The Araceae of Peninsular Malaysia

The Araceae of Peninsular Malaysia

The Araceae of Peninsular Malaysia

Mashhor Mansor Peter C. Boyce Ahmad Sofiman Othman Baharuddin Sulaiman

Penerbit Universiti Sains Malaysia Pulau Pinang

Academic Imprint Series

Distributors: The Universiti Sains Malaysia Co-operative Bookshop Ltd. Universiti Sains Malaysia 11800 USM Pulau Pinang Malaysia MPH Distributors Sdn. Bhd.

© Penerbit Universiti Sains Malaysia, 2012 Perpustakaan Negara Malaysia

Cataloguing-in-Publication Data

The araceae of Peninsular Malaysia / Mashhor Mansor ... [et al.] Bibliography: p. 129.

ISBN 978-983-861-530-3 1. Araceae. 2. Araceae–Ecology. 584.64 QK495.A685

Typeset in Arno Pro Penerbit Universiti Sains Malaysia (Universiti Sains Malaysia Press) 11800 USM, Pulau Pinang, Malaysia Copy Editor: Normah Marmut Cover Designer: Mohd. Murad Shahiran Proofreader: Rosni Habib Typesetter: Sabrina Ahmad Printed by: Sinaran Bros. Sdn. Bhd. 389, Lebuh Chulia 10200 Pulau Pinang Malaysia

I. Mashhor Mansor, 1950-.

Contents Foreword

vii

Introduction

ix

Chapter 1

History of Aroid Research and Fieldwork in Peninsular Malaysia 1 Chapter 2

Ecology and Life Forms

9

Chapter 3

Family Description

17

Chapter 4

The Keys

23

Chapter 5

The Checklist

31

References

129

Index

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Foreword Since 2008 Universiti Sains Malaysia (USM) has been on a journey to be a renowned sustainability-led university under the Accelerated Programme for Excellence (APEX); a publication of this nature is indeed very appropriate and timely in tandem with the mission of the university. This is especially so since Malaysia, hosting one of the world’s mega-diversities, recognizes that good taxonomic outputs are of utmost importance. The Aroids of Peninsular Malaysia will go a long way to fill information gaps in knowledge crucial to keep the rich diversity of Malaysia's flora and fauna intact, and further appreciated by others. It is relevant not only to researchers and university students, but to anyone interested in nature as part of their work or hobby. The authors have taken the challenging task to make this compilation available to the community as part of the USM engagement process. They are known for their passion as field scientists and researchers, spending many months in the forest to make this book possible. This includes travelling and exploring the remote and isolated sites that would otherwise remain hidden from the general public. Their tireless and exemplary work as field botanists to comb the deep Malaysian jungle to make the record available species as complete as possible is praiseworthy. Through this work it may come as a surprise to learn that a large plant family of great ecological and no little economic significance is still much under-researched even for the most basic general account in one of its richest natural habitats, as in Malaysia. Moreso since with the threat of climatic change, and changes in land use many of the species may not be readily available in the near future for us to appreciate, let alone study. Currently taxonomy is in crises, globally, by vii

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Foreword

the ‘trendy’ preoccupation, many times unjustifiable, to focus narrowly on the so-called high-impact-factored journals, making it difficult to persuade researchers, especially the younger ones, to undertake fundamental biological research such as this here. As a result we are fast-losing the ability to generate taxonomic knowledge that is fundamental to the production of good science. This volume is a clear attempt to provide such basic data on aroids in Peninsular Malaysia with the hope that it will both facilitate research, and inspire a new generation of Malaysian students and scientists to further the research done on this fascinating family. I believe given the expertise of the authors on this rarely written subject it would be a very useful reference, in particular, for all plant lovers the world over. Dzulkifli Abdul Razak, Professor Tan Sri Dato’ Vice-Chancellor April 8, 2011

Introduction The Araceae, the fourth largest family of monocotyledons after the orchids, grasses and sedges, is one of the most important mesophytic plant families in tropical Asia. The family has several centres of diversity, notably Borneo, Sumatera, and New Guinea, but it must be said that with the exception of a few well-sampled areas, knowledge of the family is still decidedly incomplete. It is clear from the few studies that have been and are being undertaken that the biology of the family is extraordinarily complex, but that these studies are hampered by the lack of baseline data on the taxonomy for the family. This review by no means aims to be ‘the last word’ – as if that were an even achievable (or desirable) goal. Indeed, a cursory glance will reveal that Homalomena, the largest genus in Peninsular Malaysia, in fact the largest genus in tropical Asia, is by a considerable margin also the least well-known. In short, much remains to be done.

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1

History of Aroid Research and Fieldwork in Peninsular Malaysia

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The first major listing of plants collected from Malaya (which is to say modern Peninsular Malaysia and Singapore) was that of Danish surgeon, Nathaniel Wallich (1786–1854). Wallich came to the then-Danish settlement of Frederiksnagore, near Serampore, Bengal, India in 1807 as a medical attaché. During the Napoleonic War, the British East India Company took over Frederiksnagore, and Wallich was briefly imprisoned. On his release and after spell as an assistant surgeon in the British East India Company's service, Wallich was appointed assistant to William Roxburgh, the East India Company's botanist in Calcutta. In 1817, Wallich was temporarily appointed Superintendent of East India Company's Botanical Garden at Calcutta, a post that was later confirmed. Wallich served until 1846, during which time he travelled extensively on botanical explorations, and compiled a catalogue of more than 20,000 specimens entitled A Numerical List of Dried Specimens of Plants in the East India Company’s Museum – known informally as the ‘Wallich Catalogue’. Wallich’s catalogue includes plants from virtually all of the Asian part of the then British Empire, including not only his own collections from Pinang and Singapore made in 1822, but also specimens collected mainly in 1819 in Pinang by William Jack (1795–1822), and by George Porter of the East India Company’s Garden at Calcutta, who accompanied Wallich to Singapore in 1822 and later (1823–1834) remained in Pinang, at first as a schoolmaster and latterly in charge of the newly founded Botanic Garden near Ayer Hitam. Although the aroids in Wallich’s Catalogue were partly worked up by Roxburgh (1820, 1832) and Jack (1820), in the main it was left to Joseph Dalton Hooker (1817–1911), with considerable assistance from Nicholas Edward Brown (1849–1934), to incorporate them into a modern cohesive account of the aroids for the Flora of British India (see below). Aroids commemorating Wallich include Homalomena wallichii Schott and Schismatoglottis wallichii Hook.f. The next significant Peninsular Malaysian collections of aroids are those of William Griffith (1810–1845). Griffith was a British doctor, naturalist, and botanist initially based in India but later transferred as a civil surgeon to Tenasserim, in southern Myanmar, and subsequently as a civil surgeon in Malacca. Wherever he was stationed or when travelling, Griffith made botanical collections which he either wrote up himself, and were later published posthumously as Notulae ad plantas Asiaticas (Griffith, 1851a) and Icones Plantarum Asiaticarum (Griffith, 1851b), or were written up after his death in the main by Hooker and Brown. Griffith described the genus Cyrtosperma and several novel Homalomena species, including the widespread and variable


colonial helophytic H. rostrata, and Cryptocoryne cordata. Griffith is remembered by Aglaodorum griffithii (Schott) Schott, Cryptocoryne griffithii Schott, and Homalomena griffithii (Schott) Hook.f. The Austrian botanist and gardener Heinrich Wilhelm Schott (1794–1865) was the first monographer of the aroids. He was also the first botanist to make careful comparative studies of aroid inflorescences, flowers and fruits by means of which he was able to put the family on a sound taxonomic footing. He began his outputs, beginning with a series of short notes in which some of his new genera including Anthurium and Philodendron were first published, in the cultural periodical Wiener Zeitschrift für Kunst, Literatur, Theater und Mode as part of a series of articles entitled Für Liebhaber der Botanik (Schott, 1829a, 1829b, 1829c, 1829d, 1829e, 1830). Schott’s most important works with particular impact on Malaysian aroids are accounts of Araceae in the Meletemata botanica (Schott, 1832), the Genera Aroidearum (Schott, 1858), the Synopsis Aroideaum (Schott, 1856), and the Prodromus systematis Aroidearum (Schott, 1860). A notable aspect of Schott’s work was that he used a combination of herbarium material, living plants, and field work in the study of a largely tropical plant group at a time when such a wide-ranging approach was most unusual, and received regular shipments from plant collectors in Malaya and elsewhere in the tropics. Just over ten years following Schott’s death in 1865, Adolf Engler (1844–1930) produced the first papers heralding the start of almost half a century of scientific output of staggering quantity and quality, including two species-level accounts for the Araceae in a little over 20 years (Engler, 1879, 1905, 1908, 1911, 1912, 1915, 1920a, 1920b, 1920c; Engler & Krause, 1908, 1920; Krause, 1908, 1913). Engler worked extensively on Asian aroids, adding much to our knowledge of the family in Malaya. The 1880s marked the beginning of a period intensive general fieldwork in the Peninsula, leading to a great many aroids being described from Peninsular Malaysia based upon types gathered during this era. Collectors that particularly impacted on aroids include the German explorer from Australia, Hermann H. Kunstler (1837–1887), who from 1880–1886 was employed by George King, then director of the Royal Botanic Garden in Calcutta. Kunstler collected chiefly in Perak and his collections are usually annotated as ‘Dr King’s Collector’ on herbarium labels. Contemporaneous to Kunstler, the Italian Rev. Fr Benedetto Scortechini (1845–1886) was, while on a visit in 1884 to Taiping, appointed at the behest of Sir Hugh Low as the Government Botanist for Perak. Scortechini

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collected extensively in the area of Larut, although the paucity of his field data renders his material frustrating to study. Also active was Englishman Leonard Wray ( Jr) (1853–1942), who in 1881 was appointed Superintendent of Larut Hill Garden at Taiping (Perak), and from 1883–1908 was Curator of the Perak State Museum. Between 1905–1908, Wray was Director of Museums of the then Federated Malay States. Alexander Carroll Maingay (1836–1869) graduated in medicine at Edinburgh and in 1859 entered the British Indian Medical Service in Bengal. In 1860 Maingay was transferred to China, before being appointed magistrate in charge of the prison at Malacca from 1862 to 1868. Maingay was later (1868) made superintendent of Rangoon prison, where he was murdered during a mutiny by the prisoners. As with Scortechini, Maingay’s herbarium specimens rarely have much, if any, recorded field data. At approximately the same time in Pinang, Charles Curtis (1852–1928), originally a plant collector for the firm of James Veitch & Sons, occupied the post of superintendent of gardens and forests (1884–1903) including the waterfall gardens in Penang. Curtis collected not only from Pinang but also from Pulau Langkawi, and also in what is now southern Thailand, as well as from the west coast as far as south as Selangor. Curtis retired due to ill health in 1903, after which he built up a herbarium in Penang, which in 1910 was incorporated into that of the Botanic Gardens, Singapore. This remarkable outpouring of effort added numerous novel aroids to the Peninsular Malaysia flora, with several commemorating these extraordinary fieldworkers and their patrons including Pothos curtisii Hook.f., P. kingii Hook.f., Rhaphidophora maingayi Hook.f., Arisaema scortechinii Hook.f., Homalomena scortechinii Hook.f., Scindapsus scortechinii Hook.f., Schismatoglottis scortechinii Hook.f., and Arisaema wrayi Hemsl. During this period of extraordinarily productive botanical exploration, the first comprehensive flora for the Malay Peninsula was being produced in England under Hooker’s Flora of British India project. This eventually extended to seven volumes produced between 1872 and 1897 by Hooker and other botanists working at Kew. The aroids were almost certainly written mainly by Brown, although he gets somewhat less than fulsome acknowledgment in the aroid volume (Hooker, 1893). Brown was a remarkable botanist with a wide range of interests including succulent plants, notably stapeliads and mesembryanthemum, as well as the South African Cape Flora and, as noted, aroids. In a similar vein to Schott, Brown worked on a combination of living and preserved material, and


developed an excellent relationship with the leading tropical plant nurseries. In return he was often sent material to describe. His interest in plants with horticultural potential at time when there was an obsession among the wealthy businessman and titled classes to out-do each other with ever grander collections of ‘stove’ plants, meant that he has been involved with the naming of many of the most beautiful foliage aroids, often in finely illustrated botanical periodicals, including Aglaonema nebulosum N.E.Br., Alocasia reginae N.E.Br., Alocasia reversa N.E.Br. Alocasia sinuata N.E.Br, and Homalomena insignis N.E.Br. Meanwhile, in Calcutta King had embarked on the production of the Materials for a Flora of the Malayan Peninsula, along with by other botanists in Calcutta, primarily David (later Sir David) Prain (1857–1944), a Scottish botanist who studied at the University of Aberdeen. Prain later became curator of the Calcutta herbarium (1887–1889) and was eventually appointed Director of the Royal Botanic Gardens, Kew in 1905. Upon King’s retirement the Materials project was continued by James Sykes Gamble (1847–1925), an English botanist who specialized in the flora of the Indian sub-continent. In the meantime in Singapore, Henry Nicholas Ridley (1855–1956) completed a three-volume account of the monocotyledons as part of the Materials for a Flora of the Malayan Peninsula (Ridley, 1907) and then, when already in retirement and in an extraordinarily swift period of time between 1922 and 1925, produced the five-volume Flora of the Malay Peninsula. Volume 5 of this remains the standard account for the aroids of Peninsular Malaysia (Ridley, 1925). Ridley, while a prodigious generalist, also took a specialist interest in aroids and can be rightly regarded as the first professional botanist based in the Asian tropics to take an active interest in aroids. Curiously, given his impact on Malaysian aroids, Ridley is remembered in only two aroids, Peninsular Malaysian endemic Piptospatha ridleyi N.E.Br. ex Hook.f. and Sarawak endemic Alocasia ridleyi A.Hay. Caetano Xavier Dos Remedios Furtado (1897–1980) joined the staff at Singapore in 1919 as an Assistant Curator in the Gardens Department, Straits Settlements, to work principally on palms and aroids. He took a particular interest in Homalomena, a genus of great taxonomic complexity, and also attempted to unravel the taxonomy of Alocasia macrorrhizos (L.) G.Don. (Furtado, 1935, 1939, 1941). Furtadoa M.Hotta is named for him. Furtado’s work was marked by a tendency to split taxa into infeasibly narrowly defined subordinates. He also had an infuriating habit of changing the name of subordinate taxa when he made changes to their rank, causing a great deal of taxonomic confusion. It is unfortunate that Edred John Henry Corner (1906–1996), unquestionably the

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greatest 20th century botanist of Malaya, shied away from Araceae because of Furtado’s involvement with the family. Notwithstanding that Furtado remained active in Singapore and Peninsular Malaysia well past his retirement, there was a hiatus of field-based activity focussing on aroids until American Dan H. Nicolson undertook extensive aroid collecting through the region during the early 1960s. Nicolson’s studies, in part for his doctoral work on Aglaonema Schott (Nicolson, 1964, 1969), also produced revisions of Amydrium (Nicolson, 1968c), Asian Spathiphyllum (Nicolson, 1968b), and then generic Xenophya (Nicolson, 1968a). Nicolson’s influence extends well beyond the taxonomy of aroids – he is justly regarded by many current aroid botanists as their mentor – while his globally-acknowledged expertise in botanical nomenclature and typification has been of inestimable service in deliberations on the status of early names, and the history of etymological derivations (Nicolson, 1975, 1984, 1987). Several Asian aroids bear Nicolson’s name, including the peninsula endemic Rhaphidophora nicolsonii P.C. Boyce and Sarawak-endemic Schismatoglottis nicolsonii A. Hay. Josef Bogner and Niels Jacobsen began their still ongoing work on the Cryptocoryne, with important outputs including Bogner & Jacobsen (1985, 1986); Jacobsen (1985, 1987); and Jacobsen & Bogner (1986, 1987). Josef Bogner, formerly based in Munich Botanical Garden, Germany, is one of the foremost experts on Araceae, has undertaken research throughout the tropics. In the past 25 years several other botanists have entered the Asian aroid arena, beginning with Alistair Hay in the early 1980s and Peter Boyce in the mid 1980s. Alistair Hay began by tackling the lasioids as part of his doctoral research, and then went on to study and monograph Alocasia (Hay, 1998), Colocasia (Hay, 1996), Schismatoglottis (Hay & Yuzammi, 2000), contributed significantly to studies of Pothos (Boyce & Hay, 2001), the other Schismatoglottideae (Bogner & Hay, 2000), and complied an enormously useful checklist and bibliography for Malesian aroids (Hay et al., 1995). Peter Boyce, while based at Kew Gardens, undertook numerous visits to research and monograph hemiepiphytic aroids in Sunda. He eventually settled permanently in Sarawak in the early 2000s and has since concentrated among other things on the mesophytic aroids of Sarawak (e.g., Boyce, 1996, 1998, 1999, 2001, 2006a, 2006b).


