606 Obolaria  522 Obregonia  469 Obtegomeria  577 oca 297 Oceanopapaver  389 Ochagavia  195 Ochanostachys  422 Ochlandra  208 Ochna  308 Ochnoideae 307 Ochoterenaea  370 Ochotonophila  448 Ochradenus  412 Ochreinauclea  518 Ochrocephala  606 Ochroma  390 Ochrosia  527 Ochrosperma  352 Ochthocharis  355 Ochthocosmus 338 Ochthodium  417 Ocimum  577–8 Oclemena  606 Ocotea  112 ocotillo 482–4 Octamyrtus  352 Octarrhena  158 Octoceras  417 Octoknema  422 Octolepidoideae 396 Octolepis  396 Octolobus  390 Octomeria  158 Octopoma  458 Octotropis  518 Oddoniodendron  251 Odixia  606 Odonellia  534 Odontadenia  527 Odontanthera  527 Odontarrhena  417 Odontitella  584 Odontites  584 Odontocarya  218 Odontochilus  159 Odontocline  606 Odontophorus  458 Odontorrhynchus  159 Odontosoria  47, 49 Odontostelma  527 Odontostomum  165 Odyendea  380 Odyssea  209 Oeceoclades  158 Oecopetalum  514 Oedera  606 Oedibasis  633 Oedina  430 Oedochloa  209 Oemleria  263 Oenanthe  633, 636 Oenocarpus  178 Oenothera  349 Oeonia  158 Oeoniella  158 Oerstedina  551 Oestlundia  158 Ofaiston  451 Oftia  557 Ogastemma  531

Plants of the World

779

INDEX ogbono nut  303 ogeechee lime  472 Ohwia  252 oil nato  259 oil of guaiac  247 Oiospermum  606 Okenia  461 Okinawa spinach  610 Okoubaka  427 okra 391 Olacoideae 422 Olax  422 Oldenburgia  603 Oldenlandia  518 Oldenlandiopsis  518 oldest living plant  330 Oldfeltia  606 Oldfieldia  539 Olea  546 oleander 528 Oleandra  67–8 Oleandroideae  63, 67–8, 70 Olearia  606, 612, 614 oleaster 269 Olfersia  66 Olgaea  606 olibanum 368 Oligactis  606 Oliganthes  606 Oligarrhena  509 Oligocarpus  606 Oligoceras  335 Oligochaeta  606 Oligocladus  633 Oligocodon  518 Oligomeris  412 Oligophyton  159 Oligostachyum  208 Oligothrix  606 Olimarabidopsis  417 Olinia  358 Olisbeoideae 355 Olivaea  606 olive 545–6 olive oil  546 Oliverella  430 Oliveria  633 Oliveriana  158 Olmeca  208 Olmediella  330 Olneya  252 ololiuqui 535 Olsynium  167 olulu 593 Olyra  208 Ombrocharis  577 Ombrophytum  425 ombú 459 Omiltemia  518 Omoea  158 Omphacomeria  427 Omphalea  335 Omphalocarpum  490 Omphalodes  531 Omphalogramma  494 Omphalolappula  531 Omphalopappus  606 Omphalopus  355 Omphalotrix  584 Oncaglossum  531 Oncella  430 Oncidium  152, 158 Oncinema  527 Oncinocalyx  577 Oncinotis  527

780

Oncoba  330 Oncocalamus  177 Oncocalyx  430 Oncosiphon  606 Oncosperma  178 Oncostemum  494 Oncotheca  511 Ondetia  606 ong tsoi  534 Ongokea  422 onion  4, 170–1 Onixotis  146 Onobrychis  252, 257–8 Onoclea  55, 61 Ononis  252 Onopordum  606, 610, 612 Onoseris  603 Onosma  531 Onosmodium  531 Onuris  417 Onychium  52 Onychopetalum  106 Onychosepalum  204 Oocephala  606 Oocephalus  577 Ooia  119 Oonopsis  606 Oophytum  458 Oparanthus  608, 615 Opercularia  518 Operculicarya  370 Operculina  534 Ophiobotrys  330 Ophiocarpus  252 Ophiocaryon  223 Ophiochloa  209 Ophiocolea  567 Ophioglossales  10–11, 25 Ophioglossella  158 Ophioglossum  25 Ophionella  527 Ophiopogon  174 Ophiorhiziphyllon  564 Ophiorrhiza  518 Ophiuros  209 Ophrestia  252 Ophryosporus  608 Ophrypetalum  106 Ophrys  159 Ophthalmoblapton  335 Opilia  424 Opisthiolepis  226 Opisthocentra  355 Opisthopappus  608 Opithandra  551 opium poppy  216 Opizia  209 Oplismenopsis  209 Oplismenus  209 Oplonia  565 Oplopanax  630–1 Opocunonia  299 Opoideia  633 Opopanax  633, 636 opossum wood  618 Opsicarpium  633 Opuntia  469 orache 452 orange jessamine  540 orange mint  577–8 orange wild rhea  276 orangequat 376 oranges 376 Orania  178 Oraniopsis  177 Orbea  527

Christenhusz, Fay & Chase

Orbexilum  252 Orbivestus  608 Orchadocarpa  551 orchid shower  578 orchid tree  256 Orchidantha  185 Orchidoideae  152, 154–5, 158–9 orchids  39, 87, 151–9 Orchipedum  159 Orchis  159 Orcuttia  209 Oreacanthus  565 Oreanthes  509 Orectanthe  198 oregano  573, 577–8 Oreithales 220 Oreobambos  208 Oreobliton  451 Oreobolopsis  202 Oreobolus  202 Oreocallis  226 Oreocereus  469 Oreocharis  551 Oreochloa  208 Oreochrysum  608 Oreocnide  276 Oreocome  633 Oreocomopsis  633 Oreoleysera  608 Oreomunnea  280 Oreomyrrhis  633 Oreonana  633 Oreonesion  522 Oreopanax  630–1 Oreophysa  252 Oreophyton  417 Oreopoa  208 Oreopolus  518 Oreorchis  158 Oreoschimperella  634 Oreosolen  557 Oreosparte  527 Oreostemma  608 Oreoxis  634 Oresbia  608 Oresitrophe  239 Orestias  158 Orfilea  334 Orianthera  523 Oricia  375 Oriciopsis  375 Origanum  575, 577–8 Orinoco sassafras  113 Orinus  209 Orites  226 Oritrophium  608 Orixa  375 Orlaya  634, 637 Orleanesia  158 Ormocarpopsis  252 Ormocarpum  252 Ormopterum  634 Ormosciadium  634 Ormosia  252, 258 Ormosolenia  634 Ornduffia  598 Ornichia  522 Ornithoboea  551 Ornithocarpa  417 Ornithocephalus  158 Ornithogalum  174 Ornithoglossum  146 Ornithopus  252, 257 Ornithostaphylos  509 Orobanche  582–4

oroblanco 377 Orochaenactis  608 Orogenia  634 Orontioideae 119 Orontium  119 Oropetium  209 Orophea  106 Orophochilus  565 Orostachys  240–1 Orothamnus  226 Oroxylum  567–8 Orphanodendron  253 Orphium  522 orris root  167 ortaniques 377 Ortegia  448 Ortegocactus  469 Orthacanthus  209 Orthaea  509 Orthanthera  527 Orthilia  509 Orthion  325 Orthiopteris  47 Orthocarpus  584 Orthoceras  159 Orthoclada  209 Orthogoneuron  355 Orthogynium  218 Orthomene  218 Orthopappus  608 Orthophytum  195 Orthopichonia  527 Orthopterum  458 Orthopterygium  370 Orthopterygium  370 Orthosia  527 Orthosiphon  577–8 Orthosphenia  295 Orthrosanthus  167 Orychophragmus  417 Oryctanthus  430 Oryctes  536 Oryctina  430 Oryxis  252 Oryza  206, 210 Oryzidium  209 Oryzopsis  208 Osa  518 osage orange  274 Osbeckia  355 Osbertia  608 Osbornia  352 Oschatzia  634 Oscularia  458 Oserya  311 Osmadenia  608 Osmanthus  546 Osmelia  330 Osmiopsis  608 Osmitopsis  608 Osmolindsaea  49 Osmorhiza  634, 637 Osmoxylon  630–1 Osmunda  30 Osmundales  10–11, 29 Osmundastrum  30 oso-berry 266 Ossaea  355 Ossiculum  158 Osteomeles  264 Osteophloeum  102 Osteospermum  608, 613 Ostodes  335 Ostrearia  235 ostrich fern  55, 61 Ostrowskia  592

Ostrya  282 Ostryocarpus  252 Ostryopsis  282 Osyridicarpos  427 Osyris  427 Otacanthus  554 Otachyrium  209 Otaheite gooseberry  341 Otatea  208 Oteiza  608 Otholobium  252 Othonna  608, 614 Otion  252 Otiophora  518 Otoba  102, 609 Otocarpus  417 Otochilus  158 Otoglossum  158 Otomeria  518 Otonephelium  373 Otopappus  608 Otoptera  252 Otospermum  608 Otostegia  577 Otostylis  158 Ottelia  125 Ottleya  252 Ottoa  634 Ottochloa  209 Ottonia  99 Ottoschmidtia  518 Ottoschultzia  514 Ottosonderia  458 Oubanguia  486 Ougeinia  252 Ouratea  308 Ourisia  554 Ovidia  396 Ovieda  577 Owenia  381 Oxalidales 295 Oxalis  297 Oxandra  106 Oxanthera  375 oxe-eye 613 Oxera  577 ox-eye daisy  614 Oxford and Cambridge bush 578 Oxyanthus  518 Oxybasis  451 Oxycarpha  608 Oxyceros  518 Oxychloë  200 Oxychloris  209 Oxydendrum  509 Oxygonum  435 Oxygraphis  220 Oxygyne  134 Oxylobium  252 Oxylobus  608 Oxymyrrhine  352 Oxyosmyles  531 Oxypappus  608 Oxypetalum  527 Oxyphyllum  603 Oxypolis  634 Oxyrhachis  209 Oxyrhynchus  252 Oxyria  435 Oxyspora  355 Oxystelma  527 Oxystigma  251 Oxystophyllum  158 Oxystylis  414 Oxytenanthera  208

Oxytheca  435 Oxytropis  252 Oyedaea  608 oyster nut  289 oysterplant 532 Oziroë  173–4 Ozoroa  370 Ozothamnus  608, 614   Pabstia  158 Pabstiella  158 Pachira  390 Pachites  159 Pachyanthus  355 Pachycaulos  551 Pachycentria  355 Pachycereus  469 Pachycladon  417 Pachycormus  370 Pachycornia  451 Pachyctenium  634 Pachyelasma  253, 259 Pachygone  218 Pachylaena  603 Pachylarnax  103 Pachyloma  355 Pachymitus  417 Pachynema  231 Pachyneurum  417 Pachyphragma  417 Pachyphytum  240–1 Pachyplectron  159 Pachypleurum  634 Pachypodium  527 Pachyptera  567 Pachyrhizus  252, 255–6 Pachysandra  228 Pachystachys  565 Pachystegia  608 Pachystoma  158 Pachystroma  335 Pachystylidium  334 Pachystylus  518 Pachythamnus  608 Pacifigeron  608 Packera  608 paco-paco 392 Pacouria  527 Pacourina  608 Paederia  518 Paeonia  232–3 Paepalanthus  198 Pagamea  518 Pagameopsis  518 pagoda plant  578 Pagothyra  551 pain de sucre  610 painted fern  55 painted tongue  541 Painteria  253 Pajanelia  567–8 Pakaraimaea  398 pak-choi 418 Palaeoactaea  220 Palafoxia  608 Palaquium  490 Palaua  390 Paleaepappus  608 Paleodicraeia  311 Paleoenkianthus  507 Paliavana  551 Palicourea  518 Palimbia  634 Palisota  180 Paliurus  271 Pallenis  608

INDEX palm cabbage  180 palm heart  178 palm oil  77, 178, 335, 490 palm wine  178 Palmerella  592 Palmeria  111 palmetto 178 Palmorchis  158 palo de los brujos  540 Paloue  251 Paloveopsis  251 Pamburus  375 Pamianthe  171 Pamphilia  500 Panama hat  139 Panamanthus  430 Panax  629–30 Pancheria  299 Pancicia  634 Pancovia  373 Pancratium  171 Panda  302 pandan leaves  140 Pandanales  10–11, 115, 136, 150 Pandaniditis  140 Pandanus  140 Panderia  451 Pandiaka  450 Pandorea  567–8 Pangium  324 Panicoideae  206, 208 Panicum  209–10 Panisea  158 Pankycodon  592 Panopsis  226 Panphalea  603 panpipes 211 pansy 325 Pantacantha  536 Pantadenia  335 Panurea  252 Panzerina  577 papasan 288 Papaver  216 Papaveroideae 215–16 papaya 404–5 paper daisies  613 paper mulberry  274 paperbush 396 Paphia  509 Paphinia  158 Paphiopedilum  152, 154 Papilionanthe  158 Papillilabium  158 Pappea  373 Pappobolus  608 Pappophorum  209 paprika 538 Papuacalia  608 Papuacedrus  84 Papuaea  159 Papuanthes  430 Papuasicyos  287 Papuechites  527 Papuodendron  389 papyrus 202 Paquirea  608 Pará nut  486 Pará rubber tree  335 Parabaena  218 Parabambusa  208 Paraboea  551 Paracaleana  159 Paracalia  608 Paracalyx  252

Paracarphalea  518 Paracaryum  531 Paracautleya  191 Paracephaelis  518 Parachidendron  253 Parachim  518 Parachimarrhis  518 Paracladopus  311 Paracordyline  173 Paracorynanthe  518 Paracostus  191 Paracroton  335 Paracryphia  618 Paracryphiales 618 Paractaenium  209 Paraderris  252 Paradisanthus  158 paradise tree  380 Paradisea  174 Paradombeya  390 Paradrymonia  551 Paradrypetes  305 Parafaujasia  608 Parafestuca  208 Paragenipa  518 Parageum  263 Paragonia  566 Paragramma  70 Paraguay tea  589 paragueiro 264 Paragynoxys  608 Parahancornia  527 Paraharveya  584 Parahyparrhenia  209 Paraia  112 Parajubaea  178 Parakaempferia  192 Parakibara  111 Paraknoxia  518 Paralamium  577 Paralepistemon  534 Paralophia  158 Paramachaerium  252 Paramacrolobium  251 Paramapania  202 Paramelhania  390 Parameria  527 Paramignya  375 Paramollugo  462 Paramomum  192 Paramongaia  171 Paramyristica  102 Paranecepsia  334 Paranephelium  373 Paranephelius  608 Paraneurachne  209 Paranomus  226 Parantennaria  608 Parapachygone  218 Parapentas  518 Paraphalaenopsis  158 Paraphlomis  577 Parapholis  208 Parapiptadenia  253 Parapiqueria  608 Parapolystichum  66 Paraprenanthes  608 Paraprotium  368 Paraquilegia  220 Pararistolochia  101 Parartocarpus  274 Pararuellia  565 Parasassafras  112 Paraselinum  634 Parasenecio  608 Paraserianthes  253

Parashorea  400 Parasilaus  634 Parasitaxus  71, 82 parasol tree  391, 393 Parasopubia  584 Parasponia  273 Parastemon  322 Parastrephia  608 Parastyrax  500 Parasyncalathium  608 Paratecoma  567 Paratephrosia  252 Paratheria  209 Parathesis  494 Paravitex  576 Pardanthus  167 Pardoglossum  531 Parentucellia  584 Parepigyum  527 Pariana  208 Paridoideae 143 Parietaria  276 Parinari  322 Parinariopsis  322 Paris  143 Parishia  370 Parkia  253, 255 Parkinsonia  253, 259 Parlatoria  417 Parmentiera  567–8 Parnassia  294 Parnassioideae 294 Parochetus  252 Parodia  470 Parodianthus  573 Parodiodendron  539 Parodiodoxa  417 Parodiolyra  208 Parolinia  417 Paronychia  448 Paropsia  327 Paropsiopsis  327 Paropyrum  220 Paroxygraphis  220 Parquetina  527 Parrotia  235 Parrotiopsis  235 Parrya  417 Parryella  252 Parryodes  417 parsley 636 parsnip 634–5 Parsonsia  527 Parthenice  608 Parthenium  608, 612, 614 Parthenocissus  244–5 parvifolius  304 Pasaccardoa  603 Pascalia  608 Pasithea  168 Paspalidium  209 Paspalum  209–11 pasqueflower  222 Passerina  396 Passiflora  327 Passifloroideae  327 passionflower  328 passionfruit  326, 328 Passovia  430 Pastinaca  634 Pastinacopsis  634 Patagonula  531 patana oak  487 patchouli 578 Paterson’s curse  532 Patersonia  167