Wilbert Hetterscheid, now based in Wageningen, Netherlands, has spent much of the past 25 years working towards a phylogeny and taxonomic monograph of the large and complex genus Amorphophallus. He has been successful in bringing virtually all species into cultivation, an essential task since herbarium material of these often huge plants is notoriously poorly prepared. To date, Hetterscheid has described almost 80 new species from throughout the range of the genus, including three novelties occurring in Peninsular Malaysia (Hetterscheid, 1994; Hetterscheid & Van der Ham, 2001). Baharuddin Sulaiman appeared in the world of Asian aroids in 1993. Most of his work is related to wetland Araceae, for which he produced a thesis on Riverine Araceae of Peninsular Malaysia in 1997. He also contributed work on wetland Araceae in northern Peninsular Malaysia (Baharuddin, 1997; Mashhor & Sulaiman, 1997; Sulaiman & Mashhor, 2001; Baharuddin & Boyce, 2005, 2010a, 2010b). In late 1997, Sofiman Othman began working on molecular systematics of Cryptocoryne (Othman, 1997). He later went on to study the population genetics of several Cryptocoryne species in the Malay Peninsula which are fast disappearing owing to their habitat being threatened by rapid development (Othman & Sim, 2005; Othman et al., 2007). Together with Jacobsen and Mashhor in 2009, Sofiman wrote an account of Cryptocoryne for the Malay Peninsula (Othman et al., 2009), which is now the standard reference for the genus in West Malaysia. Most recently, a Sarawakian, Wong Sin Yeng has begun undertaking the first critical study of evolution and taxagenesis in Sundaic aroids using a combination of extensive fieldwork and molecular analyses. Her data and those of her students are already changing generic frameworks and forcing re-examination of our understanding of the aroids of Sunda (Boyce & Wong, 2007; Wong & Boyce, 2010; Wong et al., 2010).

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2


Aroids for the greater part are plants dependent on abundant available water and prevailing high atmospheric humidity. Since both structurally and physiologically aroids are not well adapted for growth in arid or cold conditions, none are known occur in the most extreme environments, while those that occur in seasonally cold or arid habitats are for the most part geophytes (see below). Araceae are most diverse and abundant in the humid tropics, and it is there that the richest variety of their life forms are found. Indeed, relatively few genera inhabit temperate regions of the world and as noted these are either geophytes (Ambrosina, Arisaema, Arisarum, Arum, Biarum, Dracunculus, Eminium, Helicodiceros, and Pinellia) or helophytes (Calla, Lysichiton, Orontium, Peltandra, and Symplocarpus). The most detailed reviews yet published for ecology and life forms remain those of Croat (1990, 1992) and were summarized by Mayo et al. (1997).

Ecology The major forest types in Peninsular Malaysia are lowland, hill, and upper hill dipterocarp forest; oak-laurel forest; montane ericaceous forest; peat and peat swamp forest; and mangrove forest. There also lesser areas of freshwater swamp forest, heath forest (kerangas and kerapah), and various types of forest on limestone. Close to Kuala Lumpur there is a forest on quartz ridges (Whitmore, 1975, 1984, 1990, 1998). Up to an altitude of about 900 m asl, forests in Peninsular Malaysia are usually dominated by Dipterocarpaceae, hence the term ‘dipterocarp forests’. Dipterocarp forest in the Peninsula is classified according to altitude, with lowland dipterocarp forest up to 300 m asl, hill dipterocarp forest occurring between 300 m and 750 m asl, and upper hill dipterocarp from 750 m to 1,200 m asl. The last transitions into oak-laurel forest. Lowland dipterocarp forest is a threatened habitat. In the past much of Peninsular Malaysia was covered with lowland forest but today the majority has been cleared for other land uses, with very few areas of this forest type remaining outside of protected areas. Some of the few remaining pockets near urban centres are the Sungai Buloh Reserve, the Kanching Forest Reserve (part of which is the popular Templer's Park), and the Ampang Forest Reserve, in Selangor. Most of the dipterocarp forest remaining in Peninsular Malaysia is categorized as hill dipterocarp forest. The primarily granite Main Range or Banjaran Titiwangsa is the largest remaining continuous forest tract in Peninsular Malaysia, running

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along the backbone of the peninsula, for 500 km southwards from the border of Thailand. In Peninsular Malaysia, limestone forest areas are estimated at 26,000 ha, mostly concentrated in the northern states, and with more than 300 scattered limestone outcrops in the peninsula. A number of these consist of limestone islands in the Langkawi archipelago, with major outcrops in Kelantan, Perlis, Kedah, Perak, and northern Pahang. The outskirts of the city of Kuala Lumpur is surrounded by many greyish outcrops that many assume to be limestone. However, they are not limestone, but rather quartz – essentially glass crystal. Examples include the craggy rocks near Ampang, the Klang Gates Ridge, and the quartz ridge in Kuala Lumpur along the Kajang-Cheras road. These major quartz veins around Kuala Lumpur and Seremban were formed by the Kuala Lumpur-Mersing fault zone, with massive buckling and faulting in the earth's crust thrusting hydrothermal quartz upwards, where they then crystallized.

Aroid Ecology in Peninsular Malaysia Preliminary studies of Araceae biogeography in Peninsular Malaysia have revealed several significant biogeographical ‘realms’, as shown in the Figure 1. The richest habitat for aroids in Malaysia is moist, everwet or perhumid lowland to hill dipterocarp forest, with the greatest abundance and diversity occurring in gallery forest on steep slopes above streams and small rivers. Hay & Yuzammi (2000) speculated that for Schismatoglottis, which are commonly found on steep soil banks in forest, the minute seeds perhaps adhere and the seedlings are less likely to be buried by leaf litter. In addition to this, gallery forest provides a wide range of microhabitats, ranging from cool and constantly humid in the forest directly above the stream, to warmer with diurnal-nocturnal fluctuations in atmospheric humidity on the upper parts of the slope and ridge. The shadier, cooler, wetter, and more consistently humid niches occurring in the lower slope zone, and at the valley bottoms, favour genera such as Schismatoglottis and Homalomena, while the lighter, warmer conditions coupled with fluctuating atmospheric humidity and better drainage provided by the upper slopes favours genera such as Alocasia and Amorphophallus. The ridges themselves, while generally too dry for most terrestrial aroids are frequently host to hemiepiphytic genera such as Pothos, Scindapsus, and Rhaphidophora.


Figure 1 Primary biogeographical realms of Araceae so far identified for Peninsular Malaysia

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Lowland forest on level or gently undulating ground usually carries a less significant species load and also exhibits somewhat of lower diversity than the steep slopes, but nonetheless aroids are still frequently abundant. In particular permanently damp or wet low-lying areas often have Aglaonema and Homalomena, while the juvenile and preadult stages of Rhaphidophora and Scindapsus frequently form tangles of stems. In lowland forest with deep peat deposits (peat forest and peat swamp forest) or white sand and peat deposits (heath forest – kerangas, and its inundated facies – kerapah), there are rather few aroids, although those that do occur are in the main specialized to the highly acidic and low-nutrient soils occurring in these areas. Notable species are Homalomena rostrata, the podzol-specialist Podolasia stipitata, and Rhaphidophora lobbii. Numerous species of the genus Cryptocoryne are a significant component of so-called ‘blackwater’ streams occurring in such habitats. Generally speaking species occurring in blackwater habitats do not occur in other ecologies. Lowland freshwater swamp forest is poor in aroid diversity, although again there are aroids that appear to have become specialized to this habitat, notably Cyrtosperma merkusii and Rhaphidophora minor. Forest above 600 m asl exhibits a lower abundance and diversity of aroids, although certain genera, notably Arisaema, are more diverse and abundant there than at lower altitudes. Still higher altitude forest (above 1,000 m) has few aroids in Malaysia but carries a significant endemic species component from genera that otherwise favour the lowlands. Of particular note in the peninsula are Alocasia perakensis, Rhaphidophora nicolsonii, and Scindapsus scortechinii. Rocky riverbeds frequently have a rheophytic flora component, typically comprising species of Piptospatha and Homalomena of the Chamaecladon Supergroup, while muddy forest stream banks also favour rheophytic species of otherwise non-rheophytic genera, notably Rhaphidophora beccarii. Sandy substrates provide an ideal situation for Cryptocoryne. Disturbed open areas, such as disused padi, especially if seasonally or permanently inundated, often maintain large populations of adventive or naturalized Colocasia esculenta and Alocasia macrorrhizos. The former has been cultivated since antiquity as a carbohydrate crop and minor vegetable, while the latter was perhaps introduced to Malaysia as a medicinal plant in historical times. Another medicinal plant, albeit indigeneous, that is often encountered along the margins of utilized padi is Typhonium flagelliforme (keladi tikus).


Mangrove and the muddy banks of large, open rivers frequently support large stands two species specialist to the habitat – Aglaodorum griffithii and Cryptocoryne ciliata, and also provide a suitable habitat for species associated with freshwater swamp forest. Another important aspect of aroid ecology is the underlying geology. A large area of the peninsula is granite, a rock that is notably impermeable to water, and thus provides water shedding and subsequently a readily draining surface. This habitat, where enough soil overlies the rocks, supports a rich and varied aroid flora. However, in areas where the soil depth is thin or absent, the aroid flora decreases considerably, although rheophytic species and those adapted to lithophytism, are often locally specialized to growing on bare rock. Two geologies in particular show high levels of endemism – limestone and shale. While limestone obligate endemics have long been recognized as being of significant importance, recognition of the importance shale-obligation is new. Forested limestone in Peninsular Malaysia falls into two distinct types. In the southern part of Peninsular Malaysia, depending on aspect and exposure, limestone supports perhumid to moist evergreen forest with a large component of mesophytic aroids, including a number of limestone specialists such as Rhaphidophora burkilliana, R. angustata, and Amydrium humile. By comparison, the limestones in the north, notably those in the northwest, mainly supports dry to humid semi-deciduous forest with a predominance of geophytic taxa, notably Amorphophallus, Typhonium, and Arisaema. Mesophytes, where present, tend to be species of otherwise Thai and Myanmar distribution, for example Aglaonema costatum, A. pumilum, and Homalomena truncata.

Life Forms A large percentage of all Araceae are terrestrial mesophytes occurring in tropical humid forest. In these the stem is aerial and erect or decumbent with short but distinct green internodes; mesophytes are intolerant of atmospheric dryness, lack of root moisture and sun-exposure, and are among the first species to dieout when forests are heavily disturbed. Typical examples are Aglaonema (tribe Aglaonemateae), Homalomena (tribe Homalomeneae), and Schismatoglottis (tribe Schismatoglottideae), all taxa of the Aroideae. Among ‘primitive’ groups only tribe Spathiphylleae and terrestrial Anthurium species have this life form.

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Hemiepiphytes include species from genera of subfamilies Pothoideae and Monsteroideae, as well as many Philodendron, and most species of African tribe Culcasieae. Hemiepiphytes are characteristic plants of humid tropical forests with species varying considerably in size, from shortly climbing plants found on the major branches or trunks of trees to huge plants with attached stems growing high into the forest canopy and oftentimes produce enormously long, pendent flowering stems (e.g., Scindapsus pictus). Hemiepiphytes are divided into primary and secondary hemiepiphytes. Primary hemiepiphytes begin growth above ground level and then go on to but produce feeder roots which eventually grow down to reach the forest floor. Secondary hemiepiphytes germinate on the forest floor, grow up tree boles, become detached from the ground by rotting of the juvenile stem but then become reconnected later by feeder roots which grow down from the upper internodes. Hemiepiphytic aroids typically have anchor roots as well, and are thus often called ‘root climbers’. Flagelliform shoots, heteroblastic leaf development and shingle plants are characteristic features of hemiepiphytic Araceae, though not present in all species of each genus. Epiphytes, which never become connected to the ground by feeder roots, while abundant in the Neotropics, exemplified by Anthurium, Philodendron, and Stenospermation, a rare elsewhere. In the Asian tropics only the genus Remusatia and a few species of Scindapsus are classified as epiphytes. Many hemiepiphytes, epiphytes and geophytes are also found as lithophytes growing on rocks in forested regions wherever shade and humidity are sufficient, the rock surface providing much the same conditions for attachment as tree boles. A number of geophytes are characteristically found growing in the eroded, litter- or humus-filled cavities of limestone outcrops; examples are numerous Amorphophallus, and Typhonium species. Rheophytes (see below) are also typically lithophytic. A life form that is especially well developed in Asian aroids, notably on the island of Borneo, is that of rheophyte. Rheophytes are flood-resistant plants usually from tropical forests, growing in or along swift-running streams inhabiting the region between the lowest and the highest water levels, where plants are repeatedly saturated by regular flash floods after heavy rainfall followed by exposure to dry conditions during the season of low water level. Rheophytes are characterized by narrow, leathery leaves and a firmly attached, usually epilithic stem. In addition to tribe Schismatoglottideae, in which the majority of genera have this habit, rheophytes also occur in Homalomena, Anubias, and Holochlamys.


Aquatic, subaquatic, and helophytic genera are scattered throughout the family from very ‘primitive’ groups such as subfamily Orontioideae to very ‘advanced’ ones such as tribe Cryptocoryneae and Pistia in subfamily Aroideae. Within subfamily Lasioideae genera, e.g., Lasia, Podolasia and many species of Cyrtosperma, have a life form somewhat intermediate between the helophytic/ mesophytic and geophytic/mesophytic categories. Helophytes are widespread throughout the family in many different taxonomic groups of both temperate and tropical genera. The stem may by tuberous (e.g., Typhonium flagelliforme), rhizomatous (Aglaodorum, Homalomena rostrata, H. expedita), semi-prostrate to aerial (Lasia), or erect and arborescent (Montrichardia, Philodendron). Rhizomes are usually adaptations for colonizing muddy riverine margins as in the case of Aglaodorum, Typhonodorum, Homalomena expedita, and Montrichardia. Tuberous stems may be associated with seasonally flooded habitats and a marked dry season – technically such plants are best categorized as hyperseasonal helophytes. Many Cryptocoryne species are either aquatic (permanently in water, the leaves often submerged or held just below the surface) or amphibious (subjected to seasonal drying-out of the habitat, often accompanied by plant dormancy). The inflorescence as a whole or its upper portion is held above the water surface while all other parts are completely submerged. Cryptocoryne is the largest genus of aquatic aroids and merits more detailed consideration. There are a number of species which are usually submerged but are emergent exceptionally low water (e.g., C. affinis). The submerged leaves of such species are relatively large, whereas the emergent leaves are quite small, indicating that such conditions are unfavourable for their growth. The submerged and emergent leaves of the same species generally morphologically different in shape, size, colour and structure. Submerged leaves are softer and emergent ones more coriaceous. Many species occur in the freshwater tidal zone where there is a daily cycle of exposure and submersion. Some species occur only in freshwater, like C. affinis and C. cordata, while others can grow both in fresh and brackish water (e.g., C. ciliata). A few species are helophytes, preferring swampy conditions and growing during the dry season completely emergent in normal soil, like C. spiralis, a weed of rice fields in India. Usually Cryptocoryne species flower at low water level when the plants become emergent. The only free-floating species of Araceae are the pantropical Pistia stratiotes and the five genera of the Lemnoideae (former Lemnaceae – the duckweeds).

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Geophytes cover those genera with tuberous, rhizomatous, subterranean or partly subterranean stems. Geophytic aroids characteristically have periodic dormant periods when no leaves are present and these normally correspond to the dry season (or winter) of their habitat, however, even species from moist and wet tropical forests exhibit growth periodicity and dormancy even in non-seasonal climates (e.g., Amorphophallus and Typhonium). The majority of geophytic genera occur in temperate latitudes, notably in regions with a Mediterranean climate; typical of such genera are the European genera Ambrosina, Arum, Biarum, Dracunculus, and Eminium; warm-temperate South American Gorgonidium, Mangonia, and Synandrospadix; and temperate Asian Arisaema and Pinellia.

Chapter

Family Description

3

18

Family Description

Perennial, evergreen or seasonally dormant bisexual, monoecious, rarely paradioecious, very rarely dioecious, sometimes gigantic, very rarely submicroscopic, often laticiferous or sometimes resiniferous herbs; common life forms (modified from Schimper, 1903; Raunkiaer, 1934) include climbing or suffruticose primary and secondary hemiepiphytes (e.g., Pothos, most Rhaphidophora, etc.), epiphytes (rare in Asia, but including Remusatia and some Scindapsus), nanophanerophytes (e.g., most Aglaonema, most Apoballis), climbing and herbaceous usually mesophytic phanerophytes (e.g., most Homalomena, some Apoballis), mesophytic chamerophytes (e.g., most Schismatoglottis), lithopytes, rheophytes, and chasmophytes (many Schismatoglottis, all Piptospatha), hemicryptophytes (e.g., Hapaline), geophytes (Amorphophallus), inland (freshwater – e.g., some Homalomena, Lasia) or estuary (brackish water, e.g., Aglaodorum) helophytes, amphibious or true hydrophytes (e.g., most Cryptocoryne), hydrohemicryptophytes (Pistia), and hydrotherophytes (Lemnoideae). Stems aerial and erect to climbing or creeping with very short (plant rosulate) to very long (plant scandent) internodes, or subterranean and consisting of a subglobose to depressed-globose tuber (sometimes turnip-or carrot-like or irregular in shape) or horizontal to erect rhizome; shoot modules mono- to multi-phyllous, physiognomically monopodial with sympodia superposed, or very rarely true monopodial flowering terminally with the continuation shoot arising two nodes, rarely one node – Orontioideae below the inflorescence, or apical portion of the shoot not renewing growth after flowering and the plant persisting by the release of lateral buds low on the shoot, the remainder of which subsequently dying; terrestrial plants and helophytes sometimes arborescent with massive stem and terminal rosette of leaves (Alocasia) or arborescent with a pseudostem of petiole sheaths (Arisaema, Typhonodorum); geophytes often with solitary leaf. Leaves usually spirally arranged, sometimes distichous; normally differentiated into petiole and expanded limb, usually glabrous, rarely pubescent, tomentose, villous or with small to large and complex trichomes or papillae on the petiole or leaf blade. Prophylls and cataphylls caducous, marcescent, deciduous or persistent, sometimes beautifully mottled and patterned (e.g., Arisaema, some Alocasia), when persistent sometimes a conspicuous feature of plant and either membranous or forming fibrous mass (e.g., Rhaphidophora korthalsii). Petiole often as long as or longer than lamina, usually smooth, sometimes hairy, papillose, warty, prickly, or aculeate (e.g., Lasioideae, some Amorphophallus), often pulvinate apically (e.g., Scindapsus), basally or rarely centrally (e.g., most Homalomena); petiolar sheath normally well-developed, often at least half as long as entire petiole, sometimes ligulate apically (e.g., Piptospatha), less often very

Family Description

reduced in (e.g., Aglaonema brevispathum and relatives). Leaf blade simple to compound, extremely variable in shape – rarely filiform (e.g., some Cryptocoryne), linear (certain Typhonium), most commonly elliptic, ovate, oblong, sagittate, hastate, less commonly trifid to trisect, pedatifid to pedatisect, radiatisect, ‘dracontioid’ or ‘decompound’ (i.e., trisect with each primary division further much divided), pinnatifid to pinnatisect, bipinnatifid, tripinnatifid to quadripinnatifid (the last two not in Thailand), fenestrate (Rhaphidophora) or laciniate (i.e., fenestrate with slit-like holes – Amydrium medium); heteroblasty frequent, especially in climbing hemiepiphytes, ‘shingle leaves’ sometimes formed (e.g., some Potheae, some Monstereae); venation – midrib almost always differentiated, sometimes massive and succulent; primary veins usually arising pinnately from midrib (and then called primary lateral veins), either running into marginal vein (e.g., Homalomena) or joining distally to form a submarginal collective vein on each side (e.g., Pothos), sometimes primary veins all arising from petiole insertion and running arcuately into leaf apex (e.g., Ariopsis), rarely strictly parallel (not in Malaysia) or subparallel (Pistia), sometimes not differentiated at all (e.g., Epipremnum giganteum); secondary and tertiary veins either reticulate (e.g., Typhonium) or parallel-pinnate, that is running parallel to primaries (e.g., Piptospatha), or arising from primaries at a wide angle and then arching strongly towards leaf margin (e.g., Colocasia), sometimes forming sinuous or zig-zag interprimary collecting veins (‘colocasioid’ venation); higher order venation reticulated or forming cross connections between lower order veins (tessellate). Inflorescence terminal, solitary, or 2 to many in a synflorescence, consisting of a spadix (spike) of small flowers and subtended by a spathe (bract), usually erect, sometimes pendent (e.g., Rhaphidophora hookeri, Piptospatha), sometimes becoming pendent after anthesis (e.g., Schismatoglottis). Peduncle very short to very long, usually similar to petiole in appearance, coloration, pubescence or armature, normally longer than spadix stipe, sometimes ± suppressed and spadix stipe elongated (e.g., some Pothos). Spathe nearly always conspicuous (except some Pothos), very variable in shape and colour, simpler forms often green, reflexed or spreading, more complex forms often showy and highly coloured, erect, usually either boat-shaped or constricted centrally to form a basal tube and an apical limb; tube may enclose the female zone of the spadix or both fertile zones or rarely the entire spadix (e.g., Cryptocoryne), very occasionally much longer than limb (e.g., many Cryptocoryne species), tube margins usually convolute, sometimes connate (e.g., some Typhonium and Amorphophallus); limb usually erect and gaping, sometimes widely spreading, twisted, reflexed or rarely margins ± closed forming slit-like opening; spathe constriction may lie between or above male and female zones or occur in two

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Family Description

places (e.g., some Remusatia species); spathe entirely deciduous soon after anthesis (e.g., most Monstereae), or lower part persistent to fruiting and limb marcescent to deciduous after anthesis (Schismatoglottideae), or spathe entirely persistent until fruiting (e.g., Homalomena) or whole spathe gradually withering and rotting (most Areae). Spadix usually erect, often fleshy and relatively thick, sessile or shortly stipitate, rarely very long-stipitate, usually free, sometimes adnate basally (e.g., Hapaline, Schismatoglottis) or entirely to spathe, either ± uniform in appearance (flowers bisexual), or divided into distinct floral zones (flowers unisexual), fertile zones contiguous or separated by sterile zones, female (pistillate) zone always basal and male (staminate) zone either apical or intermediate in position; sterile zones may be basal, intermediate or apical or any combination of these, apical sterile zone usually known as a terminal appendix; rarely a single plant producing inflorescences bearing male flowers only, followed in later years by inflorescences bearing female flowers only, and vice versa (paradioecy: Arisaema). Flowers 2- to 3-merous, bisexual or unisexual, often very small, protogynous, lacking floral bracts, usually numerous (except e.g., Pistia), almost always sessile, usually densely arranged, sometimes laxly so; bisexual flowers with or without a perigone (perianth), unisexual flowers usually without a perigone but sometimes including rudimentary organs representing modified sexual parts of the other sex (e.g., staminodes of female flowers in Steudnera). Perigone composed of free or partially connate tepals, or consisting of a single cup-like structure; when free, tepals 4 to 6 (–8) and imbricate in 2 whorls, membranaceous or more commonly thickened at least apically, truncate to cucullate. Female flowers (unisexual, aperigoniate flowers) with gynoecium sometimes surrounded by a whorl of variously shaped staminodes (e.g., Steudnera), or sometimes ± regularly associated with a single clavate staminode (e.g., Homalomena). Gynoecium (bisexual and unisexual flowers) with ovary usually 1–3 locular, rarely more, 1-locular ovaries probably always pseudomonomerous; ovules 1-many per locule, orthotropous, hemiorthotropous, campylotropous, amphitropous, hemianatropous or anatropous; placenta 1-several, axile, parietal, apical, basal, or basal and apical; stylar region (tissue lying between ovary and stigmatic epidermis) usually well developed, usually at least as broad as ovary, sometimes attenuate and elongate (e.g., some Amorphophallus, Arisaema) or massive and truncate (most Monstereae species); stigma hemispheric, capitate, discoid, umbonate, more-orless strongly lobed (e.g., some Amorphophallus, Alocasia), rarely stellately lobed (e.g., Ariopsis). Appendix present in some genera (e.g., Thomsonieae, Areae, most Schismatoglottideae), ranging from massively conic to filiform, mostly erect, rarely spreading or pendent, partly or completely covered with staminodes,

Family Description

variously rugose or corrugated or entirely smooth. Sterile organs (pistillodes, staminodes, synandrodes) often forming zones between fertile zones, sometimes present below female zone (Schismatoglottideae), or on base of terminal appendix, very variable in shape, most often ± truncate and prismatic less often filiform, subulate, bristle-like or elongate-clavate, spathulate, cylindric (Aridarum) or enlarged and pearl-like, palmate to otherwise branched; the longer and more complex structures often motile. Stamens of bisexual perigoniate, bisexual non-perigoniate and unisexual perigoniate flowers usually free, equal in number and opposite to tepals (when present); filaments distinct, often ± oblong and flattened, rarely filiform, usually rapidly elongating to push anthers above perigone or gynoecium at anthesis; anthers usually terminal, basifixed, extrorse, composed of two thecae each with two microsporangia; connective usually slender, inconspicuous, often over-topped by thecae, thecae dehiscing by single longitudinal slit or apical stomial pore, with all intermediate degrees occurring. Male flowers (unisexual aperigoniate flowers): 1-8-androus, floral grouping of stamens sometimes obvious in mature inflorescence (e.g., many Philodendron and Homalomena species, sometimes not, e.g., Aglaonema); stamens free or partially to completely connate to form a synandrium. Stamens (unisexual aperigoniate flowers): usually sessile to subsessile, filament sometimes distinct (e.g., Schismatoglottis), connective sometimes ± slender but often strongly thickened, apically broad, fleshy, thecae lying opposite or adjacent on one side of stamen, dehiscing by single longitudinal slit or apical pore, rarely theca prolonged apically into a horn dehiscing by single pore (Cryptocoryneae). Synandrium (unisexual non-perigoniate flowers) usually ± sessile, sometimes formed by fusion of filaments only (e.g., Arisaema), most commonly composed of completely connate stamens and then usually apically truncate and ± prismatic (e.g., Colocasia, Alocasia), very rarely the synandria themselves connate (Ariopsis); common connective (synconnective) usually broad, fleshy (e.g., Alocasia); thecae either lateral, apical or marginal depending on the degree of elongation of the thecae and the extent to which they are overtopped by the synconnective. Infructescence usually densely packed, uncommonly few-fruited or fruits scattered, cylindric to globose, exposed by withering, basal abscission (e.g., Homalomena) or apical splitting (e.g., Alocasia) of spathe, rarely fruits dehiscent, either basally (Lagenandra) or apically (Cryptocoryne) or with seeds exposed by ± simultaneous sloughing of stylar regions of all fruits (‘monsterocarps’ in Monstereae). Fruit mostly a juicy berry, rarely mesocarp leathery, less commonly an apically- (Cryptocoryne) or basally- (Lagenandra) dehiscent carpel, or with the stylar regions sloughing (‘monsterocarp’ in Monstereae); berries normally free, rarely connate (Syngonium), often usually

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Family Description

red, orange, or purplish red, but white (e.g., some Amorphophallus), yellow (some Amorphophallus), green (some Typhonium, Pycnospatha, Lasia), blue (some Amorphophallus), purple (some Typhonium) or brownish (some Typhonium) occurring. Seed 1-many per fruit; testa thick to thin, smooth, roughened, verrucose or striate-costate, papery in seeds with highly developed embryos, sometimes decaying at maturity, or lacking altogether, sometimes arillate with a conspicuous strophiole (e.g., many Areae), rarely operculate (e.g., Pistia); embryo usually straight, sometimes curved, usually undifferentiated, rarely with highly developed plumule (e.g., Cryptocoryne ciliata) and then endosperm lacking and outer cell layers of embryo chlorophyllous; endosperm copious or absent, with all intermediate states occurring. The Araceae is subcosmopolitan in distribution although most abundant and diverse in everwet or perhumid tropics. Globally currently considered to comprise ca. 120 genera and well in excess of 4,000 species, with numerous species awaiting formal description and many more certainly awaiting discovery.

Chapter

The Keys

4

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The Keys

Key to the subfamilies and tribes of Peninsular Malaysian Araceae 1a. Plants comprising small to minute few-rooted or rootless free-floating thallus-like leafless bodies. Lemnoideae – see Generic Key 1 1b. Plants not as above. If free-floating then leaves forming a conspicuous rosette with copious roots hanging beneath.

2a. Flowers bisexual each with a perigone of conspicuous tepals. 3a. Spiny helophytes.

Lasioideae – see Generic Key 2

3b. Climbing hemiepiphytes with (mostly) elliptic leaves. Pothoideae – Pothos

2b. Flowers bisexual without a perigone of conspicuous tepals, or unisexual. 4a. Flowers bisexual, perigone if present membranous and inconspicuous; mainly climbing hemiepiphytes, rarely rheophytic. Monsteroideae – see Generic Key 3 4b. Flowers unisexual. 5a. Free-floating plants with leaves forming a conspicuous rosette with Aroideae: Pistieae – Pistia copious roots hanging beneath. 5b. Plants not so. 6a. Leaves variously divided. 7a. Leaf blade decompound. Inflorescences appearing before leaf emergence, with flowers of both sexes. Aroideae: Thomsonieae – Amorphophallus 7b. Leaf blade trifoliolate to pentafoliolate. Inflorescences produced with leaves, often single-sexed. Aroideae: Arisaemateae – Arisaema

The Keys

6b. Leaf blade simple, ranging from linear-lanceolate to hastate. 8a. Aquatic plants. Infructescences composed of dehiscent carpels. Aroideae: Cryptocoryneae – Cryptocoryne 8b. Plants not aquatic, rarely colonial Infructescences comprised of Berries.

helophytes.