Patersonioideae 167 Patima  518 Patinoa  390 Patosia  200 Patrinia  623, 625 pattypan 288 Paubrasilia  253, 257 Pauldopia  567 Paulita  634 Paullinia  373 Paulownia  581, 583 Pauridia  163 Pauridiantha  518 Paurolepis  608 Pausandra  335 Pausinystalia  518 Pavetta  518 Pavieasia  373 Pavonia  389 pawpaw  106, 405 Paxistima  295 Payena  490 Payera  518 Paypayrola  325 Paysonia  417 peach 264 peach palm  178 peachwood 257 peanut 254 pear 264–6 Pearcea  551 pearl lupine  255 pearly everlasting  613 Pearsonia  252 pecan tree  280 Pechuel-loeschea  608 Pecluma  70 Pecteilis  159 Pectinaria  527 Pectis  608, 611 Pectocaryum  531 Pedaliodiscus  562 Pedalium  562–3 Peddiea  396 Pedersenia  450 Pedicellarum  119 Pedicularis  583–4 Pedinopetalum  634 Pediocactus  470 Pediomelum  252, 256 Pedistylis  430 peepul tree  274 Pegaeophyton  417 Peganum  247, 366 Pegia  370 Pegolettia  608 Pehria  346 Peixotoa  318 pekea nut  314 Pelagodoxa  178 Pelargonium  342 Pelatantheria  158 Pelecostemon  565 Pelecyphora  470 Pelexia  159 pelican flower  101 Peliosanthes  174 Peliostomum  557 Pellacalyx  305 Pellaea  50, 53 Pellegrinia  509 Pellegrinodendron  251 pellitory  610, 613 Peltaea  390 Peltandra  119 Peltanthera  10, 550–1

Peltantheroideae 551 Peltaria  417 Peltariopsis  417 Peltastes  527 Peltiera  252 Peltoboykinia  239 Peltogyne  251 Peltophorum  253, 257 Peltophyllum  136 Peltostigma  375 Pelucha  608 Pemphis  346 Penaea  358 pencil tree  335 Penelopeia  287 Penianthus  218 Peniocereus  470 Pennantia  625 Pennellia  418 Pennilabium  158 Pennisetum  209–11 pennyroyal 577 pennywort 631 Penstemon  554 Pentabrachion  341 Pentacalia  608 Pentace  389 Pentaceras  375 Pentachaeta  608, 612 Pentachlaena  399 Pentachondra  509 Pentaclethra  253 Pentadesma  309 Pentadiplandra  410, 413 Pentaglottis  531 Pentagonia  518 Pentalepis  608 Pentalinon  527 Pentaloncha  518 Pentameris  209 Pentanema  608 Pentanisia  518 Pentanopsis  518 Pentapetes  390 Pentaphragma  594 Pentaphylax  489 Pentaplaris  390 Pentapleura  577 Pentapogon  208 Pentaptilon  600 Pentarhopalopilia  424 Pentarrhaphis  209 Pentarrhinum  527 Pentas  518 Pentasachme  527 Pentascyphus  373, 527 Pentaspadon  370 Pentastelma  527 Pentastemona  138 Pentastemonodiscus  448 Pentatrichia  608 Pentatropis  527 Penthorum  242–3 Pentodon  518 Pentopetia  527 Pentzia  608 peony 232 Peperomia  15, 99 pepino 539 Peplidium  580 Peplonia  527 Peponidium  518 Peponium  287 Peponopsis  287 pepperflower  374 pepperleaf 96

peppermint 577 pequi nut  314 Pera  331 Peracarpa  592 Perakanthus  518 Perama  518 Peraxilla  430 Perdicium  603 Perebea  274 Peregrina  318 Pereilema  209 Perenideboles  565 perennial nasturtium  403 Pereskia  470 Pereskiopsis  470 Perezia  603 perfume tree  522 Pergularia  527 Periandra  252 Perianthomega  567 Periballia  208 Pericallis  608, 613 Pericalymma  352 Pericalypta  565 Pericampylus  218 Perichlaena  567 Pericome  608, 612 Pericopsis  252 Perideridia  634–5 Peridiscus  231–2 Perilla  577–8 Perillula  577 Periomphale  596–7 Peripentadenia  300 Peripeplus  518 Peripleura  608 Periploca  527 Periplocoideae 527 Periptera  390 Peripterygia  295 Perissocarpa  308 Perissocoeleum  634 Peristeranthus  158 Peristeria  158 Peristethium  430 Peristrophe  565 Peristylus  159 Peritassa  295 Peritoma  414 Perityle  608 periwinkle  525, 528 Perkinsiodendron  500 Pernettya  509 Peronema  577 Perotis  209 Perovskia  577–8 Perralderia  608 Perriera  380 Perrierbambus  208 Perrierodendron  399 Perrierophytum  390 Perrierosedum  241 Perrottetia  384 Perryodendron  375 Persea  112 Persian lilac  381 Persian shield  565 Persicaria  435 persimmon 491–2 Persoonia  226 Persoonioideae 226 Pertusadina  518 Pertya  603 Pertyoideae 602 Peruvian basil  578 Peruvian parsnip  635

Plants of the World

781

INDEX Peruvian pepper tree  371 Pervillaea  527 Perymenium  608 Pescatoria  158 Petagnaea  634 Petalacte  608 Petaladenium  252 Petalidium  565 Petalonyx  476 Petalostelma  527 Petalostigma  539 Petalostylis  251 Petasites  608, 610, 612, 614 Petchia  527 Petelotiella  276 Petenaea  382 Peteravenia  608 Peteria  252 Petermannia  143 Peterodendron  324 Petersianthus  486 Petitia  576, 578 Petitiocodon  518 Petitmenginia  584 Petiveria  460 Petopentia  527 Petradoria  608 Petraeomyrtus  352 Petraeovitex  577 Petrea  573 Petrobium  608, 614 Petrocallis  418 Petrocodon  551 Petrocosmea  551 Petroedmondia  634 Petromarula  592 Petronymphe  174 Petrophile  226 Petrophytum  264 Petroravenia  418 Petrorhagia  448 Petrosavia  131–2 Petrosaviales 10–11, 131–2 Petrosedum  241 Petroselinum  634, 636 Petrosimonia  451 Petteria  252 Petunia  536 petunia  536, 540–1, 565 Peucedanum  634 Peucephyllum  608 Peumus  111 peyote 470 Peyritschia  208 Pfaffia  450 Phacelia  531 Phacellanthus  584 Phacellaria  427 Phacelurus  209 Phaeanthus  106 Phaedranassa  171 Phaenanthoecium  209 Phaenocoma  608 Phaenosperma  208 Phaeoptilum  461 Phaeostigma  608 Phagnalon  608 Phainantha  355 Phaius  158 Phalacraea  608 Phalacrocarpum  608 Phalacroseris  608 Phalaenopsis  152, 158 Phalaris  208, 211

782

Phaleria  396 Phalocallis  167 Phanera  250 Phanerodiscus  422 Phaneroglossa  608, 615 Phanerophlebia  66 Phanerosorus  34 Phanerostylis  608 Phania  608 Pharnaceum  462 Pharoideae 206 Pharus  206 Phaseolus  252, 254–5 Phaulopsis  565 Phaulothamnus  449, 459 pheasant’s eye  222 Phebalium  375 Phedimus  241 Phegopteris  58–9 Pheidochloa  209 Pheidonocarpa  551 Pheladenia  159 Phelline  589, 597 Phellocalyx  518 Phellodendron  375 Phellolophium  634 Phelpsiella  197 Phelypaea  584 Phemeranthus  464 Phenakospermum  184–5 Phenax  276 Pherosphaera  82 Pherotrichis  527 Phialacanthus  565 Phialanthus  518 Phialiphora  518 Philacra  308 Philactis  608 Philadelphus  473 Philbornea  337 Philcoxia  554 Philenoptera  252 Philesia  147 Philgamia  318 Philippiella  448 Phillyrea  546 Philodendron  119–20 Philoglossa  608 Philotheca  375 Philydrella  182 Philydrum  181–2 Philyra  334 Philyrophyllum  608 Phinaea  551 Phippsia  208 Phitosia  608 Phlebocarya  184 Phlebodium  70 Phlebolobium  418 Phlebotaenia  262 Phlegmatospermum  418 Phleum  208 Phloeophila  158 Phlogacanthus  565 Phlojodicarpus  634 Phlomidoschema  577 Phlomis  577–8 Phlomoides  577 Phlox  485 Phlyctidocarpa  634 Phoebanthus  608 Phoebe  112–13 Phoenicanthus  106 Phoenicaulis  418 Phoenicophorium  178 Phoenix  177

Christenhusz, Fay & Chase

Pholidocarpus  177 Pholidostachys  178 Pholidota  158 Pholisma  531 Pholistoma  531 Pholiurus  208 Phoradendron  427 Phormium  160, 164, 168, 170 Photinia  264 Phragmanthera  430 Phragmipedium  154 Phragmites  194, 209, 211 Phragmocarpidium  390 Phragmorchis  158 Phragmotheca  390 Phravenia  418 Phreatia  158 Phryma  580 Phrynella  448 Phrynium  190 Phtheirospermum  584 Phthirusa  430 Phuopsis  518 Phuphanochloa  208 Phycella  171 Phygelius  557 Phyla  573 Phylacium  252 Phylica  271 Phyllacanthus  518 Phyllagathis  355 Phyllangium  523 Phyllanoa  325 Phyllanthoideae 341 Phyllanthopsis  341 Phyllanthus  15, 340–1 Phyllarthron  567 Phyllis  518 Phyllitis  58 Phylloboea  551 Phyllobolus  457 Phyllobotryon  330 Phyllocephalum  608 Phyllocladus  82 Phyllocrater  518 Phylloctenium  567 Phyllocyclus  522 Phyllodium  252 Phyllodoce  509 Phyllolepidium  418 Phyllomelia  518 Phyllonoma  585, 587 Phyllopentas  518 Phyllopodium  557 Phyllorachis  206 Phylloscirpus  202 Phyllosma  375 Phyllospadix  128 Phyllostachys  208, 211 Phyllostegia  577 Phyllostemonodaphne  112 Phyllostylon  272 Phyllota  252 Phyllotrichum  373 Phylloxylon  252 Phylohydrax  518 Phymaspermum  608 Phymatarum  119 Phymatidium  158 Phymatocarpus  352 Phymatosorus  70 Phymosia  390 Physacanthus  565 Physaliastrum  536 Physalis  536

Physaria  418 Physena  444 Physeterostemon  355 Physocalymma  346 Physocalyx  584 Physocardamum  418 Physocarpus  264 Physoceras  159 Physochlaina  536 Physogyne  159 Physokentia  178 Physominthe  577 Physoplexis  592–3 Physopsis  576 Physoptychis  418 Physorhynchus  418 Physospermopsis  634 Physospermum  634 Physostegia  577–8 Physostemon  414 Physostigma  252 Physotrichia  634 Phytelephas  177 Phyteuma  592–3 Phytocrene  513 Phytolacca  459 Phytolaccoideae 459 Piaranthus  527 Pibiria  327 Picardaea  518 Picconia  546 Picea  79–80 pichi 541 Pichinia  119 Pichonia  490 pickerelweed 183 Pickeringia  252 Picnomon  608 Picralima  527 Picramnia  364 Picramniales 363 Picrasma  380 Picrella  375 Picria  558 Picris  608 Picrodendron  539 Picrolemma  380 Picrorhiza  554 Picrosia  608 Picrothamnus  608 Pictetia  252 Pieris  509 Pierranthus  558 Pierreodendron  380 Pierrina  486 pietsnot 387 Pigafetta  177 pigeon bean  255 pigface 458 piggyback plant  239 pignolias 80 pignut 635 Pilbara  608 Pilea  276 Pileanthus  352 Pileostegia  473 Pilgerina  427 Pilgerodendron  84 pili nut  368 Pilidiostigma  352 Piliocalyx  352 Piliostigma  250, 257 Pillansia  167 Piloblephis  577 Pilocarpus  375 Pilocosta  355

Pilophyllum  158 Pilopleura  634 Pilosella  608, 614 Pilostyles  283 pilpilvoqui 568 Pilularia  36–7 Pimelea  396 Pimelodendron  335 Pimenta  352 Pimentelia  518 pimento 538 Pimia  389 pimpernel 494 Pimpinella  634, 636 Pinacantha  634 Pinacopodium 306 Pinales  10–11, 79–80 Pinalia  158 Pinanga  178 Pinaropappus  608 Pinarophyllon  518 Pinckneya  518 Pinda  634 pine  71–2, 79–81, 211 pineapple 194–5 pineapple guava  352 pineapple lily  174 Pineda  330 Pinellia  119 pinenuts 80 Pinga  208 Pinguicula  569 Pinillosia  608 pink-ball 393 Pinochia  527 piñones 80 pinto beans  254 Pintoa  247 Pinus  79–80 Pinzona  231 Piora  608 Piper  15, 99 Piperales  10–11, 98 pipevine 101 Pippenalia  608 Piptadenia  253 Piptadeniastrum  253 Piptadeniopsis  253 Piptanthus  252 Piptatherum  208 Piptocalyx  93 Piptocarpha  608 Piptocoma  608 Piptolepis  608 Piptophyllum  209 Piptoptera  451 Piptospatha  119 Piptostigma  106 Piptothrix  608 Pipturus  276 Piqueria  608, 614 Piqueriella  608 Piqueriopsis  608 Piranhea  539 Piresia  208 Piresiella  208 Pirinia  448 Piriqueta  327 Piscidia  252 Pisonia  461 Pisoniella  461 pisse-en-lit 610 pistachio 370 Pistacia  370 Pistia  119–20 Pistorinia  241

Pisum  252, 254 pitanga 352 Pitardella  518 Pitavia  375 Pitaviaster  375 pitaya 470 Pitcairnia  194–5 pitcher plant  439, 501 Pithecellobium  253, 255 Pithecoseris  608 Pithocarpa  608 pitomba  352, 373 Pitraea  573 Pittocaulon  608 Pittoniotis  518 Pittosporopsis  513 Pittosporum  629 Pityopsis  608 Pityopus  509 Pityranthe  389 Pityrocarpa  253 Pityrodia  576 Pityrogramma  50, 52 Placea  171 Placocarpa  518 Placodiscus  373 Placospermum  226 Pladaroxylon  608, 614 Plagiantha  209 Plagianthus  390 Plagiobasis  608 Plagiobotrys  531 Plagiocarpus  252 Plagiocheilus  608 Plagiocladus  341 Plagiogyria  39, 42 Plagiogyrioideae 41–2 Plagiolirion  171 Plagiolophus  608 Plagiopteron  295 Plagiorhegma  219 Plagioscyphus  373 Plagiosetum  209 Plagiosiphon  251 Plagiostachys  192 Plagiostyles  335 Plagius  608 Plakothira  476 Planaltina  518 Planaltoa  608 Planchonella  490 Planchonia  486 Planea  608 Planera  272 plane-tree 224 Planocarpa  509 Planodes  418 Plantago  554 plantain lily  174 plantains 188 Platanocarpus  224 Platanthera  159 Platanus  224–5 Platea  514 Plateilema  608 Plathymenia  253 Platonia  309 Platostoma  577 Platycapnos  216 Platycarpha  608 Platycarpum  518 Platycarya  280 Platycaulos  204 Platycelyphium  252 Platycerieae  70 Platycerium  70

INDEX Platychorda  204 Platycladus  84 Platycodon  592–3 Platycoryne  159 Platycrater  473 Platycyamus  252 Platydesma  375 Platygyna  334 Platylepis  159 Platylobium  252 Platylophus  299 Platymiscium  252 Platymitra  106 Platypholis  584 Platypodanthera  608 Platypodium  252 Platypterocarpus  295 Platyrhiza  158 Platysace  629, 634 Platyschkuhria  608 Platysepalum  252 Platyspermation  596–7 Platystele  158 Platystemma  551 Platystemon  216 Platytheca  300 Platythelys  159 Platytinospora  218 Platyzoma  52 Plazia  603 Plecostachys  608 Plectaneia  527 Plectocephalus  608 Plectocomia  177 Plectocomiopsis  177 Plectorrhiza  158 Plectranthus  575, 577–8 Plectrelminthus  158 Plectritis  623 Plectrocarpa  247 Plectroniella  518 Plectrophora  158 Pleea  121 Pleiacanthus  608 Pleioblastus  208 Pleiocarpa  527 Pleioceras  527 Pleiochiton  355 Pleiocoryne  518 Pleiogynium  370 Pleiokirkia  366, 380 Pleioluma  490 Pleiomeris  494 Pleione  158 Pleioneura  448 Pleiospermium  375 Pleiospilos  458 Pleiostachya  190 Pleiotaxis  603 Plenckia  295 Pleocarphus  603 Pleocnemia  66–7 Pleodendron  95 Pleogyne  218 Pleonotoma  567 Pleopeltis  70 Pleradenophora  335 Plerandra  629–31 Plethadenia  375 Plethira  355 Plettkea  448 Pleuranthodendron  330 Pleuranthodes  271 Pleuranthodium  192 Pleuricospora  509 Pleurisanthes  513