9a. Plants mostly suffruticose. Fruits conspicuous red or pink berries not surrounded by a persistent spathe. Aroideae: Aglaonemateae – see Generic Key 4 9b. Plants not suffruticose. Fruits various, if red or orange red berries then surrounded by a persistent lower spathe. 10a. Entire spathe closing after anthesis and then persisting until fruit maturity. Aroideae: Homalomeneae – see Generic Key 5 10b. Spathe limb mostly shedding during or soon after anthesis, and lower spathe persisting until fruit maturity; if upper spathe persisting then attached portion of petiolar sheath very short and the remainder free-ligular or absent and the protective role taken on by prophyll/cataphyll. 12a. Staminate flowers forming synandria. 13a. Spathe not obviously with a lower tubular part separated from the spathe limb by a conspicuous constriction. Spathe limb persistent. Pistillate flowers very few, arranged in a single vertical row. Fruits ripening white, enclosed in the persistent folded lower spathe but not exposed by the lower spathe actively splitting. Aroideae: Caladieae – Hapaline

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The Keys

13b. Spathe with a well-defined lower tubular part separated from the spathe limb by a pronounced constriction and forming a chamber enclosing all or most of the pistillate flower zone. Spathe limb deciduous prior to fruit maturation. Pistillate flowers many, arranged in a dense spiral. Fruits ripening red and odourless or yellow-brown odorous, exposed by the persistent lower spathe actively splitting from top to bottom. Aroideae: Colocasieae – see Generic Key 6 12b. Staminate flowers not forming synandria 14a. Tuberous-stemmed geophytes. Sterile flowers between the staminate and pistillate flower zones filamentous and tangled. Spadix with a long, naked appendix. Aroideae: Areae – Typhonium 14b. Mesophytes, rheophytes, lithophytes and chasmophytes with stems never tuberous. Sterile flowers between the staminate and pistillate flower zones if present never filamentous and tangled. Spadix appendix, if present, never naked. Aroideae: Schismatoglottideae – see Generic Key 7

The Keys

Keys to the genera of Peninsular Malaysian Araceae Generic Key 1 – Lemnoideae

1a. Roots 1–21 per plant body; plant body with 1–21 veins; the daughter plant bodies and inflorescences originating from 2 lateral pouches at the base of the plant body. 2a. Roots 1 per plant body; plant bodies with 1–3 veins, without a scale at the base Lemna. 2b. Roots (1–)2–21 per plant body; plant bodies with (3–)5–21 veins, surrounded at its base by a small scale (prophyll) covering the point of attachment of the roots. 3a. Plant bodies 1–1.5 times as long as wide with 10–21 veins and 7–12 Spirodela roots of which 1(–2) perforate the scale. 3b. Plant bodies 1.5–2 times as long as wide, with (3–)5–7 veins and Landoltia (1–)2–7( –12) roots all of which perforate the scale. 1b. Roots absent; plant body without veins; the daughter plant bodies riginating from a single terminal pouch or cavity at the base of the plant body; inflorescence originating in a cavity on the plant body upper surface. Wolffia Generic Key 2 – Lasioideae

1a. Stems suffruticose, erect to decumbent, usually with prickly conspicuous internodes; leaf blade hastate to pinnatifid; spathe caducous, rarely marcescent; placentation apical; fruit usually spinulous. Lasia 1b. Stem a condensed rhizome, rarely with distinct internodes, and these then unarmed; leaves entire, sagittate to hastate; spathe persistent to marcescent; placentation not apical; fruit smooth. 2a. Spathe interior white, infructescence nodding, fruits barely emerging from between the tepals, ripening dull purple. Seeds crested or warty. Plant with spines mixed straight and up-turned. Plants mainly of freshwater swamp forest. Cyrtosperma

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The Keys

2b. Spathe interior red-purple, infructescence erect, fruits emerging fully from between the tepals, ripening bright red. Seeds smooth. Plant with spines mixed straight and down-turned. Plants of peat and peatswamp Podolasia forest. Generic Key 3 – Monsteroideae

1a. Spathe in bud stout, short to long-stout-pedunculate, not conspicuously long-beaked, if beak present then less than 1/6 length of entire spathe, opening with inside yellow, orange, greenish or white, only moderately waxy. Flowers naked. Infructescence a monsterocarp or, if with discrete berries, then these not truncate, and ripening white. Trichosclereids present (but sparse in Amydrium). 2a. Adult leaf blade pinnate and conspicuously perforated. Infructescence with discrete berries ripening white. Trichosclereids sparse. Amydrium 2b. Adult leaf blade entire, sometimes conspicuously perforated; or pinnate and then ‘not’ conspicuously perforated. Infructescence a monsterocarp, with stylar plates sloughing away to reveal seeds. Trichosclereids abundant. 3a. Ovules 2–6 or more, placentation parietal. Fruits with more than one seed. 4a. Ovules 8 or more, superposed on 2 (rarely 3) parietal placentas. Seeds many, ellipsoid, straight, 1.3–3.2 mm long, 0.6–1 mm wide; testa brittle, smooth. Rhaphidophora 4b. Ovules 2–4(–6) at base of a single intrusive placenta. Seeds few, curved, 3–7 mm long, 1.5–4 mm wide; testa bony and ornamented. Epipremnum 3b. Ovules solitary, placentation basal. Fruits with a solitary seed. Scindapsus

The Keys

1b. Spathe in bud slender, long-slender-pedunculate, conspicuously long-beaked (beak to 1/3 length of entire spathe), opening with inside greenish white or white, and conspicuously glossy-waxy. Flowers with a membranous perigone of fused tepals. Infructescence with discrete, truncate berries ripening red or Anadendrum orange. Trichosclereids absent. Generic Key 4 – Aglaonemateae

1a. Colonial helophytes with creeping spongy rhizomes and erect, spongy petioles. Fruits large, ripening green and functioning as floating dispersal units. Aglaodorum 1b. Terrestrial forest herbs mainly with erect firm stems, and erect solid petioles. Fruits medium-sized, ripening red or rarely pink, not water dispersed. Aglaonema Generic Key 5 – Homalomeneae

1a. Staminate flower consisting of solitary stamen overtopped by flask-shaped pistillode; ovary 1-locular, placenta basal. Furtadoa 1b. Staminate flower consisting of (1)2–6 stamens, pistillodes absent; ovary incompletely 2–5 locular, placentas parietal and axile. Homalomena Generic Key 6 – Colocasieae

1a. Infructescences erect, fruits medium-sized, ripening orange-red, odourless. Alocasia 1b. Infructescences pendent, fruits small, ripening yellowish brown, smelling variously of overripe fruit or butyric acid. Colocasia Generic Key 7 – Schismatoglottideae

1a. Wings of petiolar sheath always extended into a free ligular portion; seeds with micropylar appendage. Fruiting spathe forming a splashcup. Plants obligate rheophytes. Piptospatha

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The Keys

1b. Wings of petiolar sheath fully or almost fully attached to the petiole; seed without a micropylar appendage. Fruiting spathe not forming a splash-cup. 2a. Inflorescences on very slender peduncles, nodding at anthesis, peduncle at spathe insertion flexing 180O from vertical axis. Infructescences narrowly campanulate,nodding. Plants of podsols. Hestia 2b. Inflorescences erect at anthesis, peduncle stout. Infructescences fusiform with a constricted orifice, if campanulate, then thick-walled and erect, never nodding. Plants of various substrates but never on podsols. Schismatoglottis

Chapter

The Checklist

31

5

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The Checklist

Aglaodorum Schott Aglaodorum Schott, Gen. Aroid.: 58 (1858). (Plate 1) Taxonomy: One species occurring from southern Cambodia and Vietnam to Peninsular Malaysia, Sumatera, and much North Borneo, although absent from Sabah. Most recent account: Nicolson (1969). Ecology: Colonial helophyte in tidal open swamps. Notes: A single species closely allied to Aglaonema, Aglaodorum griffithii (Schott) Schott differs from Aglaonema in vegetative and fruit features seemingly associated with a swamp-dwelling habitat. Typically Aglaodorum occurs as a colonial helophyte in open swamps, especially on mud in freshwater and brackish tidal zones, usually in full sun. The stems and petioles are filled with air-spaces. The fruits, with very thick flesh, are green and evidently dispersed by floating. Unlike Aglaonema the petioles and peduncles are relatively long, elevating the leaf blades and inflorescences above water. Aglaodorum griffithii (Schott) Schott, Gen. Aroid.: 58 (1858). Aglaonema griffithii Schott, Syn. Aroid.: 123 (1856). Aglaonema palustre Teijsm. & Binn., Natuurw. Tijdschr. Ned.-Indië 25: 399 (1863). Distribution: South Vietnam and Cambodia to Peninsular Malaysia, Sumatera, and North West Borneo.

Plate 1. Aglaodorum Schott. A–E. Aglaodorum griffithii (Schott) Schott. A. Plants in habitat. B. Inflorescence at pistillate anthesis. C. Ripe infructescence. The large green fruits function as floating dispersal bodies. D. Germinating seed. Note that the fruit flesh has split to allow the emergence initially of the several hooked cataphylls. Only later does the root-radical emerge.

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The Checklist

Aglaonema Schott Aglaonema Schott, Wiener Z. Kunst 1829: 892 (1829). (Plate 2) Taxonomy: Twenty-two species in tropical and subtropical Asia from North East India through to New Guinea. Four species in Peninsular Malaysia, one endemic. Most recent account: Nicolson (1969). Ecology: Primary and secondary perhumid to everwet, occasionally swampy, broadleaf evergreen forest. Aglaonema costatum N.E.Br., Gard. Chron., III, 1892: 426 (1892). Aglaonema costatum var. foxii Engl., Pflanzenr., IV, 23Dc: 17 (1915). Aglaonema costatum f. foxii (Engl.) Jervis, Aglaonema: 25 (1980). Aglaonema costatum var. maculatum Engl., Pflanzenr., IV, 23Dc: 17 (1915). Distribution: Thailand, South Vietnam, and Peninsular Malaysia (Pulau Langkawi). Aglaonema flemingianum A. Hay, Gard. Bull. Singapore 50: 1 (1998). Distribution: Peninsular Malaysia (Terengganu). Endemic. Aglaonema nebulosum N.E.Br., Ill. Hort. 31: 67 (1887). Aglaonema nanum Hook.f., Fl. Brit. India 6: 530 (1893). Aglaonema minus var. nanum (Hook.f.) Ridl., Mat. Fl. Malay. Penins. 2: 22 (1907). Aglaonema pictum var. nanum (Hook.f.) Ridl., Fl. Malay Penins. 5: 102 (1925). Aglaonema nebulosum f. nanum (Hook.f.) Nicolson, Smithsonian Contr. Bot. 1: 31 (1969). Aglaonema minus Hook.f., Fl. Brit. India 6: 530 (1893). Aglaonema scortechinii Hook.f., Fl. Brit. India 6: 530 (1893). Aglaonema minus var. scortechinii (Hook.f.) Ridl., Mat. Fl. Malay. Penins. 2: 22 (1907). Aglaonema pictum var. scortechinii (Hook.f.) Ridl., Fl. Malay Penins. 5: 102 (1925). Aglaonema obovatum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 322 (1922).

Plate 2. Aglaonema Schott. A–B. Aglaonema nitidum ( Jack) Kunth. A. Adult fruiting plant (over 1 m tall). In some populations, as here, the fruits are ripe when pale pink, whereas mainly fruits ripen through to scarlet. B. Inflorescence at pistillate anthesis. C–E. Aglaonema nebulosum N.E.Br. C. Fruiting plant in habitat. Aglaonema nebulosum is restricted to kerangas. D. Inflorescence at pistillate anthesis. Note the globose staminate portion greatly exceeds the spathe; this occurs at the onset of pistillate anthesis and is enabled by a rapid extension of the spadix stipe. E. Inflorescence at pistillate anthesis with the spathe artificially removed. Note the much-extended stipe.

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Distribution: Sumatera, Peninsular Malaysia, and North Borneo Aglaonema nitidum ( Jack) Kunth, Enum. Pl. 3: 56 (1841). Calla nitida Jack, Malayan Misc. 1(1): 24 (1820). Arum integrifolium Link, Enum. Pl. 2: 394 (1822). Aglaonema integrifolium (Link) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 20 (1832). Aglaonema oblongifolium Schott, Wiener Z. Kunst 1829: 829 (1829). Aglaonema princeps Kunth, Enum. Pl. 3: 55 (1841). Aglaonema oblongifolium var. curtisii N.E. Br., Gard. Chron., III, 21: 70 (1897).

Aglaonema nitidum f. curtisii (N.E.Br.) Nicolson, Baileya 15: 126 (1968).

Aglaonema oblanceolatum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 322 (1922). Aglaonema oblanceolatum f. maximum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 323 (1922). Aglaonema nitidum f. cinereum Jervis, Aglaonema: 31 (1980). Distribution: Thailand to Indo-China, Sumatera, Peninsular Malaysia, Java, and Borneo. Aglaonema simplex (Blume) Blume, Rumphia 1: 152 (1837). Caladium simplex Blume, Catalogus: 103 (1823). Aglaonema simplex f. typicum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 323 (1922), nom. inval. Caladium princeps Kunth, Enum. Pl. 3: 55 (1841). Aglaonema schottianum Miq., Fl. Ned. Ind. 3: 216 (1856). Aglaonema schottianum var. genuinum Engl., Pflanzenr., IV, 23Dc: 20 (1915), nom. inval. Aglaonema malaccense Schott, Bonplandia (Hannover) 7: 30 (1859). Aglaonema schottianum var. malaccense (Schott) Engl., Pflanzenr., IV, 23Dc: 21 (1915). Aglaonema longicuspidatum Schott, Prodr. Syst. Aroid.: 304 (1860).

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Aglaonema propinquum Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 280 (1864). Aglaonema fallax Schott ex Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 439 (1879). Aglaonema birmanicum Hook.f., Fl. Brit. India 6: 529 (1893). Aglaonema nicobaricum Hook.f., Fl. Brit. India 6: 530 (1893). Aglaonema simplex f. nicobaricum (Hook.f.) Jervis, Aglaonema: 34 (1980). Aglaonema angustifolium N.E.Br., Bull. Misc. Inform. Kew 1895: 18 (1895). Aglaonema schottianum f. angustifolium (N.E.Br.) Engl., Pflanzenr., IV, 23Dc: 20 (1915). Aglaonema simplex f. angustifolium (N.E.Br.) Jervis, Aglaonema: 34 (1980). Aglaonema tenuipes Engl., Bot. Tidsskr. 24: 275 (1902). Aglaonema siamense Engl., Bot. Tidsskr. 24: 275 (1902). Aglaonema angustifolium var. undulatum Ridl., Mat. Fl. Malay. Penins.: 21 (1907). Aglaonema borneense Engl., Pflanzenr., IV, 23Dc: 22 (1915). Aglaonema schottianum var. brownii Engl., Pflanzenr., IV, 23Dc: 21 (1915). Aglaonema schottianum var. winkleri Engl., Pflanzenr., IV, 23Dc: 21 (1915). Aglaonema subfalcatum Engl., Pflanzenr., IV, 23Dc: 23 (1915). Aglaonema pierreanum Engl., Pflanzenr., IV, 23Dc: 24 (1915). Aglaonema brevivaginatum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 324 (1922). Aglaonema elongatum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 326 (1922). Aglaonema emarginatum Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 324 (1922). Aglaonema grande Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 325 (1922). Aglaonema latius Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 325 (1922).

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Aglaonema nieuwenhuisii Engl. ex Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 320 (1922). Aglaonema simplex f. nieuwenhuisii (Engl. ex Alderw.) Jervis, Aglaonema: 34 (1980). Aglaonema simplex f. inaequale Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 323 (1922). Aglaonema subarborescens Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 326 (1922). Scindapsus tonkinensis K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem 11: 331 (1932). Aglaonema alpinum Elmer, Leafl. Philipp. Bot. 10: 3611 (1938), no latin descr. Distribution: Nicobar Islands through to West Maluku. Alocasia (Schott) G.Don Alocasia (Schott) G.Don in R. Sweet, Hort. Brit., ed. 3: 631 (1839), nom. cons. (Plate 3) Ensolenanthe Schott, Bonplandia (Hannover) 9: 368 (1861). Schizocasia Schott, Bonplandia (Hannover) 10: 148 (1862). Xenophya Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 124 (1863). Panzhuyuia Z.Y. Zhu, J. Sichuan Chinese Med. School 4(5): 49 (1985). Taxonomy: At least 100 species distributed from the subtropical eastern Himalayas throughout subtropical and tropical Asia into the tropical western pacific and eastern Australia. Four species in Peninsular Malaysia, plus one naturalized non-indigenous. None endemic (compared with Borneo, a major centre of diversity, with 41 indigenous species, of which 37 endemic). Most recent account: Hay (1998). Ecology: Primary and secondary perhumid to everwet broadleaf forest and open swamps; some species are tolerant of quite strong seasonality. Predominantly in

Plate 3. Alocasia (Schott) G.Don. A–E. Alocasia longiloba Miq. A & B. Variation in leaf markings and leaf blade shape. Alocasia longiloba is a highly variable species, notable for producing localized populations with highly distinctive morphologies but which, when viewed as part of the species as a whole co-exist with a range of intermediates. C. Inflorescences emerge from subtended by conspicuous prophylls and cataphylls in many populations (as here on Santubong, North West Sarawak). D. Inflorescence at pistillate anthesis with spathe partly removed. Note that constriction between the lower spathe and limb coincides with the sterile interstice separating the pistillate and staminate flower zones. Note, too, that the staminate flower zone is entirely outside the lower spathe. The conspicuously stellate stigma lobes are a feature of the Longiloba Group. E. Ripe infructescence. Note that the lower spathe has been shed some time before to leave a scar below the fruits. This is in contrast to other Alocasia in which the lower spathe persists until fruit maturity, at which point it actively splits downwards into strips that recurve to reveal the fruits.

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the lowlands, extending from sea level to lower and mid-montane zones. Many species are specific to certain geologies, notably limestone. Distinguishing Alocasia from Colocasia Traditionally, these two genera which are undoubtedly closely allied and frequently confused with one another, have been separated on the basis of ovule number and placentation – many ovules on parietal placentas in Colocasia, few on basal placentas in Alocasia (e.g., Mayo et al., 1997: 90). These characteristics are not really of practical use in field identification. However they translate in fruiting plants into markedly different dispersal syndromes, apparently (though not observed in West Malesia) involving birds in Alocasia, in marked contrast to the mammal dispersal syndrome of Colocasia where the fruits are smelly and inconspicuously coloured with many tiny seeds in slimy mucilage (Hay, 1996). In respect of synflorescence architecture, Alocasia may be readily distinguished from Colocasia by its bimodular synflorescence subunits. Inflorescence multiplication in Colocasia is achieved in such a way that the whole synflorescence is equivalent to one bimodular unit in Alocasia. Where the inflorescence terminating the vegetative module has only one further inflorescence in the axil of its subtending cataphyll in Alocasia (with the synflorescence being built up by relay axes), in Colocasia the second inflorescence has a third in the axil of its prophyll and so on up to c. 8 in Colocasia gigantea. The relay axis in Colocasia is vegetative and thus the whole synflorescence is displaced to a quasi-lateral position on one side of the shoot. Alocasia inornata Hallier f., Meded. Rijks-Herb. 26: 7 (1915). Alocasia nobilis Hallier f., Meded. Rijks-Herb. 26: 6 (1915). Distribution: Sumatera and southern Peninsular Malaysia. Alocasia longiloba Miq., Fl. Ned. Ind. 3: 207 (1856). Caladium veitchii Lindl., Gard. Chron. 1859: 740 (1859). Alocasia veitchii (Lindl.) Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 125 (1863). Alocasia lowii var. veitchii (Lindl.) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 508 (1879). Alocasia korthalsii Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 124 (1863).

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Alocasia lowii Hook.f., Bot. Mag. 89: t. 5376 (1863). Caladium lowii Lem., Ill. Hort. 1863: t. 360 (1863). Alocasia lowii var. picta Hook.f., Bot. Mag. 91: t. 5497 (1865). Alocasia singaporensis Linden, Gartenflora 14: 252 (1865). Alocasia amabilis W. Bull, Cat. 143: 9 (1878). Alocasia thibantiana Mast., Gard. Chron., n.s., 1878(1): 527 (1878). Alocasia denudata Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2:507 (1879). Alocasia putzeysii N.E.Br., Ill. Hort. 29: t. 439 (1882). Alocasia eminens N.E.Br., Gard. Chron., III, 1887(1): 105 (1887). Alocasia lucianii Pucci ex Rodigas, Ill. Hort. 34: 75 (1887). Alocasia pucciana André, Rev. Hort. 1887: 466 (1887). Alocasia curtisii N.E.Br., Bull. Misc. Inform. Kew 1894: 247 (1894).

Alocasia cuspidata Engl., Bot. Jahrb. Syst. 25: 25 (1898).

Alocasia cochinchensis Pierre ex Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23E: 103 (1920). Alocasia denudata var. elongata Engl., Pflanzenr., IV, 23E: 100 (1920). Alocasia leoniae Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23E: 109 (1920), prov. syn. Alocasia longifolia Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23E: 103 (1920), prov. syn. Alocasia spectabilis Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23E: 109 (1920), prov. syn. Distribution: China (Yunnan to Guangdong) to Indo-China and Thailand, Peninsular Malaysia, through to C. Malesia.

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Alocasia macrorrhizos (L.) G.Don in R. Sweet, Hort. Brit., ed. 3: 631 (1839). Arum macrorrhizon L., Sp. Pl.: 965 (1753). Caladium macrorrhizon (L.) R.Br., Prodr.: 336 (1810). Colocasia macrorrhizos (L.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 18 (1832). Arum peregrinum L., Sp. Pl.: 966 (1753). Colocasia peregrina (L.) Raf., Fl. Tellur. 3: 65 (1837). Philodendron peregrinum (L.) Kunth, Enum. Pl. 3: 51 (1841). Arum mucronatum Lam., Encycl. 3: 12 (1789). Colocasia mucronata (Lam.) Kunth, Enum. Pl. 3: 40 (1841). Arum indicum Lour., Fl. Cochinch.: 536 (1790). Colocasia indica (Lour.) Kunth, Enum. Pl. 3: 39 (1841). Colocasia indica (Lour.) Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 160 (1842). Alocasia indica (Lour.) Spach, Hist. Nat. Vég. 12: 47 (1846). Alocasia indica var. typica Engl., Pflanzenr., IV, 23E: 87 (1920). Arum cordifolium Bory, Voy. îles Afrique 1: 376 (1804). Alocasia cordifolia (Bory) Cordem., Fl. Réunion: 136 (1895). Caladium odoratum Lodd., Bot. Cab. 5: t. 416 (1820). Arum rapiforme Roxb., Fl. Ind. ed. 1832, 3: 497 (1832). Colocasia rapiformis (Roxb.) Kunth, Enum. Pl. 3: 40 (1841). Alocasia rapiformis (Roxb.) Schott, Prodr. Syst. Aroid.: 157 (1860). Calla badian Blanco, Fl. Filip.: 658 (1837). Calla maxima Blanco, Fl. Filip.: 658 (1837). Colocasia indica var. rubra Hassk., Pl. Jav. Rar.: 145 (1840). Alocasia indica var. rubra (Hassk.) Engl., Pflanzenr., IV, 23E: 88 (1920). Colocasia boryi Kunth, Enum. Pl. 3: 41 (1841). Philodendron punctatum Kunth, Enum. Pl. 3: 48 (1841). Alocasia metallica Schott, Oesterr. Bot. Wochenbl. 4: 410 (1854). Alocasia indica var. metallica (Schott) Schott, Prodr. Syst. Aroid.: 145 (1860). Alocasia pallida K. Koch & C.D. Bouché, Index Seminum (B) 1854(App.): 5 (1855).

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Alocasia variegata K. Koch & C.D. Bouché, Index Seminum (B) 1854(App.): 5 (1855). Alocasia indica var. variegata (K. Koch & C.D. Bouché) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 502 (1879). Caladium indicum K. Koch, Berliner Allg. Gartenzeitung 25: 136 (1857), nom. inval. Caladium plumbeum K. Koch, Berliner Allg. Gartenzeitung 25: 136 (1857), nom. inval. Alocasia plumbea Van Houtte, Fl. Serres Jard. Paris, II, 11: 93 (1875). Alocasia indica var. heterophylla Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 502 (1879). Caladium metallicum Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 502 (1879). Alocasia grandis N.E.Br., Gard. Chron., n.s., 1886(2): 890 (1886), nom. illeg. Alocasia marginata N.E.Br., Gard. Chron., III, 1887(2): 712 (1887). Alocasia indica var. diversifolia Engl., Pflanzenr., IV, 23E: 88 (1920). Alocasia uhinkii Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23E: 110 (1920). Distribution: Malesia through to Queensland (Murray Group). Not indigenous in Peninsular Malaysia. Notes: Alocasia macrorrhizos is never encountered in the wild but rather is always closely associated with human settlement. Alocasia perakensis Hemsl., J. Bot. 25: 205 (1887). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Alocasia puber (Hassk.) Schott, Syn. Aroid.: 47 (1856). Colocasia pubera Hassk., Cat. Hort. Bot. Bogor.: 302 (1844). Alocasia margaritae L. Linden & Rodigas, Ill. Hort. 33: 155 (1886). Alocasia ovalifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 47 (1904).

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Alocasia crassinervia Engl., Pflanzenr., IV, 23E: 82 (1920). Distribution: Peninsular Malaysia, West and central Java. Amorphophallus Blume ex Decne. Amorphophallus Blume ex Decne., Nouv. Ann. Mus. Hist. Nat. 3: 366 (1834), nom. cons. (Plates 4 & 5) Thomsonia Wall., Pl. Asiat. Rar. 1: 83 (1830), nom. rej. Pythonium Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 17 (1832), nom. illeg. Pythion Mart., Flora 14: 459 (1831), nom. rej. Candarum Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 17 (1832), nom. illeg. Kunda Raf., Fl. Tellur. 2: 82 (1837). Brachyspatha Schott, Syn. Aroid.: 35 (1856). Conophallus Schott, Syn. Aroid.: 34 (1856). Plesmonium Schott, Syn. Aroid.: 34 (1856). Corynophallus Schott, Oesterr. Bot. Wochenbl. 7: 389 (1857). Allopythion Schott, Gen. Aroid.: t. 24 (1858). Hansalia Schott, Oesterr. Bot. Z. 8: 82 (1858). Hydrosme Schott, Gen. Aroid.: t. 33 (1858). Rhaphiophallus Schott, Gen. Aroid.: t. 27 (1858). Synantherias Schott, Gen. Aroid.: t. 28 (1858). Dunalia Montrouz., Actes Soc. Linn. Bordeaux 26: 576 (1866). Proteinophallus Hook.f., Bot. Mag. 101: t. 6195 (1875). Tapeinophallus Baill., Dict. Bot. 1(5) (1877). Pseudodracontium N.E.Br., J. Bot. 20: 193 (1881).

Plate 4. Amorphophallus Blume ex Decne. A. Amorphophallus haematospadix Hook.f. cannot easily be mistaken for any other species. The red, clavate appendix which produces a faint but distinct banana-like scent from its upper half. B. Amorphophallus carneus Ridl. resembles A. excentricus (Plate 5A) from southern Thailand and Pulau Langkawi, the latter differing mainly in having tubers smaller offsets and a larger, especially broader appendix with more prominent grooves, often running the entire length or with a network of shallow grooves. C. Amorphophallus bufo Ridl. is a common highland species in the Main Range, and quite variable in colour. D. Formerly considered a Sumateran endemic, Amorphophallus manta Hett. & Ittenbach is now considered to be represented in Peninsular Malaysia. Lowland plants hitherto referred to as A. bufo are referable here. A-D © Wilbert Hetterscheid. Used with permission.

Plate 5. Amorphophallus Blume ex Decne. A. Amorphophallus excentricus Hett. is a recently described species related to A. carneus. B–C. Amorphophallus elatus Hook.f. Amorphophallus elatus may be looked upon as a small version of A. macrorrhizos, a species from northern Thailand. It differs by its smaller dimensions, lacking hairs on the appendix, the strongly 2-lobed stigma, and lacking hairs on the petiole and peduncle. D–E. Amorphophallus boyceanus Hett. resembles most closely A. elegans, but the latter has a much shorter and straight style, and a broadly triangular spathe. Javan A. variabilis is also very similar, but has a distinctly narrowed base to the spathe limb, and many more and smaller annual offsets, while the appendix usually is three times or more the length of the fertile part of the spadix. F. Amorphophallus elegans Rid. resembles A. variabilis but differs by having the spathe limb not suddenly narrowed, the spathe base interior not verrucate but smooth or furrowed, a non-lobate stigma, and leaves with unwinged rhachises with some of the leaflets petiolulate. A–F © Wilbert Hetterscheid. Used with permission.

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Taxonomy: Over 200 species distributed from tropical Africa and Madagascar, throughout the Indian subcontinent, the subtropical eastern Himalayas, subtropical and tropical Asia into western Oceania and northeastern Australia. 12 or 13 indigenous species in Peninsular Malaysia, of which four are endemic. One species (A. paeoniifolius (Dennst.) Nicolson) almost certainly non-indigenous and found only in association with human disturbance. Most recent account: Ridley (1925). Ecology: Lowland to upper hill forest perhumid to everwet broadleaf forest. Some of the peninsular species are limestone obligates. Amorphophallus boyceanus Hett., Blumea 46(2): 256 (2001). Distribution: Southern Peninsular Thailand to Peninsular Malaysia (the latter not confirmed; based on a visual record). Amorphophallus bufo Ridl., J. Fed. Malay States Mus. 4: 89 (1909). Distribution: Peninsular Malaysia. Endemic. Amorphophallus carneus Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 47 (1903). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Amorphophallus elatus Hook.f., Fl. Brit. India 6: 517 (1893). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Amorphophallus elegans Ridl., J. Straits Branch Roy. Asiat. Soc. 86: 310 (1922). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Amorphophallus excentricus Hett., Blumea 39: 255 (1994). Distribution: Southern Peninsular Thailand to Peninsular Malaysia (Pulau Langkawi).

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Amorphophallus haematospadix Hook.f., Fl. Brit. India 6: 517 (1893). Amorphophallus siamensis Gagnep., Notul. Syst. (Paris) 9: 122 (1941). Distribution: South Thailand to Peninsular Malaysia (Pulau Langkawi). Amorphophallus longituberosus (Engl.) Engl. & Gehrm. in H.G.A. Engler, flanzenr., IV, 23C: 73 (1911). Hydrosme longituberosa Engl., Bot. Tidsskr. 24: 273 (1902). Amorphophallus viridis Ridl., J. Straits Branch Roy. Asiat. Soc. 61: 63 (1912). Distribution: Bangladesh to northern Peninsular Malaysia. Amorphophallus manta Hett. & Ittenbach, Blumea 39: 263 (1994) Distribution: Sumatera and Peninsular Malaysia. Notes: Lowland plants hitherto referred to as A. bufo are in fact either A. manta or perhaps represent a related but undescribed species. Amorphophallus bufo is a montane species. Amorphophallus minor Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 46 (1903). Distribution: Peninsular Malaysia. Endemic. Amorphophallus paeoniifolius (Dennst.) Nicolson, Taxon 26: 337 (1977). Dracontium paeoniifolium Dennst., Schlüssel Hortus Malab.: 38 (1818). Arum campanulatum Roxb., Pl. Coromandel 3: 68 (1820), nom. illeg. Arum rumphii Gaudich., Voy. Uranie: 43 (1826), nom. illeg. Candarum rumphii Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 17 (1832), nom. illeg. Candarum hookeri Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 17 (1832), nom. illeg. Candarum roxburghii Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 17 (1832), nom. illeg. Amorphophallus giganteus Blume, Rumphia 1: 144 (1837), nom. illeg. Conophallus giganteus Schott ex Miq., Fl. Ned. Ind. 3: 198 (1856), nom. illeg. Kunda verrucosa Raf., Fl. Tellur. 2: 82 (1837), nom. illeg.

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Dracontium polyphyllum Dennst., Schlüssel Hortus Malab.: 38 (1818), nom. illeg. Pythion campanulatum Mart., Flora 14: 459 (1831). Amorphophallus campanulatus Decne., Nouv. Ann. Mus. Hist. Nat. 3: 366 (1834). Amorphophallus paeoniifolius var. campanulatus (Decne.) Sivad., Taxon 32: 130 (1983). Amorphophallus dubius Blume, Rumphia 1: 142 (1837). Amorphophallus sativus Blume, Rumphia 1: 145 (1837). Conophallus sativus (Blume) Schott, Prodr. Syst. Aroid.: 35 (1860). Arum decurrens Blanco, Fl. Filip.: 656 (1837). Amorphophallus decurrens (Blanco) Kunth, Enum. Pl. 3: 581 (1841). Arum phalliferum Oken, Allg. Naturgesch. 3(1): 595 (1841). Arum rumphii Oken, Allg. Naturgesch. 3(1): 595 (1841). Plesmonium nobile Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 279 (1864). Amorphophallus chatty Andrews, Ill. Hort. 19: 361 (1872). Amorphophallus virosus N.E.Br., Gard. Chron., n.s., 1885(1): 759 (1885). Amorphophallus rex Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62: 79 (1893). Amorphophallus campanulatus var. blumei Prain, Bengal Pl.: 1109 (1903). Amorphophallus malaccensis Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 46 (1903). Amorphophallus campanulatus f. darnleyensis F.M. Bailey, Queensland Agric. J., n.s., 1: 124 (1914). Amorphophallus gigantiflorus Hayata, Icon. Pl. Formosan. 6: 101 (1916). Hydrosme gigantiflora (Hayata) S.S. Ying, Mem. Coll. Agric. Natl. Taiwan Univ. 31(1): 31 (1991). Amorphophallus microappendiculatus Engl., Notizbl. Bot. Gart. Berlin-Dahlem 8: 458 (1923). Amorphophallus dixenii K. Larsen & S.S. Larsen, Reinwardtia 9: 141 (1974).