Pleurocalyptus  352 Pleurocarpaea  608 Pleurocitrus  375 Pleurocoronis  608 Pleuropappus  608 Pleuropetalum  450 Pleurophora  346 Pleurophyllum  608 Pleuropogon  208 Pleuropterantha  450 Pleurosoriopsis  70 Pleurosorus  58 Pleurospermum  632 Pleurostachys  202 Pleurostylia  295 Pleurothallis  152, 158 Pleurothallopsis  158 Pleurothyrium  112 Plicosepalus  430 Plinia  352 Plinthanthesis  209 Plinthus  458 Plocama  518 Plocaniophyllon  518 Plocoglottis  158 Plocosperma  523, 544 Ploiarium  308 Plowmania  536 Plowmanianthus  180 Pluchea  608 Plukenetia  334 plum  264–65, 299, 330, 352, 371, 373, 490, 492, 527 Plumbagella  433 Plumbaginoideae 433 Plumbago  433 plume-poppy 216 Plumeria  527 Plutarchia  509 Poa  208, 211 poached egg plant  406 Poaephyllum  158 Poales 193 Podachaenium  608 Podadenia  334 Podalyria  252 Podandrogyne  414 Podangis  158 Podanthus  608 Podistera  634 Podocaelia  355 Podocalyx  539 Podocarpus  82 Podochilus  158 Podococcus  178 Podocoma  608 Podocytisus  252 Podolasia  119 Podolepis  608, 613 Podolobium  252 Podolotus  252 Podonephelium  373 Podophorus  208 Podophylloideae 219 Podophyllum  219–20 Podopterus  435 Podorungia  565 Podosorus  70 Podostemoideae 311 Podostemum  311 Podotheca  608 Podranea  567–8 Poecilandra  308 Poecilanthe  252 Poecilolepis  608

Poeciloneuron  310 Poecilostachys  209 Poeppigia  251 Poga  284 Poggea  324 Pogogyne  577 Pogonachne  209 Pogonarthria  209 Pogonatherum  209 Pogoneura  209 Pogonia  153 Pogoniopsis  158 Pogonochloa  209 Pogonolepis  608 Pogonophora  331 Pogonopus  518 Pogonotium  177 Pogostemon  577–8 Pohlidium  209 Poikilacanthus  565 Poikilogyne  355 Poikilospermum  276 Poilanedora  413 Poilannammia  355 Poincianella  253 poinsettia 336 Poiretia  252 poison ivy  371 poison oak  371 poison sumac  371 Poissonia  252 Poitea  252 Pojarkovia  608 pokeweed 458–9 Polanisia  414 Polaskia  470 Polemannia  634 Polemanniopsis  634 Polemonium  485 Polevansia  209 Polhillia  252 Polianthes  173 Poliomintha  577–8 Poliothyrsis  330 polka dot plant  565 Pollia  180 Pollichia  448 Polpoda  462 Polyachyrus  603 Polyalthia  106 Polyanthina  608 Polyarrhena  608 Polyaster  375 Polybotrya  66 Polycalymma  608 Polycardia  295 Polycarena  557 Polycarpaea  448 Polycarpon  448 Polycephalium  513 Polyceratocarpus  106 Polychrysum  608 Polyclathra  287 Polyclita  509 Polycnemoideae 450 Polycnemum  450 Polyctenium  418 Polycycnis  158 Polydora  608 Polygala  262 Polygaloides  262 Polygonanthus  284 Polygonatum  174 Polygonella  435 Polygonoideae 435 Polygonum  435

Polylepis  263 Polylophium  634 Polylychnis  565 Polymeria  534 Polymita  458 Polymnia  608 Polyosma  615–16 Polyotidium  158 Polypleurum  311 polypod ferns  45 Polypodiales  10–11, 36 Polypodiidae  22, 29 Polypodioideae  63, 67–8, 70 Polypodium  33, 63, 70 polypody 63 Polypogon  208 Polypompholix  570 Polyporandra  513 Polypremum  548 Polypsecadium  418 Polyrhabda  450 Polyscias  629–31 Polyspatha  180 Polysphaeria  518 Polyspora  496 Polystachya  158 Polystemonanthus  251 Polystichopsis  66 Polystichum  63, 66 Polytaenia  634 Polytaenium  54 Polytaxis  608 Polytepalum  448 Polytoca  209 Polytrias  209 Polyura  518 Polyzygus  634 Pomaderris  271 Pomaria  252 Pomatocalpa  158 Pomax  518 pomegranate 346–7 pomelo 376–7 Pometia  373 pommerac 352 Pommereschea  192 Pommereulla  209 pompom tree  396 Ponapea  178 Poncirus  375 ponderosa lemon  378 Ponera  158 Ponerorchis  159 Pongamiopsis  252 Pontechium  531 Pontederia  183 Ponthieva  159 ponytail palm  174 Pooideae  193, 208 poor-man’s orchid  150 Popoviolimon  433 Popowia  106 poppy 215–16 Populina  565 Populus  330 Porana  534 Porandra  180 Poranopsis  534 Poranthera  341 Poraqueiba  514 porcelain rose  193 Porcelia  106 porcupine flower  565 Porlieria  247 Porocystis  373

Porolabium  159 Porophyllum  608 Porpax  158 Porphyroglottis  158 Porphyrostachys  159 Porphyrostemma  608 Porroglossum  158 Porrorhachis  158 Portea  195 Portenschlagiella  634 Porterandia  518 Porterella  592 Porteresia  206 portia tree  393 Portlandia  518 Portulaca  467 Portulacaria  465 Posidonia  127, 130 Poskea  554 Posoqueria  518 possumwood 618 Postiella  634 Potalia  522, 3 Potameia  112 Potamogeton  128–9 Potamophila  206 Potaninia  263 Potarophytum  197 potato  4, 210, 464, 534, 536, 541, 578 potato bean  256 potato vine  534, 541 Potentilla  15, 263, 265, 267 Poteranthera  355 Pothoidium  119 Pothomorphe  99 Pothos  119 Pottingeria  295 Pottsia  527 Pouchetia  518 Poulsenia  274 Poupartia  370 Poupartiopsis  370 Pourouma  276 Pouteria  490 Pouzolzia  276 Povedadaphne  112 Pozoa  634 Pradosia  490 Praecoxanthus  159 Prainea  274 prairie mallow  393 prairie turnip  256 Pranceacanthus  565 Prangos  634 Prantl, Karl  429 Praravinia  518 Prasium  575, 577 Prasophyllum  159 Pratia  593 Praxeliopsis  608 Praxelis  608 prayer bead vine  258 prayer plants  190 Premna  576, 578 Prenanthes  608 Prepusa  522 Prescottia  159 Preslianthus  413 Presliophytum  476 Prestelia  608 Prestoea  178 Prestonia  527 Preussiella  355 Preussiodora  518

prickly gooseberry  238 prickly lettuce  609 prickly pear  470 pride of Burma  256 primrose  349, 492, 494, 551 Primula  494 Primulina  551 Primuloideae 494 Principina  202 Pringlea  418 Pringlechloa  209 Prinsepia  264 Printzia  608 Priogymnanthus  546 Prionanthium  209 Prionium  200 Prionosciadium  634 Prionostemma  295 Prionotes  509 Prioria  251, 256 Prismatocarpus  592 Prismatomeris  518 Pristimera  295 Pritchardia  177 Priva  573 privet 547 Proboscidea  561 Prockia  330 Prockiopsis  324 Procris  276 Proiphys  171 Prolobus  608 Prolongoa  608 Promenaea  158 Prometheum  240–1 Prorograminis  193 Prosartes  146, 150 Proserpinaca  243 Prosopanche  100–1 Prosopidastrum  253 Prosopis  101, 253, 255 Prospero  173–4 Prosphytochloa  208 Prostanthera  576, 578 Prostantheroideae 575–6 Prosthechea  158 Protarum  119 Protea  226 Proteales 222 Proteoideae 226 Proteopsis  608 Protium  368 Protoananas  193, 195 Protomegabaria  340 Protorhus  370 Protoschwenckia  536 Prototulbaghia  171 Protoyucca  173 Proustia  603 Prumnopitys  82 Prunella  577–8 Prunus  264 Przewalskia  536 Psacaliopsis  608 Psacalium  608 Psammagrostis  209 Psammisia  509 Psammochloa  208 Psammogeton  634 Psammomoya  295 Psammophora  458 Psammotropha  462 Psaronius  27 Psathura  518 Psathyrostachys  208

Plants of the World

783

INDEX Psathyrotes  609 Psathyrotopsis  609 Psednotrichia  609 Psephellus  609, 613 Pseudabutilon  390 Pseudaechmea  195 Pseudagrostistachys  334 Pseudaida  518 Pseudanamomis  352 Pseudanthistiria  209 Pseudanthus  539 Pseudartabotrys  106 Pseudarthria  252 Pseudechinolaena  209 Pseudelephantopus  609 Pseudeminia  252 Pseudephedranthus  106 Pseuderanthemum  565 Pseuderia  158 Pseuderucaria  418 Pseudima  373 Pseudiris  167 Pseudoacanthocereus  470 Pseudoarabidopsis  418 Pseudobahia  609 Pseudobersama  381 Pseudobetckea  623 Pseudoblepharispermum  609 Pseudobombax  390 Pseudobotrys  586 Pseudobrickellia  609 Pseudocalyx  564 Pseudocamelina  418 Pseudocarpidium  576 Pseudocarum  634 Pseudocaryopteris  577–8 Pseudocedrela  381 Pseudocentrum  159 Pseudocherleria  448 Pseudochirita  551 Pseudoclappia  609 Pseudoclausenia  381 Pseudocodon  592 Pseudoconnarus  296 Pseudoconyza  609 Pseudocorchorus  389 Pseudocydonia  264 Pseudocymbium  390 Pseudocymopterus  634 Pseudodacryodes  368 Pseudodanthonia  208 Pseudodendron  21 Pseudodichanthium  209 Pseudodicliptera  565 Pseudodiplospora  518 Pseudodissochaeta  355 Pseudodraba  418 Pseudoeriosema  252 Pseudoernestia  355 Pseudofortuynia  418 Pseudofumaria  216 Pseudogaltonia  174 Pseudognaphalium  609 Pseudogomphrena  450 Pseudogonocalyx  509 Pseudogoodyera  159 Pseudogynoxys  609, 614 Pseudohamelia  518 Pseudohandelia  609 Pseudohydrosme  119 Pseudolachnostylis  341 Pseudolaelia  158 Pseudolarix  80 Pseudolithos  527 Pseudolmedia  274

784

Pseudolotus  252 Pseudomacrolobium  251 Pseudomalmea  106 Pseudomantalania  518 Pseudomarrubium  577 Pseudomelasma  584 Pseudomertensia  531 Pseudomiltemia  518 Pseudomonotes  400 Pseudomuscari  173–4 Pseudomussaenda  518 Pseudonemacladus  592 Pseudonesohedyotis  518 Pseudonoseris  609 Pseudopanax  630 Pseudopancovia  373 Pseudoparis  180 Pseudopentameris  209 Pseudophegopteris  59 Pseudophoenix  177 Pseudophyllanthus  341 Pseudopimpinella  634 Pseudopiptadenia  253 Pseudopiptocarpha  609 Pseudoplantago  450 Pseudoprosopis  253 Pseudoprospero  173–4 Pseudopteris  373 Pseudopyxis  518 Pseudoraphis  209 Pseudorchis  159 Pseudorhipsalis  470 Pseudoridolfia  634 Pseudorlaya  634 Pseudorontium  554 Pseudoruellia  565 Pseudosalacia  295 Pseudosamanea  253 Pseudosasa  208 Pseudosbeckia  355 Pseudoschoenus  202 Pseudosclerochloa  208 Pseudoscolopia  330 Pseudosedum  241 Pseudoselago  557 Pseudoselinum  634 Pseudosempervivum  418 Pseudosericocoma  450 Pseudosmelia  330 Pseudosmodingium  370 Pseudosopubia  584 Pseudosorghum  209 Pseudospondias  370 Pseudostachyum  208 Pseudostellaria  448 Pseudostifftia  609 Pseudostriga  584 Pseudotaxus  86 Pseudotectaria  66–7 Pseudotrachydium  634 Pseudotrillium  143 Pseudotrimezia  167 Pseudotsuga  80 Pseudoturritis  418 Pseudovanilla  154 Pseudovesicaria  418 Pseudovigna  252 Pseudoweinmannia  299 Pseudowintera  96 Pseudoxandra  106 Pseudoxytenanthera  208 Pseudoyoungia  609 Pseudoziziphus  271 Pseudozoysia  209 Pseuduvaria  106 Psiadia  609

Christenhusz, Fay & Chase

Psiadiella  609 Psidium  352 Psiguria  287 Psilactis  609 Psilanthele  565 Psilocarphus  609 Psilocaulon  457 Psilochilus  158 Psilolaemus  531 Psilolemma  209 Psilopeganum  375 Psilophyton  26 Psilostrophe  609 Psilotales 10 Psilotrichopsis  450 Psilotrichum  450 Psilotum  3, 26, 72 Psiloxyloideae 351 Psiloxylon  351 Psilurus  208 Psittacanthus  430 Psophocarpus  252, 255 Psoralea  252 Psoralidium  252 Psorothamnus  252 Psychilis  158 Psychine  418 Psychopsiella  158 Psychopsis  158 Psychopterys  318 Psychotria  15, 318, 518, 520 Psychrogeton  609 Psydrax  518 Psylliostachys  433 psyllium 554 Psyllocarpus  518 Ptaeroxylon  375 Ptelea  375 Pteleocarpa  524 Pteleopsis  346 Ptelidium  295 Pterachaenia  609 Pteralyxia  527 Pterandra  318 Pteranthus  448 Pterichis  159 Pteridella  53 Pteridium  49 Pteridoblechnum  61 Pteridocalyx  518 Pteridoideae 50–2 Pteridophyllum  215–16 Pteridrys  67 Pteris  33, 40, 50–52, 59, 66, 586 Pternopetalum  634 Pternra  355 Pterocactus  470 Pterocarpus  252, 257 Pterocarya  280 Pterocaulon  609 Pterocelastrus  295 Pteroceltis  273 Pterocephalus  623 Pteroceras  158 Pterochaeta  609 Pterocissus  245 Pterocypsela  609 Pterodiscus  562–3 Pterodon  252 Pterogastra  355 Pteroglossa  159 Pterogyne  253 Pterolepis  355 Pterolobium  253

Pteronia  609 Pteropepon  287 Pteropyrum  435 Pterorhachis  381 Pterospermum  390 Pterospora  509 Pterostegia  435 Pterostemma  158 Pterostemon  237 Pterostylis  159 Pterostyrax  500 Pterozonium  52 Pterygiella  584 Pterygocalyx  522 Pterygodium  159 Pterygopappus  609 Pterygopleurum  634 Pterygostemon  418 Pterygota  390 Pteryxia  634 Ptilimnium  634 Ptilochaeta  318 Ptilostemon  609 Ptilothrix  202 Ptilotus  450 Ptisana  27 Ptychococcus  178 Ptycholobium  252 Ptychopetalum  422 Ptychopyxis  334 Ptychosema  252 Ptychosperma  178 Ptychotis  634 Ptyssiglottis  565 Pubistylus  518 Puccinellia  208 Puccionia  414 puccoon 532 puchurin nut  112 Puelia  206 Puelioideae 206 Pueraria  252, 256 Puffia  518 Pugionium  418 pulaka 120 pulasan 373 Pulchranthus  565 Pulicaria  609 Pulicarioidea  609 Pullea  299 Pulmonaria  531 Pulsatilla  220, 222, 588 Pultenaea  252 pumpkin 288 pumpkin seed  289 punah 481–2 Punica  346 Pupalia  450 purchury bean  112 Purdiaea  505 Purdieanthus  522 purging nut  335 purple appleberry  629 purple bell-vine  555 purple cone flower  611 purple loosestrife  347 purple nutsedge  202 purple velvet plant  614 purple wreath  573 purpletop vervain  573 Purpureostemon  352 Purshia  263 purslane  448, 464, 467 purum hijos  481 Puschkinia  174 pussytail 452

putaweta 589 Putranjiva  315 Puya  195–6 Pycnandra  490 Pycnanthemum  577 Pycnanthus  102 Pycnarrhena  218 Pycnobotrya  527 Pycnocoma  334 Pycnocycla  634 Pycnophyllopsis  448 Pycnophyllum  448 Pycnoplinthopsis  418 Pycnoplinthus  418 Pycnorhachis  527 Pycnosorus  609 Pycnospatha  119 Pycnosphaera  522 Pycnospora  252 Pycnostachys  577–8 Pycreus  202 Pygmaeorchis  158 Pygmaeothamnus  518 Pyracantha  264 Pyragra  518 Pyramidanthe  106 Pyramidoptera  634 Pyranthus  252 Pyrenacantha  513 Pyrenaria  496 Pyrenean saxifrage  239 Pyrgophyllum  191 Pyrola  509 Pyrolirion  171 Pyroloideae 509 Pyrorchis  159 Pyrostegia  567–8 Pyrostria  518 Pyrrhanthera  209 Pyrrhopappus  609 Pyrrocoma  609 Pyrrorhiza  184 Pyrrosia  63, 70, 257 Pyrularia  427 Pyrus  264 Pytinocarpa  609 Pyxidanthera  499   qantu 485 qat 295 Quadribractea  609 quaking aspen  330 Qualea  350 quamash 174 quandong 527 Quaqua  527 Quararibea  390 quaruba 349–50 Quassia  380 Quaternella  450 Quechua  159 Quechualia  609 Queen of the Andes  196 queen palm  178 Quekettia  158 Quelchia  603 Quercus  278, 589 Quesnelia  195 Quetzalia  295 Quezeliantha  418 Quiabentia  470 Quiina  307 Quiinoideae  307 quilineja 145 Quillaja  248 quince 266