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Distribution: India to western Oceania. Not indigenous in Peninsular Malaysia. Notes: Much of the distributional range is based on feral populations originating from earlier introduction as a carbohydrate crop. Amorphophallus perakensis Engl., Pflanzenr., IV, 23C: 90 (1911). Distribution: Peninsular Malaysia. Endemic. Amorphophallus prainii Hook.f., Fl. Brit. India 6: 516 (1893). Amorphophallus loerzingii Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 367 (1920). Amorphophallus panomensis Gagnep., Notul. Syst. (Paris) 9: 120 (1941). Amorphophallus bintangensis Ipor, Tawan & Meekiong, Folia Mal. 11(1): 43 (2010). Distribution: Sumatera and Peninsular Malaysia. Amorphophallus sparsiflorus Hook.f., Fl. Brit. India 6: 516 (1893). Amorphophallus microspadix Engl. & Gehrm. in H.G.A. Engler, Pflanzenr., IV, 23C: 94 (1911), nom. illeg. Distribution: Peninsular Malaysia. Endemic. Amydrium Schott Amydrium Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 127 (1863). (Plate 6) Epipremnopsis Engl., Pflanzenr., IV, 23B: 1 (1908). Taxonomy: Five species occurring from southern Myanmar to South West China as far east as New Guinea. Two species in Peninsular Malaysia. Most recent account: Nicolson (1968c). Ecology: Primary and secondary perhumid broadleaf forests, less often in kerangas, at low to moderate altitudes. On a variety of geologies including limestone.

Plate 6. Amydrium Schott. A–D. Amydrium medium (Zoll. & Moritzi) Nicolson. Amydrium medium is unusual among hemiepiphytic monsteroids in remaining small, usually climbing less than 4 m, and thus flowering low on the tree trunk. Adult plants have the leaves both perforated and pinnately divided (A). Juvenile plants have less elaborately divided leaves (B). As in most Monstereae the spathe is shed during anthesis (C), but unlike most the fruits are indehiscent berries (D).

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Amydrium humile Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 127 (1863). Epipremnum humile (Schott) Hook.f., Fl. Brit. India 6: 549 (1893). Rhaphidophora humilis (Schott) Ridl., Mat. Fl. Malay. Penins. 3: 41 (1907). Distribution: Sumatera and Peninsular Malaysia. Amydrium medium (Zoll. & Moritzi) Nicolson, Blumea 16: 124 (1968). Scindapsus medius Zoll. & Moritzi in H. Zollinger, Syst. Verz.: 82 (1854). Anadendrum medium (Zoll. & Moritzi) Schott, Bonplandia (Hannover) 5: 45 (1857). Epipremnum medium (Zoll. & Moritzi) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 250 (1879). Epipremnopsis media (Zoll. & Moritzi) Engl., Pflanzenr., IV, 23B: 1 (1908). Rhaphidophora huegelii Schott, Prodr. Syst. Aroid.: 384 (1860). Scindapsus huegelii (Schott) Ender, Index Aroid.: 74 (1864). Epipremnopsis huegelii (Schott) Engl., Pflanzenr., IV, 23B: 138 (1908). Rhaphidophora korthalsiana Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 250 (1879). Epipremnum truncatum Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23B: 63 (1908). Epipremnopsis subcordata M. Hotta, Acta Phytotax. Geobot. 22: 2 (1966). Distribution: Southern Myanmar through to Maluku. Anadendrum Schott Anadendrum Schott, Bonplandia (Hannover) 5: 45 (1857). (Plate 7) Taxonomy: At least 30 species in tropical and subtropical East Asia, from Myanmar to South West China, as far east as the Philippines. Many species in Sunda are as yet undescribed. Perhaps eight species in Peninsular Malaysia, but taxonomy chaotic. Most recent account: Ridley (1925). Ecology: Primary and secondary perhumid broadleaf forest at low to moderate altitudes. At least some species are limestone obligates.

Plate 7. Anadendrum Schott. A. An undescribed species allied to Anadendrum affine Schott. Note the distinctive fan-like leaf arrangement, the slender peduncle to the inflorescences, and the conspicuous beaked spathe. B. Another undescribed species in which the stamen filaments are conspicuously extended at staminate anthesis, bringing the anthers out from between the individual flowers. C. The fruits of Anadendrum are immediately distinguishable by the truncate top to the berry, and the linear, transverse stigma. The red fruits are fully ripe.

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Anadendrum angustifolium Engl., Bot. Tidsskr. 24: 272 (1902). Distribution: Southern Peninsular Thailand and Peninsular Malaysia. Notes: Provisionally accepted. Anadendrum latifolium Hook.f., Fl. Brit. India 6: 540 (1893). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Anadendrum marginatum Schott, Oesterr. Bot. Wochenbl. 7: 116 (1857). Scindapsus marginatus (Schott) Ender, Index Aroid.: 74 (1864). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Anadendrum microstachyum (de Vriese & Miq.) Backer & Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 371 (1920). Pothos monopetalus Reinw. ex Miq., Fl. Ned. Ind. 3: 188 (1856). Scindapsus microstachyus de Vriese & Miq. in F.A.W. Miquel, Fl. Ned. Ind. 3: 188 (1856). Pothos malaianus Miq., Fl. Ned. Ind., Eerste Bijv.: 596 (1861). Anadendrum malaianum Backer & Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 372 (1920), nom. illeg. Distribution: Peninsular Malaysia and Java. Notes: Provisionally accepted.

Apoballis Schott Apoballis Schott, Oesterr. Bot. Zeit. 8: 318. (1858). (Plate 8) Taxonomy: About 13 species from southern Myanmar to South Thailand, Sumatera (the majority), and Peninsular Malaysia. Two species in Peninsular Malaysia. Most recent account: Hay & Yuzammi (2000) – treated as a synonym of Schismatoglottis. See also Wong & Boyce (2010).

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Plate 8. Apoballis Schott. A. Apoballis mutata (Hook.f.) S.Y. Wong & P.C. Boyce. Note that the spathe barely opens at anthesis. B. Apoballis brevipes (Hook.f.) S.Y. Wong & P.C. Boyce. All Apoballis species have long initially erect, later decumbent stems not rooting as they go. C. Apoballis acuminatissima (Schott) S.Y. Wong & P.C. Boyce. Inflorescence at staminate anthesis. Note that the spathe limb barely opens and that the interstice (just visible) between the staminate and pistillate flower zones is naked except for a few scattered staminodes. A–B © David Scherberich. C. © David Broederbauer. Used with permission.

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Ecology: Primary and secondary perhumid to evermoist broadleaf forest at low to moderate altitudes. Notes: Apoballis is defined by the decidous leaf sheath and prominent naked interstice interspersed sparsely with staminodes. The spathe usually hardly opens and is semi-persistent into fruiting. All species so far investigated have spiny pollen (smooth in Schismatiglottis). Interestingly, Apoballis investigated produce a floral odour reminiscent of benzaldehyde (almond oil), contrasting to Schismatoglottis (methyl esterase – model airplane glue). This, together with differences in spathe mechanics (Boyce & Hay, 2007), suggests pollinator differences. Apoballis brevipes (Hook.f.) S.Y. Wong & P.C. Boyce, Bot. Stud. (Taipei) 51: 253 (2010). Schismatoglottis brevipes Hook.f., Fl. Brit. Ind. 6: 538 (1893). Schismatoglottis minor Hook.f., Fl. Brit. India 6: 538 (1893). Distribution: North Sumatera and Peninsular Malaysia. Apoballis mutata (Hook.f.) S.Y. Wong & P.C. Boyce, Bot. Stud. (Taipei) 51: 254 (2010). Schismatoglottis mutata Hook.f., Fl. Brit. Ind. 6: 538 (1893). Schismatoglottis batoeensis Engl., Pflanzenr., IV, 23Da: 111 (1912). Schismatoglottis linguiformis Engl., Pflanzenr., IV, 23Da: 93 (1912). Schismatoglottis ridleyana Engl., Pflanzenr., IV, 23Da: 116 (1912). Schismatoglottis rotundifolia Engl., Pflanzenr., IV, 23Da: 122 (1912). Schismatoglottis brooksii Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 341 (1922). Schismatoglottis leptophylla Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 210 (1922). Schismatoglottis monticola Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 202 (1922). Distribution: Southern Myanmar, South Thailand, Sumatera, and Peninsular Malaysia.

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Arisaema Mart. Arisaema Mart., Flora 14(2): 459 (1831). (Plate 9) Alocasia Neck. ex Raf., Fl. Tellur. 3: 64 (1837), nom. rej. Dochafa Schott, Syn. Aroid.: 24 (1856). Muricauda Small, Fl. S.E. U.S.: 227 (1903). Flagellarisaema Nakai, J. Jap. Bot. 25: 6 (1950). Heteroarisaema Nakai, J. Jap. Bot. 25: 6 (1950). Pleuriarum Nakai, J. Jap. Bot. 25: 5 (1950). Ringentiarum Nakai, J. Jap. Bot. 25: 6 (1950). Taxonomy: In excess of 200 species occurring from North America south to Mexico, East and northeastern Africa, the Arabian Gulf, through India to China and Japan and south through South East Asia to the Philippines. Seven species recorded for Peninsular Malaysia, of which two endemic. Most recent account: Ridley (1925). Ecology: Mid-elevation to montane perhumid or everwet evergreen broadleaf forest. Notes: Arisaema is predominantly warm temperate Asian, with the greatest number of species and highest diversity occurring in China and Japan. The majority of Arisaema species are remarkable for their ability to change the sex of the inflorescences dependent on the size, maturity and overall vigour of the plant. Plants flowering for the first time, and weak mature individuals produce staminate inflorescences whereas robust plants in good health flower pistillate. This phenomenon is termed paradioecy.

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Plate 9. Arisaema Blume. A & B. Arisaema fimbriatum Mast. subsp. fimbriatum. A species restricted to limestone and remarkable for the filamentous structures along the entire length of the spadix appendix. C. Arisaema anomalum Hemsl. A species endemic to Peninsular Malaysia. D. Arisaema roxburghii Kunth. E. Arisaema wrayi Hemsl. is one of a series of Asian species in which then spadix appendix in greatly extended and hangs down from the mouth of the spathe. Its role in pollination is still not understood but it may function as a ‘ladder’ to enable the pollinators to gain access and egress from the inflorescence. A–B, D–E © Guy Gusman. C. © David Scherberich. Used with permission.

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Arisaema anomalum Hemsl., J. Bot. 25: 205 (1887). Distribution: Peninsular Malaysia. Endemic. Arisaema filiforme (Reinw.) Blume, Rumphia 1: 102 (1836). Arum filiforme Reinw. in C.L. Blume, Catalogus: 103 (1823). Arisaema filiforme var. typicum Engl., Pflanzenr., IV, 23F: 169 (1920), nom. inval. Arisaema makoyanum Kunth, Index Seminum (B) 1845: 9 (1845). Arisaema stictopodum Miq., Fl. Ned. Ind. 3: 219 (1856). Arisaema fallax Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 123 (1863). Arisaema filiforme var. fallax (Schott) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 542 (1879). Arisaema filiforme f. pedatisectum Kuntze, Revis. Gen. Pl. 2: 739 (1891). Arisaema filiforme f. ternatum Kuntze, Revis. Gen. Pl. 2: 739 (1891). Arisaema filiforme var. sumatranum Engl., Bot. Jahrb. Syst. 37: 142 (1905). Arisaema filiforme var. verrucosum Steenis, Bull. Jard. Bot. Buitenzorg, III, 17: 451(1948). Distribution: Sumatera, Peninsular Malaysia, Java, and Borneo. Arisaema fimbriatum Mast., Gard. Chron., n.s., 1884(2): 680 (1884). subsp. fimbriatum Arisaema putii Gagnep., Notul. Syst. (Paris) 9: 127 (1941). Distribution: Southwestern Thailand to Peninsular Malaysia (Pulau Langkawi). Arisaema laminatum Blume, Rumphia 1: 99 (1836). Distribution: Sumatera, Peninsular Malaysia, Java, Nusa Tenggara, North Borneo, and Philippines (Palawan).

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Arisaema roxburghii Kunth, Enum. Pl. 3: 18 (1841). Arum cuspidatum Roxb., Fl. Ind. ed. 1832, 3: 506 (1832), nom. illeg. Arisaema cuspidatum Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 536 (1879), nom. illeg. Arisaema kunstleri Hook.f., Fl. Brit. India 6: 497 (1893). Arisaema roxburghii var. kunstleri (Hook.f.) Ridl., Fl. Malay Penins. 5: 88 (1925). Arisaema harmandii Engl., Bot. Jahrb. Syst. 25: 27 (1898). Arisaema pierreanum Engl., Pflanzenr., IV, 23F: 159 (1920). Distribution: South Thailand to southern Indo-China and Peninsular Malaysia. Arisaema scortechinii Hook.f., Fl. Brit. India 6: 503 (1893). Distribution: Peninsular Malaysia. Endemic. Arisaema wrayi Hemsl., J. Bot. 25: 205 (1887). Distribution: Southern Peninsular Thailand, Peninsular Malaysia, and North Borneo (Sabah).

Colocasia Schott Colocasia Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 18 (1832), nom. cons. (Plate 10) Leucocasia Schott, Oesterr. Bot. Wochenbl. 7: 34 (1857). Taxonomy: Probably fewer than 15 species distributed from the subtropical eastern Himalayas throughout subtropical and tropical Asia into the tropical western pacific and eastern Australia, with all of this distribution accounted for by C. esculenta (L.) Schott, a carbohydrate and less often green vegetable crop which is cultivated throughout the subtropics and tropics of both hemispheres. Two species in Peninsular Malaysia, one not indigenous. Most recent account: Hay (1996).

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Plate 10. Colocasia Schott. A & B. Colocasia esculenta (L.) Schott. The widespread taro or keladi cina, while not indigenous to Peninsular Malaysia is perhaps the best known aroid in the region. The waxy surface of the leavers shed water into ‘mercury droplets’. The yellow inflorescences smell of overripe pineapple, and are pollinated by Colocasiomyia flies that have a mutalistic reliance on the plant. C & D. Colocasia gigantea (Blume) Hook.f. is an indigenous species, often associated with limestone, although not obligately so. The brilliant white inflorescences are produced successively in fan held beneath the abaxially pruinose leaves. As with C. esculenta, they are pollinated by mutalistic Colocasiomyia.

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Ecology: Colocasia esculenta is predominantly in the lowlands, extending from sea level to lower and mid-montane zones whereas C. gigantea (Blume) Hook.f. is often abundant in seasonally dry forest on limestone, although not obligately so. Colocasia esculenta (L.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 18 (1832). Arum esculentum L., Sp. Pl.: 965 (1753). Caladium esculentum (L.) Vent., Descr. Pl. Nov.: t. 30 (1801). Leucocasia esculenta (L.) Nakai, Bull. Natl. Sci. Mus. Tokyo 31: 127 (1952). Arum colocasia L., Sp. Pl.: 985 (1753). Aron colocasium (L.) St.-Lag., Ann. Soc. Bot. Lyon 7: 119 (1880). Colocasia colocasia (L.) Huth, Helios 11: 134 (1893), nom. inval. Caladium colocasia (L.) W. Wight, Contr. U. S. Natl. Herb. 9: 206 (1905), nom. illeg. Arum chinense L., Amoen. Acad. 4: 234 (1754). Arum peltatum Lam., Encycl. 3: 13 (1789). Colocasia peltata (Lam.) Samp., Herb. Portug.: 12 (1913). Arum lividum Salisb., Prodr. Stirp. Chap. Allerton: 260 (1796). Caladium nymphaeifolium Vent., Descr. Pl. Nov.: t. 30 (1801). Arum nymphaeifolium (Vent.) Roxb., Hort. Bengal.: 65 (1814). Colocasia nymphaeifolia (Vent.) Kunth, Enum. Pl. 3: 37 (1841). Caladium acre R.Br., Prodr.: 336 (1810). Colocasia acris (R.Br.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 18 (1832). Arum colocasioides Desf., Tabl. École Bot., ed. 3: 7 (1829). Caladium colocasioides (Desf.) Brongn., Nouv. Ann. Mus. Hist. Nat. 3: 156 (1834). Caladium violaceum Desf., Tabl. École Bot., ed. 3: 7 (1829). Caladium glycyrrhizum Fraser, Bot. Misc. 1: 259 (1830). Calla virosa Roxb., Fl. Ind. ed. 1832, 3: 517 (1832). Colocasia virosa (Roxb.) Kunth, Enum. Pl. 3: 41 (1841). Zantedeschia virosa (Roxb.) K. Koch, Index Seminum (B) 1854 (App.): 9 (1855). Steudnera virosa (Roxb.) Prain, Bengal Pl.: 1113 (1903). Calla gaby Blanco, Fl. Filip.: 659 (1837). Colocasia vulgaris Raf., Fl. Tellur. 3: 65 (1837).