Quinchamalium  428 Quincula  536 Quinetia  609 quinghao 611 quinine 519 quinoa 451 Quinqueremulus  609 Quintinia  618 Quipuanthus  355 Quisqueya  158 Quivisianthe  381   raapstelen 419 Rabiea  458 Racemobambos  208 racemosum; 551 Rachelia  609 Rachicallis  518 Racinaea  195 Radamaea  584 Radcliffea  335 Raddia  208 Raddiella  208 Radermachera  567–8 radhuni 636 radicchio 610 Radinosiphon  167 Radiola  337 Radiovittaria  54 radish 418 Radlkofera  373 Radlkoferotoma  609 Radyera  390 Raffenaldia  418 raffia  179 Rafflesia  245, 332 Rafinesquia  609 Rafnia  252 ragweed 614 ragwort 614 Raiguenrayun  602 Raillardella  609 rain tree  255 rainforest plum  352 Rainiera  609 raishan 210 raisin bush  371 raisin tree  271 raisins  245, 620 Rajania  134 rambai 341 ramie 276 ramin 396 Ramirezella  252 Ramisia  461 Ramonda  551 ramontchi 330 Ramorinoa  252 Ramosmania  518 rampion 592 ramtil 611 Ranalisma  122 Randia  518 Randonia  412 Rangaeris  158 Rangoon creeper  346 Ranopisoa  557 ransamala 234 Ranunculales 214 Ranunculoideae 220 Ranunculus  220, 222 Ranzania  219 Raoulia  609, 614 Raouliopsis  609 Raparia  418 Rapatea  197

INDEX Rapateoideae 197 rapeseed 419 Raphanorhyncha  418 Raphanus  418 Raphia  177 Raphidiocystis  287 Raphiocarpus  551 Raphionacme  527 Raphistemma  527 Rapistrum  418 Rapona  534 Raputia  375 Raputiarana  375 Raritebe  518 raspberry 266 Rastrophyllum  609 ratany 246 rathkihiriya 193 Ratibida  609, 614 rattan 178 rattan vine  271 rattlesnake master  637 Ratzeburgia  209 Rauhia  171 Rauhiella  158 Rauia  375 Raukaua  630 Raulinoa  375 Raulinoreitzia  609 Rauvolfia  527 Rauvolfioideae  527 Ravenala  184–5 Ravenea  177 Ravenia  375 Raveniopsis  375 Rawsonia  324 Rayjacksonia  609 Raylaya  389 Razafimandimbisonia  518 Razisea  565 Reaumuria  432 Rebutia  470 Recchia  260 Recordia  573 Recordoxylon  253 red bark  519 red bead tree  258 red boppel nut  226 red cedar  381 red rose of Lancaster  267 red snakeweed  573 red toon  381 red valerian  625 redbud 256 redbush 256 redcurrant 238 Redfieldia  209 redoul 286 redstar 535 Reedia  202 Reevesia  389 Regelia  352 Registaniella  634 Regnellidium  36–7 Rehdera  573 Rehderodendron  500 Rehia  208 Rehmannia  582–4 Reichardia  609–10 Reichenbachia  461 Reimarochloa  209 Reineckea  174 Reinhardtia  178 Reinwardtia  337 Reinwardtiodendron  381 Reissantia  295

Reissekia  271 Reitzia  208 Relchela  208 Reldia  551 Relhania  609 Remijia  518 Remirema  534 Remusatia  119 Remya  609 Renanthera  158 Renealmia  191–3 Rennellia  518 Rennera  609 Renschia  577 Rensonia  609 Requienia  252 Reseda  412 Resetnikia  418 Resia  551 Resnova  173–4 Restella  396 Restio  204 Restionoideae 203–4 Restrepia  158 Restrepiella  158 Retama  252 retamo wax  247 Retanilla  271 Retiniphyllum  518 Retispatha  177 Retrophyllum  71 Retzia  523, 560 Revealia  609 Reyemia  557 Reyesia  536 Reynaudia  209 Reynosia  271 Rhabdadenia  527 Rhabdocaulon  577 Rhabdodendron  443 Rhabdophyllum  308 Rhabdosciadium  634 Rhabdothamnopsis  551 Rhabdothamnus  551 Rhabdotosperma  557 Rhachidosoroideae 55–6 Rhachidosorus  56 Rhadinopus  518 Rhadinothamnus  375 Rhaesteria  158 Rhagadiolus  609 Rhagodia  451 Rhammatophyllum  418 Rhamnella  271 Rhamnidium  271 Rhamnoneuron  396 Rhamnus  271 Rhamphicarpa  584 Rhamphogyne  609 Rhamphorhynchus  159 Rhanteriopsis  609 Rhanterium  609 Rhaphidophora  119 Rhaphidura  518 Rhaphiodon  577 Rhaphiolepis  264 Rhaphiostylis  514 Rhaphispermum  584 Rhaphithamnus  573 Rhapidophyllum  177 Rhapis  177 Rhaponticoides  609 Rhaponticum  609 Rhaptonema  218 Rhaptopetalum  486 Rheochloa  209

Rhetinocarpha  609 Rhetinolepis  609 Rheum  435 Rhexia  355 Rhigiocarya  218 Rhigiophyllum  592 Rhigospira  527 Rhigozum  567 Rhinacanthus  565 Rhinactinidia  609 Rhinanthus  583–4 Rhinephyllum  458 Rhinerrhiza  158 Rhinerrhizopsis  158 Rhinotropis  262 Rhipidantha  518 Rhipidocladum  208 Rhipidoglossum  158 Rhipsalis  470 Rhizanthella  151, 159 Rhizanthes  332 Rhizobotrya  418 Rhizocephalus  208 Rhizomatophora  634 Rhizophora  305 Rhodalsine  448 Rhodamnia  352 Rhodanthe  609, 613 Rhodanthemum  609 Rhodiola  241 rhodium wood  535 Rhodocactus  470 Rhodochiton  554 Rhodocolea  567 Rhodocoma  204 Rhododendron  159, 509 Rhododon  577 Rhodogeron  609 Rhodohypoxis  163 Rhodolaena  399 Rhodoleia  234–5 Rhodomyrtus  352 Rhodopentas  518 Rhodophiala  171 Rhodopis  252 Rhodoscirpus  202 Rhodospatha  119 Rhodosphaera  370 Rhodostemonodaphne  112 Rhodothamnus  509 Rhodotypos  264 Rhoiacarpus  427 Rhoicissus  244–5 Rhoiptelea  280 Rhombochlamys  565 Rhomboda  159 Rhombolytrum  208 Rhombophyllum  458 Rhoogeton  551 Rhopalephora  180 Rhopaloblaste  178 Rhopalocarpus  394 Rhopalocnemis  425 Rhopalopilia  424 Rhopalostylis  178 rhubarb  229, 239, 435, 635 Rhus  370 Rhynchanthera  355 Rhynchanthus  192 Rhynchocalyx  358 Rhynchocladium  202 Rhynchocorys  584 Rhynchoglossum  551 Rhynchogyna  158 Rhyncholacis  311

Rhyncholaelia  158 Rhynchophora  318 Rhynchopsidium  609 Rhynchoryza  208 Rhynchosia  252, 258 Rhynchosida  390 Rhynchospermum  609 Rhynchospora  202 Rhynchostele  158 Rhynchostylis  158 Rhynchotechum  551 Rhynchotheca  344 Rhynchotropis  252 Rhynia  26 Rhyniophytes  72 Rhysolepis  609 Rhysopterus  634 Rhysotoechia  373 Rhyssolobium  527 Rhyssostelma  527 Rhytachne  209 Rhyticaryum  513 Rhytidanthera  308 Rhytidiphyllum  551 Rhytidocaulon  527 ribbonwood 393 Ribena 238 Ribes  238 rice bean  255 rice paper plant  630 riceflower  396 Richardia  518 Richardsiella  209 Richea  509 Richeria  340 Richterago  603 Richteria  609 Richtersveldia  527 Ricinocarpos  335 Ricinodendron  335 Ricinus  334 Ricotia  418 Ridleyandra  551 Ridleyella  158 Ridolfia  634 Riedelia  192 Riedeliella  252 Riencourtia  609 Rigiopappus  609 Rimacola  159 Rindera  531 Rinorea  325 Rinoreocarpus  325 Rinzia  352 Rio Grande cherry  352 Riocreuxia  527 Riodocea  518 Ripogonum  148 Riqueuria  518 Risleya  158 Ristantia  352 Ritchiea  413 Ritonia  565 Rivasmartinezia  634 Rivea  534 Rivina  460 Robbairea  448 Robbrechtia  518 Robeschia  418 Robinia  252, 258–9 Robinsonecio  609 Robinsonella  390 Robinsonia  614 Robiquetia  158 roble  276–77, 568 Robsonodendron  295

robusta coffee  519 Robynsia  518 Robynsiophyton  252 Rochefortia  531 Rochelia  531 Rochonia  609 rock jasmine  494 rocket  412, 418–19 Rockinghamia  334 rockmelon 288 rockroot 459 Rodgersia  239 Rodriguezia  158 Roebuckiella  609 Roella  592 Roemeria  216 Roeperocharis  159 Rogeria  562 Rogersonanthus  522 Rogiera  518 Rohdea  174 Roigella  518 Rojasianthe  609 Rojasimalva  390 Rolandra  609 Roldana  609 Rollinia  106 Roman chamomile  611 Roman shields  419 Romanoa  334 Romanschulzia  418 Romanzoffia  531 Romeroa  567 Romnalda  174 Romneya  215–16 Romulea  165, 167 Ronabea  518 Rondeletia  518 Rondonanthus  198 Ronnbergia  195 Roodebergia  609 rooibos 256 root chicory  610 Roraimaea  522 Roridula  502 Rorippa  418 Rosa  263 rosa jamaica  391 Rosales 262 rosary bean  258 Roscheria  178 Roscoea  192 rose  262, 267 rose hips  267 rose of Jericho  21 rose of Sharon  393 rose pepper  371 roseapple 352 roselle 391 rosemary  577–78, 636 Rosenbergiodendron  518 Rosenia  609 Roseodendron  567, 568 roseroot 241 rosewood 258 Rosifax  450 Rosmarinus  636 Rosselia  368 Rossioglossum  158 Rostkovia  200 Rostraria  208 Rosularia  240 –1 rosy dipelta  625 Rotala  346 Rotheca  577–8 Rothia  252

Rothmaleria  609 Rothmannia  518 Rottboellia  209 Roucheria  337 rough lemons  378 Roupala  226 Rourea  296 Roussea  589–91 Rousseauxia  355 Rousselia  276 Rouya  634 rowan 265 royal palms  179 Roycea  451 Royena  492 Roylea  577 Roystonea  178 Ruagea  381 rubber fig  274 rubber vine  528 Rubia  518 Rubiteucris  577 Rubovietnamia  518 Rubus  263 Rudbeckia  609, 613–14, 622 Rudgea  518 Rudolfiella  158 Rudolf-kamelinia  418 Ruellia  563, 565 Ruelliopsis  565 Rufodorsia  551 Rugelia  609 Rugoloa  209 Ruizia  390 Ruizodendron  106 Ruizterania  350 Rulingia  389 rum  152, 210–11 Rumex  435 Rumfordia  609 Rumia  634 Rumohra  63, 66 Rungia  565 Rupertia  252 Rupestrea  355 Rupicapnos  216 Rupicola  509 Ruppia  127, 129, 130–1 Ruprechtia  435 Rusbya  509 Ruschia  458 Ruschianthus  458 Ruscus  173–4 rush  24–5, 123, 193, 200, 205 Ruspolia  565 Russelia  554 Russian olive  269 Russian sage  578 Russian tarragon  611 Russowia  609 Rustia  518 Ruta  375 rutabaga 418 Rutaneblina  375 Ruthalicia  287 Rutheopsis  634 Ruthiella  592 Rutidea  518 Rutidosis  609 Rutoideae 375 Ruttya  565 Ruyschia  481 Ryania  330 Rydingia  577

Plants of the World

785

INDEX rye 210 Ryparosa  324 Rytidocarpus  418 Rytidosperma  209 Rytigynia  519 Rzedowskia  295   sa’ada 388 Saba  527 Sabal  13, 177–9 Sabatia  522 Sabazia  609 Sabia  223 Sabicea  519 Sabinaria  177 Saccellium  531 Saccharum  209–10 Saccifolium  522 Sacciolepis  209 Saccocalyx  577 Saccolabiopsis  158 Saccolabium  158 Saccoloma  46–7 sacha inchi  335 Sachsia  609 Sacleuxia  527 Sacoglottis  323 Sacoila  159 Sacosperma  519 sacred bamboo  220 sacred fig  274 sacred lotus  87, 223–4 Sadiria  494 Sadleria  61 safflower  612 saffron 167 safu nuts  368 sage 577–8 Sageraea  106 Sageretia  271 Sagina  448 Sagittaria  122 sago  73–4, 178 sago palm  178 Sagotia  335 saguaro 470 Saharanthus  433 sainfoin 257 saintpaulias 551 Saintpauliopsis  564 Sairocarpus  554 Sajanella  634 sakaki 489 Sakoanala  252 Salacca  177 Salacia  295 Salacighia  295 Salaciopsis  295 salad greens  239, 241, 392, 418, 609–10, 636 salad rocket  418 Salcedoa  603 Saldinia  519 salep 152 Salicornia  451 Salix  216, 330 Salmea  609 salmonberry 266 Salomonia  262 Salpianthus  461 Salpichlaena  61, 179 Salpichroa  536 Salpiglossis  536 Salpinga  355 Salpixantha  565

786

salsify 611–12 Salsola  451 Salta  435 saltcedar 432 Saltera  358 Saltia  450 Salvadora  408 Salvadoropsis  295 Salvertia  350 Salvia  575, 577–8 Salvinia  38 Salviniales  10–11, 36–8 Salvinioideae 38 Salweenia  252 Salzmannia  519 Samadera  380 Samaipaticereus  470 Samanea  253, 255, 259 sambong 612 Sambucus  619–22 Samolus  494 Sampantaea  334 Samuelssonia  565 Samyda  330 Sanango  551 Sanangoöideae 551 Sanblasia  190 Sanchezia  565 Sanctambrosia  448 sand apple  492 sand food  531 sandalwood  378, 426–7 Sandbergia  418 sandbox tree  336 Sanderella  158 Sandersonia  146 Sandoricum  381 Sandwithia  335 sangre de drago  257 Sanguinaria  215–16 Sanguisorba  263 sanicle 637 Sanicula  634, 637 Saniculiophyllum  239 Saniculoideae 634 Sanjappa  253 Sankowskya  539 Sannantha  352 Sanrafaelia  106 Sansevieria  173–4 Santalales 421 Santaleae 427 Santalum  427 Santiria  368 Santisukia  567 santol 381 Santolina  609, 611 Santomasia  313 Santosia  609 Santotomasia  158 Sanvitalia  609, 613 sao 371 sapele 381 Saphesia  458 Sapindales 364 Sapindoideae 373 Sapindus  373 Sapium  335 sapodilla 489–90 Saponaria  448 Saposhnikovia  634 Sapotoideae 490 sappanwood 257 Sapphoa  565 Sapranthus  106 Sapria  332