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Colocasia esculenta var. aquatilis Hassk., Pl. Jav. Rar.: 150 (1840). Colocasia himalensis Royle, Ill. Bot. Himal. Mts.: 407 (1840). Colocasia vera Hassk., Flora 25(2 Beibl. 1): 81 (1842). Colocasia euchlora K. Koch & Linden, Index Seminum (B) 1854(App.): 4 (1855). Alocasia illustris W. Bull, Cat. 1873: 4 (1873). Caladium violaceum Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 492 (1879). Colocasia gracilis Engl., Bot. Jahrb. Syst. 1: 185 (1881). Colocasia neocaledonica Van Houtte, Cat. 1885: (1885). Alocasia dussii Dammer, Gartenflora 1892: 312 (1892). Colocasia antiquorum var. patens Makino in Y. Iinuma, Somoku-Dzusetsu, ed. 3, 4: 1253 (1912). Colocasia antiquorum var. rosea Makino in Y. Iinuma, Somoku-Dzusetsu, ed. 3, 4: 1253 (1912). Colocasia aegyptiaca Samp., Herb. Portug., Ap.: 3 (1914). Colocasia antiquorum var. multifolia Makino, J. Jap. Bot. 1: 4 (1916). Colocasia antiquorum var. rupicola Haines, Bot. Bihar Orissa 5: 867 (1924). Colocasia esculenta var. rupicola (Haines) H.B. Naithani, Fl. Pl. India: 454 (1990). Colocasia antiquorum var. stolonifera Haines, Bot. Bihar Orissa 5: 867 (1924). Colocasia esculenta var. stolonifera (Haines) H.B. Naithani, Fl. Pl. India: 454 (1990). Colocasia antiquorum f. acuatica Makino, J. Jap. Bot. 3: 15 (1926). Colocasia antiquorum f. eguimo Makino, J. Jap. Bot. 3: 15 (1926). Colocasia antiquorum f. oyasetage Makino, J. Jap. Bot. 3: 15 (1926). Colocasia antiquorum f. purpurea Makino, J. Jap. Bot. 3: 15 (1926).

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Colocasia antiquorum f. yamamotoi Makino, J. Jap. Bot. 3: 15 (1926). Colocasia esculenta f. ebiimo Makino, Pract. Hort. 16(10): 996 (1940). Colocasia esculenta f. rotundifolia Makino, Pract. Hort. 16(10): 996 (1940). Colocasia tonoimo Nakai, Iconogr. Pl. Asiae Orient. 3: 231 (1940). Distribution: Throughout tropical and subtropical Asia. Cultivated in the tropics of both hemispheres. Notes: Colocasia esculenta has been utilized as a crop (in Malay: keladi, keladi cina, taro) for millennia and has undoubtedly gone feral on such numerous occasions that it is now unclear to where it is, or was, originally native. Colocasia gigantea (Blume) Hook.f., Fl. Brit. India 6: 524 (1893). Caladium giganteum Blume, Catalogus: 103 (1823). Leucocasia gigantea (Blume) Schott, Oesterr. Bot. Wochenbl. 7: 34 (1857). Colocasia prunipes K. Koch & C.D. Bouché, Index Seminum (B) 1854: 4 (1855). Arisaema fouyou H. Lév., Fl. Kouy-Tchéou: 38 (1914). Distribution: Southern China to Indo-China and Thailand to Peninsular Malaysia, and Sumatera.

Cryptocoryne Fisch. ex Wydler Cryptocoryne Fisch. ex Wydler, Linnaea 5: 428 (1830). (Plate 11) Myrioblastus Wall. ex Griff., Proc. Linn. Soc. London 1: 264 (1845). Taxonomy: Perhaps 60 species occurring India and Sri Lanka through tropical Asia to New Guinea and northwards through Thailand and Indochina to southwestern China. Currently there are 24 taxa recognized for the Peninsula, including 10 species, one nothospecies, two varieties, and one nothovariety, of which six species, one variety and one nothovariety are endemic. Most recent account: Othman, Jacobsen & Mashhor (2009).

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Plate 11. Cryptocoryne Fisch. ex Wydler. A. Cryptocoryne schulzei de Wit, is a Johor endemic. B. Cryptocoryne elliptica N.E.Br., another Peninsular Malaysian endemic. C. Cryptocoryne × purpurea Ridl is a stabilized naturally-occurring hybrid of C. cordata × C. griffithii. D. Cryptocoryne affinis N.E.Br. is also endemic to the Peninsula. A–D © Ahmad Sofiman bin Othman.

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Ecology: Peninsular Malaysian Cryptocoryne are all aquatics (very rarely helophytes: C. ciliata (Roxb.) Fisch. ex Wylder) in freshwater or brackish streams in lowland to mid-elevation perhumid or everwet broadleaf forest. Only rarely do species occur in larger rivers in open habitats. Notes: The inflorescence and fruits of Cryptocoryne are immediately diagnostic. The spathe comprises an inflated basal tube, the kettle, with united margins, an upper tube, sometimes twisted, with fused margins, and a terminal limb which opens into a flat, ovate surface that may either reflex, remain erect, when it may be spirally twisted, or arch forwards to form a hood and is usually terminated with a short to very long tail; a distinctive raised to callus-like collar is often found at the transition between the upper tube and limb. The fruits of Cryptocoryne are unusual in the family in being dehiscent, the individual carpels opening, with the fully open infructescence resembling a star. Cryptocoryne affinis N.E.Br. in J.D. Hooker, Fl. Brit. India 6: 494 (1893). Cryptocoryne haerteliana Jacobsen ex Milkuhn, Wochenschr. Aquar. Terrarienkunde 43: 288 (1949). Cryptocoryne affinis subsp. haerteliana ( Jacobsen ex Milkuhn) Schopfel, Inform. Z.A.G. Wasserpflanzen 1977(4): 5 (1977). Distribution: Peninsular Malaysia. Endemic. Cryptocoryne ciliata (Roxb.) Fisch. ex Wydler, Linnaea 5: 428 (1830). Ambrosina ciliata Roxb., Pl. Coromandel 3: 90 (1820). Arum ciliatum (Roxb.) Loudon, Encycl. Pl., Suppl. 2: 150 (1841). Ambrosina ciliaris Spreng., Syst. Veg. 3: 771 (1826). Cryptocoryne alata Griff., Iconogr. Pl. Asiae Orient. 3: t. 170 (1851). Cryptocoryne elata Griff., Not. Pl. Asiat. 3: 134 (1851). Cryptocoryne drymorrhisa Zipp. ex Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 122 (1863). Cryptocoryne ciliata var. latifolia Rataj, Stud. Ceskoslov. Akad. Ved 3: 38 (1975). Distribution: India to New Guinea.

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Cryptocoryne cordata Griff., Not. Pl. Asiat. 3: 138 (1851). var. cordata. Cryptocoryne kerrii Gagnep., Notul. Syst. (Paris) 9: 132 (1941). Cryptocoryne siamensis var. kerrii (Gagnep.) Rataj, Stud. Ceskoslov. Akad. Ved 3: 93 (1975). Cryptocoryne siamensis Gagnep., Notul. Syst. (Paris) 9: 132 (1941). Cryptocoryne blassii de Wit, Aquarien-Terrar.-Z. 13(4): 115 (1960). Cryptocoryne evae Rataj, Folia Geobot. Phytotax. 9: 314 (1974). Cryptocoryne evae var. recordata Rataj, Folia Geobot. Phytotax. 9: 314 (1974). Cryptocoryne siamensis var. ewansii Rataj, Stud. Ceskoslov. Akad. Ved 3: 93 (1975). Cryptocoryne stonei Rataj, Stud. Ceskoslov. Akad. Ved 3: 95 (1975). Distribution: Southern Peninsular Thailand to southeastern Peninsular Malaysia, Java, and Borneo. var. diderici (de Wit) N. Jacobsen, Aqua Pl. 27: 151 (2002). Cryptocoryne diderici de Wit, Meded. Bot. Tuinen Belmonte Arbor. 13: 279 (1970). Distribution: Northern Peninsular Malaysia. Endemic. Cryptocoryne decus-silvae de Wit, Aquarium (Netherlands) 46: 177 (1976). Distribution: Peninsular Malaysia. Endemic. Cryptocoryne elliptica N.E.Br. in J.D. Hooker, Fl. Brit. India 6: 495 (1893). Distribution: Peninsular Malaysia. Endemic. Cryptocoryne griffithii Schott, Syn. Aroid.: 1 (1856). Distribution: Peninsular Malaysia, Singapore, and Borneo (South Kalimantan).

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Cryptocoryne longicauda Becc. ex Engl., Bull. Soc. Tosc. Ortic. 4: 302 (1879). Cryptocoryne caudata N.E.Br., J. Linn. Soc., Bot. 18: 242 (1880). Cryptocoryne johorensis Engl., Pflanzenr., IV, 23F: 244 (1920). Distribution: Sumatera, Peninsular Malaysia, and Borneo. Cryptocoryne minima Ridl., J. Straits Branch Roy. Asiat. Soc. 54: 61 (1910). Cryptocoryne zewaldiae de Wit, Aquarienpfl., ed. Deutsch: 206 (1971). Cryptocoryne gasseri N. Jacobsen, Bot. Not. 132: 144 (1979). Cryptocoryne amicorum de Wit & N. Jacobsen, Nordic J. Bot. 2: 455 (1982). Distribution: Sumatera and Peninsular Malaysia. Cryptocoryne nurii Furtado, Gard. Bull. Straits Settlem. 8: 147 (1935). Distribution: Southern Peninsular Malaysia. Endemic. Cryptocoryne × purpurea Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 44 (1904). [C. cordata × C.griffithii]. Cryptocoryne hejnyi Rataj, Folia Geobot. Phytotax. 9: 313 (1974). nothovar. purpurea. [C. cordata × C. griffithii]. Distribution: Peninsular Malaysia. Endemic. Cryptocoryne schulzei de Wit, Aquarium (Netherlands) 42: 14 (1971). Distribution: Peninsular Malaysia ( Johor). Endemic. Cryptocoryne × timahensis Bastm., Gard. Bull. Singapore 53: 11 (2001). [C. cordata × C. nurii]. Distribution: Singapore.

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Notes: Yet to be recorded for Peninsular Malaysia, although it should be sought where the parents occur. Cryptocoryne zukalii Rataj, Folia Geobot. Phytotax. 9: 313 (1974). Distribution: Peninsular Malaysia. Endemic.

Cyrtosperma Griff. Cyrtosperma Griff., Not. Pl. Asiat. 3: 149 (1851). (Plate 12) Arisacontis Schott, Bonplandia (Hannover) 5: 129 (1857). Taxonomy: About 13 species in tropical Southeast Asia as far east as western Oceania, with the majority restricted to Papuasia. One species in Peninsular Malaysia. Most recent account: Hay (1988). Ecology: Lowlands in freshwater swamp forest margins, sometimes persisting in flooded pasture. Notes: Cyrtosperma is a genus of large to massive rhizomatous herbs with mottled and usually heavily armed petioles and hastate to sagittate leaves. Inflorescences are solitary and carried on a peduncle similar to the petioles. The majority of Cyrtosperma occur in Papuasia, with only one species in Peninsular Malaysia – C. merkusii (Hassk.) Schott, a large plant (2 m or more tall) almost always found in open swampy places in association with habitation, has leaves held erect and petioles normally only lightly armed. Cyrtosperma merkusii (Hassk.) Schott, Oesterr. Bot. Wochenbl. 7: 61 (1857). Lasia merkusii Hassk., Cat. Hort. Bot. Bogor.: 59 (1844). Cyrtosperma lasioides Griff., Not. Pl. Asiat. 3: 150 (1851). Arisacontis chamissonis Schott, Bonplandia (Hannover) 5: 129 (1857). Cyrtosperma chamissonis (Schott) Merr., Philipp. J. Sci., C 9: 65 (1914).

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Plate 12. Cyrtosperma Griff. A–D. A. Young plant showing the distinctive leaves. B. Detail of the petiole bases with their distinctive ascending prickles. C. Inflorescence at pistillate anthesis. Note the conspicuous stigma droplets. D. Mature infructescence. The nodding posture, marcescentpersistent spathe, and fruits barely emerging from the surrounding tepals are diagnostic.

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Cyrtosperma edule Schott, Bonplandia (Hannover) 9: 367 (1861). Cyrtosperma cuspidilobum Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 284 (1864). Cyrtosperma dubium Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 284 (1864). Cyrtosperma intermedium Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 284 (1864). Cyrtosperma merkusii var. intermedium (Schott) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 271 (1879). Cyrtosperma merkusii var. giganteum Nadeaud, J. Bot. (Morot) 11: 116 (1897). Cyrtosperma nadeaudianum J.W. Moore, Bernice P. Bishop Mus. Bull. 102: 22 (1933). Cyrtosperma bantamense Koord., Bull. Jard. Bot. Buitenzorg, III, 1: 160 (1919). Distribution: West Malesia to western Oceania.

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Epipremnum Schott Epipremnum Schott, Bonplandia (Hannover) 5: 45 (1857). (Plate 13) Anthelia Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 127 (1863). Taxonomy: Approximately 15 species occurring throughout tropical Asia from North East India to the Ryukyu Islands southwards to eastern Australia (Queensland) and Oceania (Cook Islands: Rarotonga). Two species in Peninsular Malaysia. Most recent account: Boyce (1998). Ecology: Low to mid-elevation perhumid broadleaf forest, and occasionally persisting in disturbed areas or growing lithophytically in exposed situations. The widespread Epipremnum pinnatum is often associated with limestone. Notes: Confusion can occur between Epipremnum and the other Malesian monsteroid genera. If fruits are mature, seed characters separate Epipremnum and Rhaphidophora. Epipremnum has fruits with few large, strongly curved, seeds with a bony, smooth to ornamented testa. The fruits of Rhaphidophora each contain many small ellipsoid seeds with a brittle, smooth testa. Alternatively, immature fruits can be dissected and the number of ovules counted (few in Epipremnum, almost always many in Rhaphidophora). Certain Scindapsus [notably S. latifolius M.Hotta (Borneo), S. splendidus Alderw. and S. roseus Alderw. (both Sumatera)] are very similar in appearance to the entire leaved Epipremnum species. The only way to differentiate these Scindapsus and Epipremnum species, aside from field experience, is to observe inflorescences or, better, semi-mature infructescences. Scindapsus has fruits with a solitary curved seed. However, certain Epipremnum species (e.g. E. ceramense and E. falcifolium) seem to habitually abort all but one ovule and produce fruits with a solitary seed. Species of Anadendrum Schott are often collected as ‘Rhaphidophora’, ‘Scindapsus’ or ‘Epipremnum’. Anadendrum, together with most Pothos L. and Pedicellarum M.Hotta (both subfamily Pothoideae), are the only simple-leaved Asian aroid climbers with reticulate venation (Amydrium humile, also with reticulate venation, can occasionally be observed as a low climber, but then with a solitary inflorescence). Additionally, Anadendrum lacks trichosclereids and can be distinguished from all Monstereae using a leaf tear. Using floral characters Anadendrum (each flower with a membranous perigon of fused tepals) is a

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Plate 13. Epipremnum Schott. A–C. Epipremnum pinnatum (L.) Engl. A. Mature plant. B. detail of active shoot apex with the conspicuous dense fibrous cataphyll, prophyll and petiolar sheath fibre. C. Synflorescence with two developing infructescences and a inflorescence at staminate anthesis. D. Detail of spadix base at late staminate anthesis. Note the conspicuous exserted stamens and that lowermost flowers produce no stamens. These flowers are functionally sterile (gynoecia also lacking ovules) and are associated with the production of sticky droplets that are likely involved with pollinator reward.