Christenhusz, Fay & Chase

Saprosma  519 sapucaia nut  486 Saraca  251, 259 Saracha  536 Saranthe  190 Sararanga  140 Sarawakodendron  295 Sarcandra  114 Sarcanthemum  609 Sarcanthopsis  158 Sarcaulus  490 Sarcocapnos  216 Sarcocephalus  519 Sarcochilus  158 Sarcochlamys  276 Sarcococca  228 Sarcocornia  451 Sarcodes  509 Sarcodraba  418 Sarcodum  252 Sarcoglottis  159 Sarcoglyphis  158 Sarcolaena  399 Sarcolobus  527 Sarcolophium  218 Sarcomelicope  375 Sarcomphalus  271 Sarcopera  481 Sarcopetalum  218 Sarcophrynium  190 Sarcophyte  425 Sarcophyton  158 Sarcopilea  276 Sarcopoterium  263 Sarcopteryx  373 Sarcopyramis  355 Sarcorhachis  99 Sarcorrhiza  527 Sarcosperma  490 Sarcospermoideae 490 Sarcostemmma  527 Sarcostigma  513 Sarcostoma  158 Sarcotheca  297 Sarcotoechia  373 Sarcotoxicum  413 Sargentodoxa  217 Saribus  177 Sarmienta  551 Sarocalamus  208 Sarracenia  501 sarsaparilla 149 sarson 418 Sartidia  209 Sartoria  252 Sartorina  609 Sartwellia  609 Saruma  101 Sasa  208, 211 Sasaella  208 saskatoon 266 Sassafras  109, 112–3 sassafras  109, 112–13 Satakentia  178 Satanocrater  565 sataw beans  255 satiné 274 Satranala  177 satsumas 378 Satureja  577–8 Satyria  509 Satyrium  152, 159 Saugetia  209 Saundersia  158 Saurauia  504 Sauroglossum  159

Sauromatum  119–20 Saururus  98 sausage tree  568 Saussurea  609 Sautiera  565 Sauvagesia  308 Sauvallea  180 Savannosiphon  167 Savia  341 Savignya  418 savory 577–8 saw palmetto  178 sawarri nut  314 saw-wort 612 Saxicolella  311 Saxifraga  239 Saxifragales 231 saxifrage 238–9 Saxifragodes  239 Saxifragopsis  239 Saxofridericia  197 Saxofriedericioideae 197 Scabiosa  623, 625 Scadoxus  171 Scaevola  599–600 Scagea  539 Scalesia  609, 615 Scaligeria  632 scallion 171 scallop 288 Scambopus  418 Scandianthus  533 Scandix  634 Scaphiarabis  418 Scaphiophora  82, 134 Scaphispatha  119 Scaphium  390 Scaphocalyx  324 Scaphochlamys  192 Scaphopetalum  389 Scaphosepalum  158 Scaphyglottis  158 Scaraboides  634 Scariola  603 scarlet runner bean  254 scarlet trumpet  485 Schaefferia  295 Schaffnerella  209 Schaueria  565 Schaueriopsis  565 Schedonnardus  209 Schefferomitra  106 Schefflera  629–31 Schefflerodendron  252 Schelhammera  83 Schenkia  522 Scherya  609 Schickendantzia  144 Schickendantziella  144, 171 Schiedea  448 Schiedeella  159 Schiekia  184 Schima  496 Schimpera  418 Schindlera  460 Schinopsis  370 Schinus  370 Schinziella  522 Schippia  177 Schisandra  92–4 Schischkinia  609 Schismatoclada  519 Schismatoglottis  119 Schismocarpus  476 Schismus  209

Schistocarpaea  271 Schistocarpha  609 Schistophragma  554 Schistostemon  323 Schistostephium  609 Schistotylus  158 Schizachne  208 Schizachyrium  209 Schizacme  523 Schizaea  36 Schizaeales 10–11 Schizaeaoideae 36 Schizanthus  536 Schizeilema  634 Schizenterospermum  519 Schizocalyx  519 Schizocarphus  173–4 Schizocarpum  287 Schizochilus  159 Schizocolea  519 Schizogyne  609 Schizolaena  399 Schizolobium  253 Schizomeria  299 Schizomussaenda  519 Schizonepeta  577 Schizopepon  287 Schizopetalon  418 Schizophragma  473 Schizopsera  609 Schizosepala  554 Schizostachyum  208 Schizostylis  167 Schizotorenia  558 Schizotrichia  609 Schizozygia  527 Schkuhria  609 Schlechtendalia  603 Schlechteranthus  458 Schlechterella  527 Schlechterina  327 Schlegelia  570–1 Schleichera  373 Schleinitzia  253 Schlimia  158 Schlumbergera  470 Schmalhausenia  609 Schmardaea  381 Schmidtia  209 Schmidtottia  519 Schnabelia  577 Schnella  250 Schoenefeldia  209 Schoenia  609 Schoenobiblus  396 Schoenocaulon  143 Schoenocephalium  197 Schoenolaena  634 Schoenolirion  174 Schoenoplectiella  202 Schoenoplectus  202 Schoenorchis  158 Schoenoselinum  634 Schoenoxiphium  202 Schoenus  202 Schoepfia  428, 509 Scholleropsis  183 Scholtzia  352 Schotia  251, 255 Schottariella  119 Schoutenia  390 Schouwia  418 Schradera  519 Schraderanthus  536 Schrebera  546 Schreiteria  464

Schrenkia  634 Schtschurowskia  634 Schubertia  527 Schuitemania  159 Schultesia  522 Schultesianthus  536 Schultesiophytum  139 Schulzia  634 Schumacheria  231 Schumannianthus  190 Schumanniophyton  519 Schunkea  158 Schuurmansia  308 Schuurmansiella  308 Schwackaea  355 Schwalbea  584 Schwantesia  458 Schwartzia  481 Schweiggeria  325 Schweinfurthia  554 Schwenckia  536 Schwendenera  519 Sciadocephala  609 Sciadopitys  83 Sciadotenia  218 Sciaphila  136 Scilla  7, 173–4 Scilloideae 173–4 Scindapsus  119–20 Sciothamnus  634 Scirpodendron  202 Scirpoides  202 Scirpus  202 Scitaminae  184 Scleranthopsis  448 Scleranthus  448 Scleria  202 Sclerocactus  470 Sclerocarpus  609 Sclerocarya  370 Sclerochiton  565 Sclerochloa  208 Sclerochorton  634 Sclerocroton  335 Sclerodactylon  209 Sclerolaena  451 Sclerolepis  609 Sclerolinon  337 Scleronema  390 Sclerophylax  536 Scleropogon  209 Scleropyrum  427 Sclerorhachis  609 Sclerosperma  178 Sclerostegia  451 Sclerotheca  592 Sclerotiaria  634 Scoliaxon  418 Scoliopus  150 Scoliosorus  54 Scolochloa  208 Scolophyllum  558 Scolopia  330 Scolosanthus  519 Scolymus  609–12 Scoparia  554 Scopellaria  287 Scopelogena  458 Scopolia  536 Scopulophila  448 Scorodocarpus  422 Scorodophloeus  251, 256 Scorpiothyrsus  355 Scorpiurus  252 Scorzonera  609–11 scorzonera 611

INDEX Scotch marigold  611 Scotch thistle  610 Scots lovage  635 Scottellia  324 scouring rush  24 Scribneria  208 Scrithacola  634 Scrobicaria  609 Scrofella  554 Scrophularia  557 Scrotochloa  206 Scurrula  430 scurvy grass  419 Scutachne  209 Scutellaria  577–8 Scutellarioideae 575 Scutia  271 Scuticaria  158 Scutinanthe  368 Scybalium  425 Scyphanthus  476 Scyphiphora  519 Scyphocephalium  102 Scyphonychium  373 Scyphostachys  519 Scyphostegia  330 Scyphosyce  274 Scytanthus  571 Scytopetalum  486 sea aster  610 sea beet  451 sea buckthorn  269 sea guarrie  492 sea hearts  258 sea lavender  433 sea purslane  448 sea samphire  636 sea-holly 636 seakale 418 Searsia  370 seashore palm  178 seaside grape  435 Sebaea  522 Sebastiania  335 Sebastiano-schaueria  565 Secale  208, 210 Secamone  527 Secamonoideae 527 Secamonopsis  527 Secondatia  527 Securidaca  262 Securigera  252 Securinega  341 Seddera  534 Sedella  240–1 sedge 200 Sedum  15, 240–1 seed ferns  71–3 Seegeriella  158 Seemannaralia  630 Seetzenia  247 Seetzenioideae 247 Seguieria  460 Sehima  209 Seidelia  335 Seidenfadenia  158 Seidenfadeniella  158 Seidlitzia  451 Selaginella  570 Selaginellales 10–11, 20–1 Selago  21, 557 Selenia  418 Selenicereus  470 Selenipedium  154 selfheal 578

Selinopsis  634 Selinum  634 Selliera  599–600 Selliguea  68 Selloa  609 Sellocharis  252 Semania  355 semaphore plant  259 Semecarpus  370 Semele  173–4 Semenovia  634 Semialarium  295 Semiaquilegia  220 Semiarundinaria  208 semillas de jicaro  567 Semiramisia  509 Semiria  609 Sempervivoideae 241 Sempervivum  240–1 Senaea  522 senduduk 354–5 Senecio  15, 603, 609, 614 Senefeldera  335 Senefelderopsis  335 Senegalia  253, 256 Senna  251, 259 senposai 418 Senra  389 sensitive fern  61 Senyumia  551 Seorsus  352 Sepikea  551 Septotheca  390 Septulina  430 Sequencia  195 Sequoia  84 Sequoiadendron  84 Serapias  159 Serenoa  177 Sergia  592 Serianthes  253 Sericanthe  519 Sericocarpus  609 Sericocoma  450 Sericocomopsis  450 Sericodes  247 Sericolea  300 Sericorema  450 Sericostachys  450 Seringia  389 Serissa  519 Serjania  373 Serpocaulon  70 serpolet oil  578 serradella 257 Serratula  609, 612 Serruria  226 Sersalisia  490 Sertifera  158 service tree  265 serviceberry 266 sesame  562, 610 Sesamoides  412 Sesamothamnus  562–3 Sesamum  562 Sesbania  252 Seseli  634 Seselopsis  634 Seshagiria  527 Sesleria  208 Sessea  536 Sesuvium  458 Setaria  209–10 Setariopsis  209 Setchellanthus  406, 413 Sevada  451

seven sons plant  625 Severinia  375 Seville orange  376 Seychellaria  136 Seymeria  584 Seymeriopsis  584 Seyrigia  287 Shafera  609 Shaferocharis  519 shallot 171 Shangrilaia  418 Shangwua  609 sharon fruit  492 shasta daisy  614 she cabbage  614 shea butter  490 shea tree  490 Sheareria  609 Shehbazia  418 Sheilanthera  375 shell ginger  193 shell-flower bush  560 she-oak 280–1 shepherd’s purse  419 Shepherdia  269 Sherardia  519 Sherbournia  519 Shibataea  208 shillelagh 278 Shinnersia  609 Shinnersoseris  609 Shirakiopsis  335 shirataki noodles  120 shiso 578 Shiuyinghua  581 Shizhenia  159 Shonia  335 shoo-fly plant  541 Shorea  400 Shortia  499 Shoshonea  634 shrimp plant  565 shrub tobacco  541 shrubsage 578 Shuaria  551 shungiku 610 Shuteria  252 siala tree  578 Siamanthus  192 Siamosia  450 Siapaea  609 Sibara  418 Sibiraea  264 Sibthorpia  554 Sicana  287 Sichuan pepper  378 Sichuania  527 Sicrea  390 Siculosciadium  634 Sicydium  287 Sicyos  287 Sida  390 Sidalcea  390 Sidasodes  390 Sidastrum  390 Siderasis  180 Sideritis  577–8 Sideroxylon  490 Sidotheca  435 Siebera  609, 634 Siegfriedia  271 Siemensia  519 Sievekingia  158 Sieversandreas  584 Sieversia  263 Sigesbeck, Johann

Georg 608 Sigesbeckia  609 Sigillaria  21 Sigmatanthus  375 Silaum  634 Silene  448 Silentvalleya  209 Siler  634 Siliquamomum  192 silk cotton tree  393 silk floss tree  393 silk tassel bush  515 silk vine  528 Siloxerus  609 Silphiodaucus  634 Silphium  609 silvaberry 266 silver squill  174 silver vine  504 silverseed gourd  288 silverweed 266 Silvianthemum  618 Silvianthus  545 Silviella  584 Silvorchis  168 Silybum  609–10 Simaba  380 Simarouba  380 Simethis  159 Simicratea  295 Simira  519 Simirestis  295 Simmondsia  443 Simplicia  208 Simsia  609 Sinacalia  609, 614 Sinadoxa  620 Sinapidendron  418 Sinapis  418 Sinarundinaria  208 Sinclairia  609 Sincoraea  195 Sindechites  527 Sindora  251 Sindoropsis  251 Singapore holly  318 Singchia  158 Sinningia  551 Sinoacanthus  565 Sinoadina  519 Sinoarabis  418 Sinobambusa  208 Sinocalamus  208 Sinocalycanthus  107 Sinocarum  634 Sinochasea  208 Sinocrassula  240–1 Sinodolichos  252 Sinofranchetia  217 Sinogentiana  522 Sinojackia  500 Sinojohnstonia  531 Sinolimprichtia  634 Sinomenium  218 Sinopanax  630 Sinoradlkofera  373 Sinosassafras  112 Sinosenecio  609 Sinoswertia  522 Sinowilsonia  235 sinqua 288 Siolmatra  287 Sipanea  519 Sipaneopsis  519 Sipapoantha  522 Siparuna  108

Siphanthera  355 Siphocampylus  592 Siphocodon  592 Siphocranion  577 Siphonandra  509 Siphonandrium  519 Siphoneugena  352 Siphonochiloideae 191 Siphonochilus  191 Siphonodon  295 Siphonostegia  584 sipo 381 Sipolisia  609 Siraitia  287 Sirdavidia  106 Sirhookera  158 Sirindhornia  159 Sirochloa  208 sirop de capillaire  50 sisal 174 Sison  634 Sisymbrella  418 Sisymbriopsis  418 Sisymbrium  418 Sisyndite  247 Sisyranthus  527 Sisyrinchium  167 Sisyrolepis  373 Sium  634–5 siwak 408 Skeptrostachys  159 Skiatophytum  458 Skimmia  375 skirret 634 skullcap 578 Skytanthus  527 Sladenia  487 Sleumeria  513 Sleumerodendron  226 slipperworts 551 Sloanea  300 sloe 264–5 Smallanthus  609, 611 Smeathmannia  327 Smelophyllum  373 Smelowskia  418 Smicrostigma  458 Smilacina  173 Smilax  148–9 Smirnowia  252 Smithatris  191 Smithia  252 Smithiantha  551 Smithsonia  158 Smitinandia  158 Smodingium  370 smooth pigweed  452 Smyrniopsis  634 Smyrnium  634–5 Smythea  271 snake gourd  288 snake palm  178 snapdragon 555 sneezeweed 614 sneezewort 610 snot apple  393 snow peas  254 snowbell 494 snowberry 625 Snowdenia  209 soapbark tree  248 soapberry 374 soapnut 374 soapwort 448 Soaresia  609 Sobennikoffia  158

Sobolewskia  418 Sobralia  158 society garlic  171 Socotrella  527 Socratea  178 Socratina  430 Soderstromia  209 Soejatmia  208 Soemmeringia  252 Sogerianthe  430 Sohnsia  209 Sokinochloa  208 Solanales 534 Solandra  536 Solanecio  609 Solanum  15, 536 Solaria  171 Soldanella  494 Soleirolia  276 Solena  287 Solenangis  158 Solenanthus  531 Solenidium  158 Solenocarpus  370 Solenocentrum  159 Solenogyne  609 Solenomelus  167 Solenophora  551 Solenopsis  592 Solenostemma  527 Soleredera  173 Solfia  178 Solidago  609, 613–14 solidaster 613 Soliva  609 sollya 629 Solmsia  396 Solms-Laubachia  418 Solomon’s seal  174 Solonia  494 Sommera  519 Sommerfeltia  609 Sommieria  178 Somrania  551 Sonchella  609 Sonchus  609–10, 614 Sonderina  634 Sonderothamnus  358 Sondottia  609 Sonerila  355 Sonneratia  346 Sophora  252, 259 Sophronanthe  554 Sopubia  584 Sorbaria  264 Sorbus  264 sorcerer’s tree  540 Sorghastrum  209 Sorghum  209–10 sorghum 584 Soridium  136 Sorindeia  370 Sorocea  274 Sorocephalus  226 Soroseris  609 Soroveta  204 sorrel 435–6 Soterosanthus  158 Sotoa  220 sotol 174 souari nut  314 Soulamea  380 Souliea  159 sour cherry  264 Souroubea  481 soursop 105–6

Plants of the World

787

INDEX southern beech  147, 277, 428 southernwood 610 Sowerbaea  174 sowthistle 610 soya 254 Soyauxia  232 soybean 254 Soymida  381 Spachea  318 Spananthe  634 Spaniopappus  609 Spanish cedar  381 Spanish cherry  490 Spanish flag  573 Spanish mahogany  381 Spanish moss  196 Spanish oyster  611 Spanish sage  578 Spanish salsify  612 Spanish tamarind  520 Sparattanthelium  110 Sparattosperma  567 Sparattosyce  274 Sparaxis  167 Sparganium  194 Sparganophoros  609 Sparrmannia  389 sparrowgrass 174 Spartidium  252 Spartina  209 Spartium  252, 258 Spartochloa  209 Spartothamnella  577 Spatalla  226 Spathacanthus  565 Spathantheum  119 Spathanthus  197 Spathelia  375 Spathia  209 Spathicarpa  119 Spathichlamys  519 Spathionema  252 Spathiostemon  335 Spathiphyllum  119 Spathodea  567–8 Spathoglottis  158 Spatholirion  180 Spatholobus  252 Spathra  355 spear thistle  610 spearmint 577 Specklinia  158 Speea  171 speedwell  551, 555 Spegazziniophytum  335 Speirantha  174 Spelaeanthus  551 spelt 210 Spenceria  263 Speranskia  335 Spergula  448 Spergularia  448 Spermacoce  519 Spermadictyon  519 Spermolepis  634 Spetaea  174 Sphacanthus  565 Sphaenolobium  634 Sphaeradenia  139 Sphaeralcea  390 Sphaeranthus  609 Sphaerantia  352 Sphaereupatorium  609 Sphaerobambos  208 Sphaerocardamum  418

788

Sphaerocaryum  209 Sphaerocoma  448 Sphaerocoryne  106 Sphaerolobium  252 Sphaeromeria  609 Sphaerophysa  252 Sphaeropteris  39, 43 Sphaerosacme  381 Sphaerostylis  335 Sphaerothylax  311 Sphagneticola  609, 614 Sphagnum  125 Sphallerocarpus  634 Sphalmium  226 Sphedamnocarpus  318 Spheneria  209 Sphenocentrum  218 Sphenoclea 542 Sphenodesme  576 Sphenomeris  49 Sphenopholis  208 Sphenophytes  24 Sphenopus  208 Sphenostemon  589, 618 Sphenostylis  252, 255 Sphenotoma  509 Sphinctacanthus  565 Sphinctanthus  519 Sphinctospermum  252 Sphinga  253 Sphingiphila  567 Sphyranthera  335 Sphyrarhynchus  158 Sphyrospermum  509 Spicebush 113 Spiculaea  159 spicy jatropha  336 spider flower  414 spider plant  174 Spigelia  523 spikenard  614, 623 Spilanthes  609 Spiloxene  163 spinach  449, 451–2 Spinacia  451 spindle 293 Spinifex  209 spinks 419 Spiracantha  609 Spiradiclis  519 Spiraea  264 Spiraeanthemum  299 Spiraeanthus  264 Spiraeopsis  299 Spiranthera  375 Spiranthes  159 Spiranthoideae 158 Spirematospermum  184, 187 Spirodela  119 Spirogardnera  427 Spirolobium  527 Spiroseris  609 Spirospermum  218 Spirostigma  565 Spirotecoma  567 Spirotheca  390 Spirotropis  252 Spodiopogon  209 Spondiadoideae 370 Spondianthus  340 Spondias  370 sponge gourd  289 Spongiocarpella  252 Spongiola  158 Spongiosperma  527

Christenhusz, Fay & Chase

Sporadanthoideae 203–4 Sporadanthus  204 Sporobolus  209 Sporoxeia  355 Spragueanella  430 Sprekelia  171 Sprengelia  509 spruce 80 Spryginia  418 spurge  332, 336 spurge laurel  396 Spuriodaucus  634 Spyridium  271 Squamellaria  519 Squamopappus  609 squash 287–8 squashberry 620 squill 174 squirting cucumber  289 St Bernard’s lily  174 St Helena boxwood  541 St Helena ebony  393 St Helena redwood  393 St John’s wort  313 Staavia  618 Staberoha  204 Stachyandra  539 Stachyanthus  513 Stachyarrhena  519 Stachycephalum  609 Stachydeoma  577 Stachyococcus  519 Stachyopsis  577 Stachyothyrsus  253 Stachyphrynium  190 Stachypteris  40 Stachys  575–7, 578 Stachystemon  539 Stachytarpheta  573 Stachyurus  362 Stackhousia  295 Stadiochilus  192 Stadmania  373 Staehelina  609 Staelia  519 stag horn fern  70 stag’s-horn clubmoss  19 Stahlia  253 Stahlianthus  191 Stalkya  159 Staminodianthus  252 Standleya  519 Standleyanthus  609 Stanfieldiella  180 Stangeria  74–5 Stanhopea  158 Stanleya  418 Stanmarkia  355 Stapelia  527 Stapelianthus  527 Stapfiella  327 Stapfochloa  209 Staphisagria  220 Staphylea  361 star-anise  9–4, 378 star-apple 490 star-of-Bethlehem 174 statice 433 Staticoideae 433 Staudtia  102 Staufferia  427 Stauntonia  217 Stauracanthus  252 Stauranthera  551 Stauranthus  375 Staurochlamys  609

Staurogyne  564 Stawellia  168 Stayneria  458 Steenisia  519 Stefanoffia  634 Steganotaenia  634 Steganthera  111 Stegnosperma  459 Stegolepis  197 Steinchisma  209 Steirachne  209 Steiractinia  609 Steirodiscus  609 Stelechocarpus  106 Stelestylis  139 Stelis  152, 158 Stellaria  448 Stellera  396 Stelmagonum  527 Stemodia  554 Stemodiopsis  558 Stemona  138, 510, 586 Stemonocoleus  251 Stemonoporus  400 Stemonurus  585–6 Stenachaenium  609 Stenandrium  565 Stenanona  106 Stenanthemum  271 Stenanthium  143 Stenaria  519 Stenia  158 Stenocactus  470 Stenocarpus  226 Stenocephalum  609 Stenocereus  470 Stenochlaena  55, 61 Stenocline  609 Stenocoelium  632 Stenodon  355 Stenodrepanum  253 Stenoglottis  159 Stenogonum  435 Stenogyne  577 Stenomeria  527 Stenomeris  134 Stenomesson  171 Stenopadus  603 Stenopetalum  418 Stenophalium  609 Stenops  609 Stenoptera  159 Stenorrhynchos  159 Stenosemis  634 Stenosepala  519 Stenoseris  609 Stenosolenium  531 Stenospermation  119 Stenostegia  352 Stenostephanus  565 Stenostomum  519 Stenotaenia  634 Stenotalis  204 Stenotaphrum  209, 211 Stenothyrsus  565 Stenotis  519 Stenotus  609 Stenotyla  158 Stephanachne  208 Stephania  218 Stephanocaryum  531 Stephanocereus  470 Stephanococcus  519 Stephanodaphne  396 Stephanodoria  609 Stephanomeria  609

Stephanopodium  320 Stephanostegia  527 Stephanostema  527 Stephanothelys  159 stephanotis 528 Sterculia  390 Sterculioideae 390 Stereocaryum  352 Stereochilus  158 Stereochlaena  209 Stereosandra  158 Stereospermum  567–8 Sterigmapetalum  305 Sterigmostemum  418 Steriphoma  413 Sternbergia  171 Stetsonia  470 Steudnera  119 Stevenia  418 Steveniella  159 Stevensia  519 Stevia  609, 611 stevia  611, 614 Steviopsis  609 Stewartia  496 Stewartiella  634 Steyerbromelia  195 Steyermarkia  519 Steyermarkina  609 Steyermarkochloa  209 Sticherus  32 Stichianthus  519 Stichoneuron  138 Stichorkis  158 Stictocardia  534 Stifftia  603 Stiftioideae 602 Stigmaphyllon  318 Stigmatodactylus  159 Stigmatodon  195 Stigmatopteris  66 Stigmatorhynchus  527 Stilbanthus  450 Stilbe  560 Stilbocarpa  634 Stillingia  335 Stilpnogyne  609 Stilpnolepis  609 Stilpnopappus  609 Stilpnophyllum  519 Stimpsonia  494 stinking roger  612 Stipa  208, 211 Stipagrostis  209 Stipecoma  527 Stipulicida  448 Stirlingia  226 Stirtonanthus  252 Stixis  412 Stizolophus  609 Stizophyllum  567 stock 419 Stocksia  373 Stockwellia  352 Stoebe  609 Stoeberia  458 Stoibrax  634 Stokesia  609, 614 Stolzia  158 Stomatanthes  609 Stomatium  458 Stomatochaeta  603 stoneseed 532 Stonesia  311 Stonesiella  252 Storckelia  251

stork’s bill  343 Storthocalyx  373 Stramentopappus  609 Strangea  226 Strasburgeria  360 Stratiotes  120 Straussiella  418 strawberry 265–6 strawberry guava  352 strawberry tomato  538 strawberry tree  509 Streblacanthus  565 Streblorrhiza  252 Streblosa  519 Streblosiopsis  519 Streblus  274 Strelitzia  175, 184–5 Strempeliopsis  527 Strephonema  346 Streptanthus  418 Streptocarpus  551 Streptocaulon  527 Streptochaeta  206 Streptoechites  527 Streptoglossa  609 Streptogyna  208 Streptolirion  180 Streptoloma  418 Streptolophus  209 Streptopetalum  327 Streptopus  149 Streptosiphon  565 Streptosolen  536 Streptostachys  209 Streptothamnus  421 stretchberry 546 Striga  584 string beans  254 string-of-pearls 614 striped hemlock  637 striped squill  174 Strobilacanthus  565 Strobilanthes  565 Strobilanthopsis  565 Strobilopsis  557 Stromanthe  190 Stromatopteris  32 Strombocactus  470 Strombosia  422 Strombosioideae 422 Strombosiopsis  422 Strongylodon  252, 259 Strophanthus  527 Strophioblachia  335 Strophostyles  252 Strotheria  609 Strumaria  170–1 Strumpfia  519 Struthanthus  430 Struthiola  396 strychnine 523–4 Strychnopsis  218 Strychnos  523 Stryphnodendron  253 Stuartina  609 Stuckenia  129 Stuckertiella  609 Stuessya  609 stuffing gourd  288 Stuhlmannia  253 Sturt’s desert rose  393 Stussenia  355 Stylapterus  358 Stylidium  595–6 Stylisma  534 Stylobasium  260

INDEX Styloceras  228 Stylochaeton  119 Stylocline  609 Stylogyne  494 Stylomecon  216 Stylophorum  216 Stylosanthes  252, 257 Stylosiphonia  519 Stylotrichium  609 Stypandra  168 Styphelia  509 Styphelioideae 509 Styphnolobium  252, 259 Styppeiochloa  209 Styrax  500 Suaeda  451 Suarezia  158 Suberanthus  519 Subulina  448 Succisa  623, 625 Succisella  623 Succowia  418 Suchtelenia  531 Suckleya  451 Sucrea  208 Sudamerlycaste  158 Suddia  206 Suessenguthia  565 Suessenguthiella  462 sugar beet  451 sugar maple  374 sugarcane  116, 210–11, 560 sugarplum 341 Sukhorukovia  451 Suksdorfia  239 Sulla  252, 257 sulla clover  257 Sullivantia  239 sumac 371 Sumatroscirpus  202 Sumbaviopsis  335 summer cypress  452 summer squash  288 Summerhayesia  158 sun marigold  613 sunberry 266 Sundacarpus  82 sundew 436–7 sunflower  611, 614 sunn hemp  258 sunrise lime  376 Suregada  335 Suriana  260, 365 Suriname cherry  352 Surreya  450 Susswassertang 63 susumber 539 Sutera  557 Sutherlandia  252 Sutrina  158 Suzukia  577 Svenkoeltzia  159 Svitramia  355 Swainsona  253, 259 Swallenia  209 swamp thistle  610 swamp-spiraea 266 Swartzia  253 swede 418 sweet alyssum  419 sweet calabash  328 sweet cherry  264 sweet chestnuts  278 sweet cicely  636 sweet corn  210

sweet corn root  190 sweet gale  279 sweet lemon  378 sweet marjoram  578 sweet orange  376 sweet pea  258–9 sweet potato  534 sweet sultan  613 sweet William  448 sweetgum 233–4 Sweetia  253 sweetroot 637 sweetsop 106 Swertia  522 Swietenia  381 Swinglea  375 Swintonia  370 Swiss-cheese plant  120 Syagrus  178 sycamore  224, 274 Sycopsis  235 Sylvichadsia  253 Symbolanthus  522 Symmeria  434 Symmerioideae 434 Symonanthus  536 Sympegma  451 Sympetalandra  253 Symphionema  226 Symphonia  309 Symphorema  576 Symphorematoideae 575 Symphoricarpos  623, 625 Symphyandra  593 Symphyllocarpus  609 Symphyllophyton  522 Symphyochlamys  390 Symphyoloma  634 Symphyonematoideae 226 Symphyopappus  609 Symphyotrichum  609, 614 Symphysia  509 Symphytum  531 Symplectrodia  209 Symplocarpus  119 Symplococarpon  489 Symplocos  498 Synandra  577 Synandrodaphne  396 Synandrospadix  119 Synaphea  226 Synapsis  571 Synaptantha  519 Synaptolepis  396 Synaptophyllum  457 Syncalathium  609 Syncarpha  609, 613 Syncarpia  352 Syncephalum  609 Synclinostyles  634 Synclisia  218 Syncolostemon  577 Syncretocarpus  609 Syndyophyllum  335 Synechanthus  178 Synedrella  609 Synedrellopsis  609 Syneilesis  609, 614 Syngonanthus  198 Syngonium  119 Syngramma  120 Synima  373 Synotis  609 Synoum  381 Synsepalum  490 Synstemon  418

Synthlipsis  418 Syntriandrium  218 Syntrichopappus  609 Synurus  609 Syreitschikovia  609 Syringa  546 Syringantha  519 Syringodea  167 Syringodium  131 Syrmatium  253 Syrrheonema  218 Systellantha  494 Systeloglossum  158 Systenotheca  435 Syzygium  351–3 Szovitsia  634   tabardillo 614 Tabebuia  566–8 Tabernaemontana  527 Tabernanthe  527 tabonuco 368 Tacarcuna  341 tacay nut  335 Tacazzea  527 Tacca  599 Taccarum  119 Tachia  522 Tachiadenus  522 Tachigali  253, 257 Tacinga  470 Tadehagi  253 Taeniatherum  208 Taenidia  634 Taeniophyllum  158 Taeniorhachis  209 Taeniorrhiza  158 Taenitis  52 Tagetes  609–10, 612–3 Tahina  177 tahini 563 Tahiti lime  378 Tahitia  389 Tahitian chestnut  255 Tahitian vanilla  152 Tainia  158 Taiwania  84 takamaka 310–11 Takhtajan, Armen Leonovich 97 Takhtajania  96–7 Takhtajaniella  418 Talamancalia  609 Talauma  103 Talbotia  137 Talbotiella  251 Talinopsis  468 Talinum  467 Talipariti  389 Talisia  373 tallest tree  72 Taltalia  144 Tamamschjanella  634 Tamamschjania  634 Tamananthus  609 tamarillo 539 tamarind  255, 520 Tamarindus  251, 255 tamarisk 432 Tamarix  432 Tamaulipa  609 Tamayorkis  158 tambalacoque 491 Tambourissa  88 Tamijia  191, 193 Tamijioideae 191

Tamilnadia  519 tammana 339 Tammsia  519 Tamonea  573 tampoi 341 Tamridaea  519 Tamus  134 Tanacetopsis  609 Tanacetum  609, 610, 612–3 Tanaecium  567 Tanakaea  239 tangelo 377 tangerines 378 tangors 377 tanner bush  286 tanner’s dock  435 Tannodia  335 Tanquana  458 tansy 612 tapegrass 125 Tapeinia  167 Tapeinidium  49 Tapeinochilos  191 Tapeinosperma  494 Tapeinostemon  522 Tapheocarpa  180 Tapinanthus  430 tapioca 335 Tapirira  370 Tapirocarpus  367 Tapiscia  384 Taplinia  609 Tapoides  335 Taprobanea  158 Tapura  320 Tara  253, 257 Taralea  253 Tarasa  390 tarata 629 Taravalia  375 Taraxacum  609–10 Taraxia  349 Taraxis  204 Tarchonanthus  603, 612 Tarenaya  414 Tarenna  519 Tarennoidea  519 Tarigidia  209 Tarlmounia  609 taro  116, 119, 513 tarragon 611–12 tarthuth 243–4 tarweed 611 Tasmannia  96 Tassadia  527 Tateanthus  355 Tatianyx  209 Tauschia  634 Tavaresia  527 Taverniera  253 tawari  359, 360 Taxandria  352 Taxillus  430 Taxodium  71, 84 Taxus  86, 149 tayberry 266 Tayloriophyton  355 Tchihatchewia  418 té de burro  344 tea oil  496 tea plant  496 Teagueia  158 teak 578 teasel  623, 625 tea-tree oil  353

Teclea  375 Tecleopsis  375 Tecoma  567–8 Tecomanthe  567 Tecomella  567–8 Tecophilaea  164–5 Tectaria  63, 67 Tectarioideae  63, 66–8 Tectonoideae 576 Tecticornia  451 Tectiphiala  178 Tectona  575–6, 578 Tecunumania  287 Teedia  557 Teesdalia  418 tef 210 Tegicornia  451 Tehuana  609 Teijsmanniodendron  576 Teixeiranthus  609 Telanthophora  609 Telectadium  527 Telekia  609, 614 Telephiae  240–1 Telephium  448 Telesonix  239 Telfairia  287 Telipogon  158 Telitoxicum  218 Tellima  239 Telmatoblechnum  61 Telmatophila  609 Telopea  226 Telosma  527 Temburongia  208 Temnadenia  527 Temnocalyx  519 Temnopteryx  519 Temochloa  208 Templetonia  253 Tenaxia  209 tendu 492 Tengia  551 Tennantia  519 Tenrhynea  609 teosinte 210 tepary bean  254 Tephroseris  609 Tephrosia  253 Tepualia  352 Tepuia  509 Tepuianthoideae 396 Tepuianthus  396 tequila 174 Teramnus  253 Teratophyllum  66–7 Terminalia  346 Terminaliopsis  346 Terniopsis  311 Ternstroemia  489 Tersonia  411 Tessaria  609 Tessiera  519 Tessmannia  251 Tessmanniacanthus  565 Tessmannianthus  355 Testudinaria  134 Testulea  308 Tetilla  239, 344 Tetraberlinia  251 Tetracarpaea  242 Tetracentron  226–7 Tetracera  231 Tetrachaete  209 Tetrachne  209 Tetrachondra  548

Tetrachyron  609 Tetraclea  577 Tetraclinis  84 Tetracme  418 Tetracoccus  539 Tetractomia  375 Tetradenia  577–8 Tetradiclis  247, 366 Tetradium  375 Tetradoxa  619–20 Tetradymia  609 Tetraena  247 Tetragastris  368 Tetragonia  458 Tetragonolobus  253, 255 Tetragonotheca  609 Tetralix  389 Tetralocularia  534 Tetrameranthus  106 Tetramerista  482 Tetramerium  565 Tetramicra  158 Tetramolopium  609 Tetranema  554 Tetraneuris  609 Tetranthus  609 Tetrapanax  630 Tetraperone  609 Tetraphyllaster  355 Tetraphyllum  551 Tetraplasandra  629 Tetrapleura  253 Tetrapogon  209 Tetrapollinia  522 Tetrapterocarpon  253 Tetrapteron  349 Tetrapterys  318 Tetraria  202 Tetraselago  557 Tetrasida  390 Tetrasiphon  295 Tetraspidium  584 Tetrastigma  244 Tetrastylidium  422 Tetrasynandra  111 Tetrataenium  634 Tetrataxis  346 Tetratheca  300 Tetrathylacium  330 Tetraulacium  554 Tetrazygia  355 Tetroncium  127 Tetrorchidium  335 Teucridium  577 Teucrium  577–8 Teuscheria  158 Texas mahonia  220 Texas ranger  557 Texsel greens  418 Teyleria  253 Thai basil  578 Thaia  158 Thailentadopsis  253 Thalassia  125 Thalassocharis  131 Thalassodendron  131 thale cress  419 Thaleropia  352 Thalia  190 Thalictroideae 220 Thalictrum  220, 222 Thaminophyllum  609 Thamnea  618 Thamnocalamus  208 Thamnochortus  204 Thamnoldenlandia  519

Plants of the World

789

INDEX Thamnopteris  58 Thamnosciadium  634 Thamnoseris  609 Thamnosma  375 Thapsia  634 Thaspium  634 thatch screwpine  140 Thaumasianthes  430 Thaumastochloa  209 Thaumatocaryon  531 Thaumatococcus  190 Thawatchaia  311 Thecacoris  340 Thecagonum  519 Thecanthes  396 Thecocarpus  634 Thecopus  158 Thecostele  158 Thedachloa  209 Theilera  592 Thelasis  158 Thelepogon  209 Thelesperma  609 Thelethylax  311 Theligonum  519 Thelionema  168 Thelocactus  470 Thelymitra  159 Thelypodiopsis  418 Thelypodium  418 Thelypteridoideae 23, 55, 58–9 Thelypteris  15, 55, 59 Thelyschista  159 Themeda  209 Themistoclesia  509 Thenardia  527 Theobroma  389 Theophrasta  494 Theophrastoideae 494 thepelakano 612 Thereianthus  167 Theriophonum  119 Thermopsis  253, 259 Theropogon  174 Therorhodion  509 Thesidium  427 Thesieae 427 Thesium  427 Thesmophora  560 Thespesia  390 Thespidium  609 Thespis  609 Thevenotia  609 Thevetia  527 Thibaudia  509 Thilachium  413 Thiloa  346 Thinicola  253 Thinopyrum  208 Thinouia  373 Thiollierea  519 Thiseltonia  609 Thismia  134, 142 Thismieae  134 Thladiantha  287 Thlaspi  418 Thlaspiceras  418 Thogsennia  519 Thomasandersia  564, 571 Thomasia  389 Thompsonella  240–1 Thonningia  425 Thoracocarpus  139 Thoreauea  527 thorn-apple 540

790

Thorncroftia  577 Thorne, Robert Folger 312 Thornea  313 thoroughwort 614 thorow-wax 637 Thottea  101 Thouinia  372–3 Thouinidium  373 Thrasya  209 Thrasyopsis  209 Threlkeldia  451 thrift 432–3 Thrinax  177 Thrixspermum  158 Thryallis  318 Thryptomene  352 Thuarea  209 Thuja  84 Thujopsis  84 Thulinia  159 Thunbergia  564–5 Thunbergianthus  584 Thunbergioideae 564 Thunia  158 Thuniopsis  158 Thurnia  200 Thurovia  609 Thurya  448 Thuspeinanta  577 Thylacopteris  70 Thylacospermum  448 Thymbra  577 thyme  573, 577–8 Thymelaea  396 Thymelaeoideae 396 Thymophylla  609 Thymopsis  609 Thymus  575, 577–8 Thyridachne  209 Thyridia  580 Thyridolepis  209 Thyrocarpus  531 Thyrsanthemum  180 Thyrsanthera  335 Thyrsodium  370 Thyrsopteridoideae 39–40 Thyrsopteris  39–41 Thyrsosalacia  295 Thyrsostachys  208 Thysanocarpus  418 Thysanoglossa  158 Thysanolaena  209 Thysanostemon  309 Thysanostigma  565 Thysanotus  174 Thyselinum  634 Tianschaniella  531 Tiarella  239 Tibetia  253 Tibetoseris  609 Tibouchina  355 Tibouchinopsis  355 tickseed 614 Ticorea  375 Tidestromia  450 tidy-tips 613 Tieghemella  490 Tietkensia  609 tigernuts 202 Tigridia  167 Tilesia  609 Tilia  389 Tiliacora  218 Tilioideae 389 Tillandsia  195–6

Christenhusz, Fay & Chase

Timonius  519 Tina  373 Tinadendron  519 Tinantia  180 tinda 288 tindori 288 tinker’s weed  625 Tinnea  577 Tinomiscium  218 Tinopsis  373 Tinospora  218 Tintinnabularia  527 tiny mice  347 Tipuana  253 Tipularia  158 Tiputinia  134 Tiquilia  531 Tirania  412 Tirpitzia  337 Tisonia  330 titan arum  120 Titanopsis  458 Titanotrichum  551 titberry 374 Tithonia  609, 614 tlanochtle 539 Tmesipteris  26, 143 toadflax  555 tobacco 539 Tobagoa  519 Tocantinia  171 Tococa  355 Tocoyena  519 Todaroa  634 Toddalia  375 Toddaliopsis  375 toddy 178 Todea  26 Toechima  373 Tofieldia  121 tokitoki 504 Toliara  209 Tolmiea  239 Tolpis  609 Tolumnia  158 Tolypanthus  430 Tomanthera  584 tomatillo 538 tomato  1, 536, 541 Tomentaurum  609 tomillo salsero  578 Tommasinia  634 Tomostima  418 Tomzanonia  158 Tonduzia  527 Tonella  554 Tonestus  609 tong ho  610 Tongoloa  634 Tonina  198 tonka bean  256 Tontelea  295 Toona  381 toothbrush tree  408 topinambour 611 Topobea  355 toquilla straw  139 torchwood 378 Tordyliopsis  634 Tordylium  634, 636 Torenia  558 Torilis  634 Tornabenea  634 Toronia  226 Torralbasia  295 Torreya  86

Torreyochloa  208 Torricellia  627 Tortuella  519 totora reeds  202 Touchardia  276 touch-me-not  259, 497 Toulicia  373 Tournefortia  531 Tourneuxia  609 Tournonia  466 Touroulia  307 Tourrettia  567 Toussaintia  106 Tovaria  409, 413 Tovarochloa  208 Tovomita  309 Tovomitopsis  309 townhall clock  621 Townsendia  609 Townsonia  159 Toxicodendron  370 Toxicoscordion  143 Toxosiphon  375 Tozzia  584 Trachelanthus  531 Trachelium  592–3 Trachelospermum  527 Trachoma  168 Trachyandra  158 Trachycarpus  177 Trachydium  634 Trachymene  629–31 Trachyphrynium  190 Trachypogon  209 Trachys  209 Trachyspermum  634, 636 Trachystemon  531 Trachystigma  551 Trachystoma  418 Trachystylis  202 Tractocopevodia  375 Tracyina  609 Tradescantia  180 Traganopsis  451 Traganum  451 Tragia  335 Tragiella  335 Tragopogon  609, 611 Tragus  209 trailing lantana  573 Trailliaedoxa  519 Transberingia  418 Trapa  346 Trapella  554 traro-voqui 396 Trattinnickia  368 Traubia  171 Traunsteinera  159 Trautvetteria  220 traveller’s joy  222 traveller’s palm  185 Traversia  609 treasure flower  613 Treculia  274 tree cotton  392 tree fern  39, 43–4 tree hollyhock  392 tree jasmine  568 tree marigold  614 tree poppy  216 tree spinach  391 tree tomato  539 tree-dahlia 614 tree-of-heaven  378, 380 Treichelia  592 Trema  273

Tremandra  300 Trematolobelia  592 Trembleya  355 Tremulina  204 Trepadonia  609 Trepocarpus  634 Treutlera  527 Trevesia  630 Trevoa  271 Trevoria  158 Triadenum  313 Triadica  335 Triaenophora  582–4 Triainolepis  519 Trianaea  536 Trianaeopiper  99 triangle palm  178 Trianoptiles  202 Triantha  121 Trianthema  458 Triaspis  318 Tribeles  615–16 Tribolium  209 Tribonanthes  184 Tribounia  551 Tribulocarpus  458 Tribuloideae 247 Tribulopsis  247 Tribulus  247 Tricalysia  519 Tricardia  531 Tricarpelema  180 Triceratella  180 Triceratorhynchus  158 Trichadenia  324 Trichanthemis  609 Trichanthera  565 Trichanthodium  609 Trichaulax  565 Trichilia  381 Trichipteris  43 Trichlora  171 Trichloris  209 Trichocalyx  565 Trichocentrum  158 Trichocephalus  271 Trichoceros  158 Trichocladus  235 Trichocline  603 Trichocoronis  609 Trichocoryne  609 Trichodesma  531 Trichodiadema  458 Trichoglottis  158 Trichogonia  609 Trichogoniopsis  609 Trichogyne  609 Tricholaena  209 Tricholaser  634 Tricholepidium  79 Tricholepis  609 Trichomanes  31 Trichoneura  209 Trichopetalum  174 Trichophorum  202 Trichopilia  158 Trichopteryx  209 Trichoptilium  609 Trichopus  134–5 Trichosalpinx  158 Trichosanchezia  565 Trichosandra  527 Trichosanthes  287 Trichoschoenus  202 Trichoscypha  370 Trichospermum  389

Trichospira  609 Trichostachys  519 Trichostema  577–8 Trichostephanus  330 Trichostigma  460 Trichotolinum  418 Trichotosia  158 Trichuriella  450 Tricliceras  327 Triclisia  218 Tricomaria  318 Tricoryne  168 Tricostularia  202 Tricyclra  287 Tricyrtis  149–50 Tridactyle  158 Tridactylina  609 Tridax  609 Tridens  209 Tridesmostemon  490 Tridimeris  106 Tridynamia  534 Trieenea  557 Trientalis  494 Trifidacanthus  253 Triflorensia  519 trifoliate orange  378 Trifolium  253, 257 Trigastrotheca  462 Triglochin  125, 127 Trigonachras  373 Trigonella  253, 256 Trigonia  319 Trigoniastrum  319 Trigoniodendron  319 Trigonobalanus  278 Trigonocapnos  216 Trigonocaryum  531 Trigonopleura  331 Trigonopterum  609 Trigonopyren  519 Trigonosciadium  634 Trigonospermum  609 Trigonostemon  335 Trigonotis  531 Trigynaea  106 Trihaloragis  243 Trihesperus  174 Trilepidea  430 Trilepis  202 Trilepisium  274 Trilisa  609 Trillium  143 Trilobachne  209 Trimenia  618 Trimeria  330 Trimezia  167 Trinia  634 Triniochloa  208 Triodanis  592 Triodia  209 Triolena  355 Triommia  368 Trioncinia  609 Triopterys  318 Triosteum  623, 625 Triphasia  375 Triphora  158 Triphyophyllum  440–1 Triphysaria  584 Triplachne  208 Tripladenia  146 Triplarina  352 Triplaris  435 Triplasiella  209 Triplasis  209

INDEX Triplateia  448 triplet lily  174 Tripleurospermum  609 Triplocephalum  609 Triplochiton  390 Triplophyllum  67 Triplopogon  209 Triplostegia  623 trip-madame 241 Tripodanthus  430 Tripodion  253 Tripogandra  180 Tripogon  209 Tripogonella  209 Tripolium  609–10 Tripora  577 Tripsacum  209 Tripteris  609 Tripterococcus  295 Tripterodendron  373 Tripterospermum  522 Tripterygium  295 Triptilodiscus  609 Triraphis  209 Triscenia  209 Trischidium  253 Trisepalum  551 Trisetaria  208 Trisetella  158 Trisetum  208 Tristachya  209 Tristagma  171 Tristania  352 Tristaniopsis  352 Tristellateia  318 Tristemma  355 Tristemonanthus  295 Tristerix  430 Tristicha  311 Tristichoideae 311 Tristira  373 Tristiropsis  373 Trisyngyne  277 Triteleia  174 Triteleiopsis  174 Trithecanthera  430 Trithrinax  177 Trithuria  14, 89 triticale 210 ×Triticosecale  210 Triticum  208–10 Tritonia  167 Tritoniopsis  167 Triumfetta  389 Triunia  226 Triuris  136 Trivalvaria  106 Trixis  603 Trizeuxis  158 Trochetia  390, 594 Trochetiopsis  390 Trochiscanthes  634 Trochocarpa  509 Trochodendrales 226 Trochodendron  226–7 Trochomeria  287 Trochomeriopsis  287 Troglophyton  609 Trollius  220, 222 Tromotriche  527 Tromsø palm  637 Tropaeolum  403 Tropheastrum  403 Trophis  274 Tropidia  158 Tropidocarpum  418

trumpet creeper  568 trumpet roble  568 Trungboa  575, 577 Trymalium  271 Trymatococcus  274 Tryonia  52 Tryssophyton  355 Tsaiorchis  159 Tsebona  490 Tsingya  373 Tsoala  536 Tsoongia  576 tsubaki oil  497 Tsuga  80 tualang 259 Tuberaria  398 Tuberculocarpus  609 Tuberolabium  168 tuberose 174 Tuberostylis  609 Tubocapsicum  536 Tuctoria  209 tucum 178 tucuso 139 Tugarinovia  609 Tulbaghia  170–1 Tulbaghieae  171 tulip  87, 150 Tulipa  149–50, 310 tulipwood 374 Tulista  158 Tumamoca  287 Tumboa  76 tummelberry 266 Tupeia  430 tupelo tree  471 Tupistra  174 Turaniphytum  609 Turbina  534 Turgenia  634 turmeric 192 Turnera  327 Turneroideae 327 turnip 418 turnip-rooted chervil  634 Turpinia  361 Turraea  381 Turraeanthus  381 Turricula  531 Turrillia  226 Turritis  418 turtlehead 555 Tussilago  609, 612 Tuxtla  609 tweedia 528 twenty-men tree  338 twin flower  622, 625 twinspur 557 Tylecodon  241 Tyleria  308 Tylerianthus  238 Tylopsacas  551 Tylosema  250, 254 Tylostigma  159 Tynanthus  567–68 Typha  193–4 Typhonium  119 Typhonodorum  119 Tyrbastes  204 Tyrimnus  609 Tzeltalia  536   Uapaca  341 Uebelinia  448 Uebelmannia  470

Uechtritzia  603 Ugamia  609 ugli fruit  377 Ugni  352 Uittienia  251 Uladendron  390 ulanzi 211 ulé rubber  274 Uleanthus  253 Ulearum  119 Uleiorchis  158 Uleophytum  609 Ulex  253, 258 ulluco 466 Ullucus  466 Ulmus  272 Umbellifera  637 Umbellularia  112 Umbilicus  240–1 umbrella-plant 202 Umtiza  253, 258 Uncaria  519 Uncarina  562–3 Uncifera  158 Uncinia  202 Ungeria  390 Ungernia  171 Ungnadia  373 Ungulipetalum  218 unicorn root  133 Unigenes  592 Uniola  209 Unonopsis  106 Unxia  609 Upuna  400 Uranthoecium  209 Uraria  253 Urbananthus  609 Urbanodendron  112 Urbinella  609 Urceola  527 Urceolina  171 urd bean  255 Urelytrum  209 Urena  390 Urera  276 Urginea  173 Uribea  253 Urmenetia  603 urn vine  529 Urobotrya  424 Urocarpus  375 Urochloa  209–10 Urochondra  209 Urodon  253 Urogentias  522 Urolepis  609 Uromyrtus  352 Urophyllum  519 Urophysa  220 Uroskinnera  554 Urospatha  119 Urospermum  609 Ursinia  609 Ursulaea  195 Urtica  276 Urvillea  373 Usteria  523 Utleya  509 Utricularia  196, 569–70 Utsetela  274 Uvaria  106 Uvariastrum  106 Uvariodendron  106 Uvariopsis  106 Uvedalia  580

uvilla 276 Uvularia  145–6   Vaccaria  448 Vaccinioideae 509 Vaccinium  509 Vachellia  253, 256 Vagaria  171 Vahadenia  527 Vahlia  533 Vahliales 533 Valantia  519 Valdivia  616 valerian 623 Valeriana  623, 625 Valerianella  623 Valerianoideae 623 Valiha  208 Vallariopsis  527 Vallaris  527 Vallea  300 Vallesia  527 Vallisneria  125 Vanasushava  634 Vancouveria  219 Vanda  152, 158 Vandasina  253 Vandellia  558 Vandopsis  158 Vangueria  519 Vangueriella  519 Vangueriopsis  519 Vanheerdea  458 Vanhouttea  551 Vania  418 Vanilla  152, 154 vanilla 152 Vanilloideae  152, 154 Vanoverberghia  192 Van-royena  490 Vantanea  323 Vanwykia  430 Vanzijlia  458 Vargasiella  158 Varilla  609 Varthemia  609 Vasconcellea  405 Vaseyochloa  209 Vasivaea  389 Vasqueziella  158 Vassilczenkoa  433 Vassobia  536 Vatairea  253 Vataireopsis  253 Vateria  400 Vateriopsis  400 Vatica  400 Vatovaea  253 Vaughania  253 Vaupesia  335 Vauquelinia  264 Vavaea  381 Vavara  565 Vavilovia  253 Vegaea  494 vegetable fern  55 vegetable ivory  179 vegetable lamb  42 vegetable oyster  611 vegetable silk  528 vegetable spaghetti  288 vegetable sponge  287 veitchberry 266 Veitchia  178 Velascoa  363 velcro 615

Veldkampia  209 Velezia  448 Vella  418 Velleia  600 Vellereophyton  609 Vellosiella  584 Vellozia  137 Vellozioideae 137 Veltheimia  174 velvet apple  492 velvet tamarind  255 velvet-fruited zanha  373 velvetleaf 393 Venegasia  609 Ventenata  208 Ventilago  271 Venus’ fly trap  437–8, 483 Vepris  375 Veratrilla  522 Veratrum  142–3, 522 Verbascum  557 Verbena  573 Verbesina  609, 614 Verhuellia  99 Vernicia  335 Vernonanthura  609 Vernonia  603, 609, 612 Vernoniastrum  609 Vernoniopsis  609 Veronica  554 Veronicastrum  554 Verreauxia  600 Verrucularia  318 Verschaffeltia  178 Verticordia  352 vervain 572–3 Vesalea  622 Veselskya  418 Vesper  634 Vesselowskya  299 Vestia  536 vetch  255, 257 vetiver 211 Vexatorella  226 Veyretella  159 Veyretia  159 Viburnum  619–22 Vicatia  634 Vicia  253–4, 257, 259, 577 Victoria  91 Vidalasia  519 Vieraea  609 Viereckia  609 Vietnamese coriander  436 Vietnamocalamus  208 Vietnamocasia  119 Vietnamochloa  209 Vietnamosasa  208 Vietsenia  355 Vigethia  609 Vigna  253–5 Viguiera  609 Viguieranthus  253 Viguierella  209 vihta 282 Villadia  240–1 Villanova  609 Villaria  519 Villarrealia  634 Villarsia  598 Villasenoria  609 Viminaria  253 vinblastine 528 Vinca  527

Vincetoxicum  527 vine spinach  452 Vinicia  609 Vinkia  243 Viola  325 violet  324–5, 551 violet cress, violet cabbage 419 violet tubeflower  541 Virectaria  519 Virgilia  253, 259 Virginia fanpetals  393 Virginian snakeroot  101 Viridivia  327 Virola  102 Virotia  226 Viscainoa  247 Visceae 427 Viscum  427 Vismia  313 Vismianthus  296 Visnea  489 Vitales 244 Vitekorchis  158 Vitellaria  490 Vitellariopsis  490 Vitex  576, 578 Viticipremna  576 Viticoideae 575–6 Vitis  244–5 Vitoideae 245 Vittadinia  609 Vittaria  50, 54 Vittarioideae  50, 54 Vittetia  609 Viviania  344 Voacanga  527 Voanioala  178 Voatamalo  539 Vochysia  350 Vogtia  609 Volkameria  577 Volkensinia  450 Volkiella  202 Volutaria  609 Vossia  209 Votomita  355 Votschia  494 Vouacapoua  253 Vouarana  373 Voyria  522 Voyriella  522 Vriesea  195 Vrydagzynea  159 Vulpia  208 Vulpiella  208   Wachendorfia  184 Wadithamnus  450 Wahlenbergia  592–3 Waireia  159 Waitzia  609 Wajira  253 wall pepper  241 wallaba oil  256 Wallacea  308 Wallaceodendron  253 Wallaceodoxa  178 wallapatta 396 Wallenia  494 Walleria  164–5 Wallichia  177 walnut 279–80 Walsura  381 Walteranthus  411 Waltheria  389

Plants of the World

791

INDEX Waltillia  195 Wamalchitamia  609 Wandersong  519 Wangenheimia  208 Warburgia  95 Warczewiczella  158 Warea  418 Warionia  609 warmia berry  371 Warmingia  158 Warneckea  355 Warnockia  577 warratah 226 Warrea  158 Warreella  158 Warreopsis  158 Warszewiczia  519 wasabi  419, 436 Washingtonia  177 water apple  352 water caltrop  347 water chestnut  347 water ferns  37 water hyacinth  183 water hyssop  555 water lemon  327 water mimosa  259 water pepper  436 water shield  90 water spinach  535 water violet  494 water wisteria  565 waterberry 352 watercress 418 waterleaf 467 waterlily 91 watermelon 288 watermint 577 waterwheel 438 water-willow 568 Watsonia  167 wattakaka 527 wax gourd  288 Weberbauera  418 Weberbauerella  253 Weberbauerocereus  470 Weberocereus  470 Weddellina  311 Weddellinoideae 311 Wedelia  609 weeping willow  330 Wehrwolfea  372 Weigela  622, 625 Weinmannia  299 Welchiodendron  352 weld 412 Weldenia  180 Welfia  178 Wellstedia  531 Welsh onion  171 Welwitschia  71, 76 Welwitschiales  10–11, 76 Welwitschiella  609 Wenchengia  577 Wendlandia  519 Wendlandiella  178 Wendtia  344 Wenzelia  375 Werauhia  195 Wercklea  390 Werneria  609 West Indian ebony  258 West Indian vanilla  152 Western Australian Christmas tree  430 Westoniella  609

792

Westringia  576 Wetria  335 Wettinia  178 Wettsteiniola  311 wheat  1, 116, 209, 210 Whipplea  473 white aster  614 white cedar  381 white horehound  578 white pear  514 white pepper  99 white rose of York  267 white turmeric  192 white willow  330 white zapote  378 whitebeam 265 whitecurrant 238 Whiteochloa  209 Whiteodendron  352 Whitesloanea  527 whitestar potato  534 whitewood  103, 300, 428, 614 Whitfieldia  565 Whitmorea  585 Whittonia  232 whitty pear  265 Whyanbeelia  539 Whytockia  551 Wiborgia  253 Wiborgiella  253 Widdringtonia  84 Widgrenia  527 widow flower  625 Wielandia  341 Wiesneria  122 Wigandia  531 Wightia  581 Wikstroemia  396 Wilbria  287 wild chamomile  611 wild dagga  578 wild ginger  101 wild goose plum  265 wild mango  371 wild raisin  620 Wilhelmsia  448 Wilkesia  609 Wilkiea  111 Willdenowia  204 Willemetia  609 Williamodendron  112 Willisia  311 Willkommia  209 willow 330 Willughbeia  527 Wilsonia  534 Wimmerella  592 Wimmeria  295 window plant  458 Windsorina  197 wine  239, 245–5, 266, 278, 282, 299, 306, 318, 435, 459, 510, 610, 620, 635 wineberry 266 Winifredia  204 Winitia  106 Winklerella  311 winter aconite  222 winter cherry  541 winter heliotrope  614 winter marjoram  578 winter melon  288 winter purslane  464 winter savory  578

Christenhusz, Fay & Chase

winter squash  288 winter stevia  614 Winter’s bark  95–6 wintercress 419 wintergreen 509 wishbone flower  558 Wislizenia  414 Wissadula  390 Wisteria  253, 259 witch hazel  235 witch’s cream  540 Withania  536 Witheringia  536 witloof 610 Witsenia  165, 167 Wittmackanthus  519 Wittrockia  195 Wittsteinia  597 woad  257, 419 Wodyetia  178 Woehleria  450 wolfberry 539 Wolffia  119, 294 Wolffiella  119 Wollastonia  609 Wollemi pine  81 Wollemia  81 wollum wollum wood  629 wonga-wonga vine  568 wood horsetail  25 wood sorrel  297 woodbine 623 Woodburnia  630 Woodfordia  346 woodruff 520 Woodsia  60 Woodsioideae  55, 59–60 woodvamp 474 Woodwardia  61 Wooleya  458 Woollsia  509 woollystar 485 Worcesterberry 238 Worcestershire Sauce  636 wormseed 452 wormwood 610 Worsleya  171 Wrightia  527 Wulfenia  554 Wulfeniopsis  554 Wullschlaegelia  158 Wunderlichia  603 Wunderlichioideae 602 Wurdastom  355 Wurmbea  145–6 wutong 393 Wyethia  609, 614   Xantheranthemum  565 Xanthisma  609 Xanthium  609 Xanthocephalum  609 Xanthoceras  372–3 Xanthoceratoideae 373 Xanthocercis  253 Xanthocyparis  84 Xanthomyrtus  352 Xanthophyllum  262 Xanthophytum  519 Xanthorhiza  220 Xanthorrhoea  168, 170, 175 Xanthorrhoeoideae 168 Xanthosia  629, 634 Xanthosoma  119 Xanthostemon  352

Xantolis  490 Xantonnea  519 Xantonneopsis  519 Xatardia  634 Xenophyllum  609 Xenoscapa  167 Xenostegia  534 Xeranthemum  609, 613 Xeroaloysia  573 Xerochlamys  399 Xerochloa  209 Xerochrysum  609, 613 Xerocladia  253 Xeroderris  253 Xerodraba  418 Xerolirion  174 Xeronema  168 Xerophyllum  143 Xerophyta  137 Xerorchis  158 Xerosicyos  287 Xerospermum  373 Xerospiraea  264 Xerothamnella  565 Xerotia  448 Xerxes  609 Xestea  522 xhoba 528 Ximenia  422 Ximenioideae 422 Xiphidium  184 Xiphochaeta  609 Xiphotheca  253 Xolocotzia  573 Xylanche  584 Xylanthemum  609 Xylia  253 Xylobium  158 Xylocalyx  584 Xylocarpus  381 Xylococcus  509 Xylomelum  226 Xylonagra  349 Xylonymus  295 Xyloolaena  399 Xylophragma  567 Xylopia  106 Xylopodia  476 Xylorhiza  609 Xyloselinum  634 Xylosma  330 Xylotheca  324 Xymalos  111 Xyridoideae 198 Xyris  198   Yabea  634 yacon 611 Yakirra  209 yala 534 yam  77, 116, 134, 534 yam bean  255 yam daisy  611 yampah 635 yanagi 276 Yanomamua  522 yardlong bean  255 Yariguianthus  609 yarrow  610, 613 Yasunia  112 ya-te-veo 483 year bean  254 Yeatesia  565 yegoma 578 yeheb nuts  255 yellow bark  519

yellow cone flower  614 yellow flamboyant  257 yellow fleabane  612 yellow gentian  522 yellow jessamine  524 yellow loosestrife  494 yellow lupine  257 yellow nutsedge  202 yellow trumpetbush  568 yellow waxbell  474 yellowhorn 374 yellowtop 419 yerba buena  577 yerba del zorillo  452 yerba maté  586, 589 yerba santa  532 Yermo  609 Yersinochloa  208 yesterday, today and tomorrow 540 yew  86, 149 Yinshania  418 ylang-ylang 106 Yoania  158 yohimbine 520 yomogi 611 Youngia  609 Ypsilandra  143 Ypsilopus  158 yuca 335 Yucca  173–4 yuccapalm 174 yuletide camellia  497 yumberry 279 Yunorchis  158 Yushania  208 yuzu 176 Yvesia  209   zabala fruit  217 Zabelia  622–3 Zagrosia  174 Zaleya  458 Zaluzania  609 Zaluzianskya  557 Zameioscirpus  202 Zamia  10–11, 75 Zamioculcas  119–20 Zanha  373 Zannichellia  129 Zanonia  287 Zantedeschia  119–20 Zanthoxylum  375 zǎo  271 Zapoteca  253 Zaqiqah  209 Zataria  577 zauschneria 349 Zea  209–10 zebra plant  565 zebrawood 296 zedoary 192 Zehnderia  311 Zehneria  287 Zelenkoa  158 Zelkova  272 Zeltnera  522 Zemisia  609 zendai 20 Zenia  251 Zenkerella  251 Zenkeria  209 Zenobia  509 Zephyra  164–5 Zephyranthes  171 Zeravschania  634

Zerdana  418 Zeugandra  592 Zeugites  209 Zeuktophyllum  458 Zeuxine  159 Zexmenia  609 Zeyheria  567–8 Zeylanidium  311 zhi mu  174 Zhumeria  577 Zieria  375 Zigadenus  143 Zilla  418 Zingeria  208 Zingiber  191–3 Zingiberales 184 Zingiberoideae 192 Zingiberopsis  191 Zinnia  609, 613 zinnia 614 Zinowiewia  295 Zippelia  98–9 ziricote 531 Zizania  208 Zizaniopsis  208 Zizia  634 Ziziphora  577–8 Ziziphus  271 Zizkaea  195 Zoegea  609 Zollernia  253 Zollingeria  373 Zombia  177 Zonanthus  522 Zonotriche  209 Zootrophion  158 Zornia  253 Zosima  634 Zostera  128 Zoysia  209, 211 Zuccagnia  253, 257 Zuccarinia  519 zucchini 288 Zuckia  451 Zuelania  330 Zuluagocardamum  418 Zygia  253 Zygocarpum  253 Zygochloa  209 Zygogynum  96 Zygopetalum  158 Zygophyllales 246 Zygophylloideae 247 Zygophyllum  247 Zygoruellia  565 Zygosepalum  158 Zygostates  158 Zygostelma  527 Zygostigma  522 Zygotritonia  167 Zyrphelis  609 Zyzyura  609 Zyzyxia  609