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singular genus and should not be confused with any other Asian climbing Araceae. Confusion can occur between Anadendrum and genera of Pothoideae (i.e., Pothos, Pothoidium Schott and Pedicellarum) that also lack trichosclereids. However, Pothos (except subg. Pothos) and Pedicellarum are instantly recognizable by the intramarginal veins crossing the primary venation. Anadendrum flowers on clinging climbing shoots whereas almost all Pothos flower on free lateral shoots. In fruit both have red, somewhat juicy berries. However, those of Anadendrum are apically truncate with a prominent linear stigma whereas Pothos has ellipsoid to globose berries with a tiny, punctiform to slightly elongated stigma. Confusion is also possible at species level. Epipremnum pinnatum is vegetatively most similar to Rhaphidophora korthalsii Schott but easilyobserved distinguishing characters are present. Mature leaves of R. korthalsii are invariably pinnatisect (variously pinnatifid, pinnatipartite or pinnatisect in E. pinnatum) with individual pinnae, even the narrowest having more than one primary lateral vein (one per pinna in E. pinnatum). The internodes of R. korthalsii lack the prominent irregular longitudinal whitish crests and older stems lack the distinctive matt to sub-lustrous pale brown papery epidermis typical of E. pinnatum. The feeder roots of R. korthalsii are scaly whereas they are lenticellate-corky in E. pinnatum. The pre-adult stage of R. korthalsii is a ‘shingle’ climber with oblong-elliptic to ovate, slightly falcate, and upwards pointing leaves overlapping in the manner of roof tiles. Juvenile plants of E. pinnatum are sprawling to climbing with long-petioled ‘conventional’ leaves. Fertile material of R. korthalsii and E. pinnatum is readily separated by the shape of the style apex [round to oval (R. korthalsii) versus angled (E. pinnatum)] and the shape of the stigma and its orientation to the spadix [punctiform and circumferential (R. korthalsii) versus linear and longitudinal (E. pinnatum)]. Epipremnum pinnatum and Rhaphidophora tetrasperma Hook. f., another pinnatifid and perforate-bladed species can be confused. Juvenile R. tetrasperma is a shingle plant similar in appearance to R. korthalsii. Flowering-size plants have smooth stems and unequal ovate-elliptic coriaceous leaf blades (longitudinally crested stems and more or less equal, ovate to oblong-elliptic and sub-membranaceous leaf blades in E. pinnatum) and a more scandent habit, with leaves scattered along sinuous stems.

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Epipremnum giganteum (Roxb.) Schott, Bonplandia (Hannover) 5: 45 (1857). Pothos giganteus Roxb., Fl. Ind. 1: 455 (1820). Monstera gigantea (Roxb.) Schott, Wiener Z. Kunst 4: 1028 (1830). Scindapsus giganteus (Roxb.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). Rhaphidophora gigantea (Roxb.) Ridl., Mat. Fl. Malay. Penins. 3: 45 (1907). Distribution: South Thailand, southern Indo-China, Sumatera(?), and Peninsular Malaysia. Epipremnum pinnatum (L.) Engl., Pflanzenr., IV, 23B: 60 (1908). Pothos pinnatus L., Sp. Pl. ed. 2: 1324 (1763). Monstera pinnata (L.) Schott, Wiener Z. Kunst 4: 1028 (1830). Scindapsus pinnatus (L.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). Rhaphidophora pinnata (L.) Schott, Bonplandia (Hannover) 5: 45 (1857). Polypodium laciniatum Burm.f., Fl. Indica: 231 (1768). Rhaphidophora laciniata (Burm.f.) Merr., Philipp. J. Sci. 19: 342 (1921). Pothos caudatus Roxb., Fl. Ind. 1: 476 (1820). Monstera caudata (Roxb.) Schott, Wiener Z. Kunst 4: 1028 (1830). Scindapsus caudatus (Roxb.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). Rhaphidophora caudata (Roxb.) Schott, Prodr. Syst. Aroid.: 382 (1860). Pothos pinnatifidus Roxb., Fl. Ind. 1: 476 (1820). Scindapsus pinnatifidus (Roxb.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). Rhaphidophora pinnatifida (Roxb.) Schott, Bonplandia (Hannover) 5: 45 (1857). Pothos decursivus Wall., Pl. Asiat. Rar. 2: 83 (1831), nom. illeg. Scindapsus decursivus Moritzi in H. Zollinger, Syst. Verz.: 82 (1854), nom. illeg. Scindapsus forsteri Endl., Ann. Wiener Mus. Naturgesch. 1: 161 (1836). Scindapsus dilaceratus K. Koch & Sello, Index Seminum (B) 1853 (App.): 5 (1853). Monstera dilacerata (K. Koch & Sello) K. Koch, Index Seminum (B) 1855 (App.): 5 (1855). Tornelia dilacerata (K. Koch & Sello) Schott, Prodr. Syst. Aroid.: 356 (1860). Rhaphidophora dilacerata (K. Koch & Sello) K. Koch in E.von Regel, Gartenflora: 5 (1864). Epipremnum mirabile Schott, Gen. Aroid.: t. 79 (1858).

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Rhaphidophora wallichii Schott, Prodr. Syst. Aroid.: 383 (1860). Rhaphidophora cunninghamii Schott, Bonplandia (Hannover) 9: 367 (1861). Rhaphidophora vitiensis Schott, Bonplandia (Hannover) 9: 367 (1861). Scindapsus bipinnatifidus Teijsm. & Binn., Cat. Hort. Bot. Bogor.: 65 (1866). Epipremnum elegans Engl., Bull. Soc. Tosc. Ortic. 4: 269 (1879). Philodendron dilaceratum Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 265 (1879). Rhaphidophora lovellae F.M. Bailey, Queensland Agric. J. 1: 453 (1897). Epipremnum mirabile f. eperforatum Engl., Bot. Jahrb. Syst. 25: 12 (1898). Epipremnum mirabile f. multisectum Engl., Bot. Jahrb. Syst. 25: 12 (1898). Epipremnum pinnatum f. multisectum (Engl.) Engl., Pflanzenr., IV, 23B: 63 (1908). Rhaphidophora merrillii Engl., Bot. Jahrb. Syst. 37: 115 (1905). Epipremnum merrillii Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23B: 137 (1908). Epipremnum angustilobum K. Krause, Bot. Jahrb. Syst. 45: 659 (1911). Epipremnum robinsonii K. Krause, Notizbl. Königl. Bot. Gart. Berlin 5: 266 (1912). Epipremnum formosanum Hayata, Icon. Pl. Formosan. 5: 239 (1915). Rhaphidophora formosana (Hayata) M. Hotta, Mem. Fac. Sci. Kyoto Univ., Ser. Biol. 4: 83 (1970), nom. illeg. Epipremnum elegans f. ternatensis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 169 (1922). Philodendron nechodomae Britton, Bot. Porto Rico 6: 335 (1926). Rhaphidophora neocaledonica Guillaumin, Bull. Soc. Bot. France 84: 160 (1937). Epipremnum glaucicephalum Elmer, Leafl. Philipp. Bot. 10: 3620 (1938), no latin descr.

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Distribution: Andaman Islands through to western Oceania.

Furtadoa M.Hotta Furtadoa M.Hotta, Acta Phytotax. Geobot. 32: 142 (1981). Taxonomy:Two species, one (F. sumatrana M. Hotta) endemic to Sumatera, one (F. mixta (Ridl.) M. Hotta) endemic to Peninsular Malaysia. Most recent account: Hotta (1985). Ecology: The Malaysian species occurs terrestrially in lowland evermoist to everwet, broadleaf forest, while the Sumateran species is a facultative rheophytic. Furtadoa mixta (Ridl.) M. Hotta, Gard. Bull. Singapore 38: 53 (1985). Homalomena mixta Ridl., J. Bot. 40: 36 (1902). Distribution: Peninsular Malaysia. Endemic.

Hapaline Schott Hapaline Schott, Gen. Aroid.: 44 (1858), nom. cons. Hapale Schott, Oesterr. Bot. Wochenbl. 7: 8 (1857), nom. rej. Taxonomy: Eight species ranging from Myanmar to China (Yunnan) and south through Thailand and Peninsular Malaysia to Brunei. One species in Peninsular Malaysia. Most recent account: Boyce (1996). Ecology: Low elevation perhumid broadleaf evergreen forest on limestone. Notes: Hapaline are diminutive to moderate-sized, slender to slightly robust tuberous or stoloniferous, clump-forming terrestrial, seasonally dormant or evergreen herbs with leaf blades ranging from thin-textured to rather thick and leathery, and ranging in shape from ovate to hastate, with posterior lobes either

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present or absent, occasionally even in the same species. Leaf blade variegation, in the form or irregular, often diffuse, silver-grey to pale green blotches, occurs in many species. Hapaline species all have white inflorescences, occasionally tinged green or greyish pink. Staminate flowers are fused into peltate synandria, with the synconnectives massively enlarged and with the thecae inserted on the lower margin of the connective tissue. Hapaline brownii Hook.f., Fl. Brit. India 6: 521 (1893). Distribution: Southern Peninsular Thailand and Peninsular Malaysia.

Hestia S.Y. Wong & P.C. Boyce Hestia S.Y. Wong & P.C. Boyce, Bot. Stud. (Taipei) 51: 250. (2010). (Plate 14) Taxonomy: One species disjunct between the Malay Peninsula, where it is known only from Perak, and Borneo, where it occurs in numerous but widely scattered localities in Sarawak and especially Brunei. Most recent account: Wong & Boyce (2010). Ecology: Restricted to raised podsols in swampy areas in kerangas formations within lowland and hill perhumid broadleaf tropical forest. Notes: A very distinctive genus easily recognised by the rather numerous nodding inflorescences on relatively very long slender wiry peduncles. There is only a weak constriction present between the lower spathe and upper spathe, and the orifice of the lower spathe is open during fruiting. The entire non-pistillate portion of the spadix often dries and adheres to the spathe limb with the whole combined unit shedding. The sole species, Hestia longifolia (Ridl.) S.Y. Wong and P.C. Boyce has vegetative modules that readily disarticulate from the deep-seated rhizome. The function of the disarticulation in this podsol-obligated species is not clear, but it is speculated that it may be an adaption to fire resistance in a highly fire-prone habitat. Perhaps enabling the shoot unit to be destroyed in some way prevents damage to the main perennating system.

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Plate 14. Hestia S.Y. Wong & P.C. Boyce. A–G. Hestia longifolia (Ridl.) S.Y. Wong & P.C. Boyce. A. Plant in habitat on podsol. B. Nodding inflorescences, the inflorescence to the left at pistillate anthesis; upper right at staminate anthesis, lower right post-anthesis with spathe beginning to degrade prior to being shed. C. Spathe slightly gaping at pistillate anthesis. D. Spadix with the spathe removed artificially to reveal the pistillate and staminate flower zones. E. Inflorescence postanthesis with spathe limb rotting at junction of lower spathe, later to be shed together with spent part of spadix. F. Infructescences with the distinctive narrowly campanulate lower spathe. G. Fruits.

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Hestia longifolia (Ridl.) S.Y. Wong & P.C. Boyce, Bot. Stud. (Taipei) 51: 252. Schismatoglottis longifolia Ridl. J. Bot. 40: 37 (1902). Distribution: Peninsular Malaysia (Perak) and Borneo (Sarawak).

Homalomena Schott Homalomena Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 20 (1832). (Plates 15 & 16) Homalonema Endl., Gen. Pl.: 238 (1837). Spirospatha Raf., Fl. Tellur. 4: 8 (1838). Cyrtocladon Griff., Not. Pl. Asiat. 3: 147 (1851). Chamaecladon Miq., Bot. Zeitung (Berlin) 14: 564 (1856). Adelonema Schott, Prodr. Syst. Aroid.: 316 (1860). Diandriella Engl., Nova Guinea 8: 20 (1910). Taxonomy: As many as 500 species distributed throughout Indo-Malesia to southern China eastwards to the Solomon Islands with centres of diversity in Sumatera, Borneo, and New Guinea, and ca 12 Neotropical species (section Curmeria but generic status very doubtful). Homalomena is taxonomically chaotic in Peninsular Malaysia with 22 names but much remains to be done. Likely many new species await description. Most recent account: Furtado (1939). Ecology: Primarily understory herbs in lowland evermoist to everwet, or less often perhumid, tropical or subtropical broadleaf forest, but also reach the midmontane zone; sometimes rheophytic, ver y rarely helophytic, occasionally relictual in regrowth and along road cuttings. Notes: Two very closely allied genera in Asia, Homalomena Schott and Furtadoa M. Hotta, mostly with strongly aromatic (terpenoids) tissues and spathes fully persistent to fruit maturity. Homalomena is widespread throughout tropical and subtropical Asia, with significant centres of endemism on Sumatera, Borneo, and

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Plate 15. Homalomena Schott (Homalomena Supergroup). A–D. Homalomena pontederifolia Hook.f. The large spathe (this ca. 11 cm,) lacking a constriction between the lower spathe and the limb, is characteristic of the Homalomena Supergroup. Note the conspicuous interpistillar staminodes in D. Note also (B) that post-anthesis the peduncle becomes declinate and thus the infructescences are pendulous.

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Plate 16. Homalomena Schott (Chamaecladon Supergroup). A–C. Homalomena humilis ( Jack) Hook.f. Spathes of the Chamaecladon Supoergroup also lack a constriction above the lower spathe but the spathes are very small (< 2 cm). A. Flowering plant, red foliage expression; B. Plant with inflorescences at various stages of development, the sequence running from right (youngest) to left (oldest). Note the small size of the spathes and the lack of a constriction. C. Detail of inflorescence at pistillate anthesis; note that spadix is fertile to the tip and that the flowers have two anthers.

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New Guinea. Furtadoa is restricted to Sumatera and Peninsular Malaysia and differs from Homalomena primarily by each staminate flower with an associated pistillode. Species of Homalomena are frequently confused with Schismatoglottis, although the infructescences of Homalomena, in which the spathes wholly persistent to fruit maturity, and then split basally and upwards to reveal the fruits are very different from those of Schismatoglottis in which the spathe limb is shed, often while still fresh, during anthesis, while the lower spathe persists into fruiting as a barrel-shaped to ellipsoid covering, splitting from the top downwards to release the ripe fruits. Homalomena is the most speciose aroid genus in tropical, with in excess of 350 species, of which fewer than half have been described. The genus comprises minute to very large clumping, tufted, rarely creeping, and very rarely climbing mostly strongly aromatic herbs with cordate to lanceolate leaves. Inflorescences generally open for only a few hours before closing again and then soon becoming pendulous with the spathe persisting, and occasionally becoming brightly coloured, around the developing infructescence. Homalomena divide into three morphotaxa (morphologically cohesive but phylogenetically untested higher taxa) termed ‘supergroups’ (see Boyce & Wong, 2008): Homalomena, Chamaecladon, and Cyrtocladon. These supergroups are defined by the following characters: The Homalomena supergroup comprises medium to large creeping to erect plants with strongly aromatic tissues, pleionanthic, or rarely hapaxanthic, shoot modules and spathes greater than 1.5 cm long, with no or only a very weak constriction between the upper and lower spathe. Spathe movement during anthesis, where known, comprises simple gaping and then closing of the spathe limb, no spadix movements have been recorded. Pistillate flowers with the associated staminode equalling or exceeding the ovary. Ovary three to four locular. Staminate flowers with three to four, rarely five to six anthers. Most of the larger species encountered in the Peninsula belong to this supergroup. The Chamaecladon supergroup comprises small to minute often creeping, less often erect plants with odourless, or very rarely aromatic, tissues; as far as is known only pleionanthic shoot modules, and spathes less than 1 cm, very rarely up to 1.5 cm long, with no constriction between the upper and lower

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spathe. Spathe movement during anthesis, where known, comprises simple gaping and closing of the spathe limb. No spadix movement recorded. Pistils with the associated staminode much shorter than ovary. Ovary two to three locular. Staminate flowers with two to three anthers. The Cyrtocladon supergroup comprises medium to very large erect to creeping plants with strongly aromatic tissues, pleionanthic (but very few studied) shoot modules and spathes greater than 2 cm long, with weak to moderate to pronounced constriction between the upper and lower spathe. All of the several species so far studied undergo a complex series of spathe and spadix movements during anthesis. Pistillate flowers with the associated staminode equalling or exceeding the ovary. Ovary three to four locular. Staminate flowers with three to four, rarely five to six anthers. Homalomena argentea Ridl., J. Bot. 40: 35 (1902). Homalomena argentea var. purpurascens Furtado, Gard. Bull. Straits Settlem. 10: 188 (1939). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena asmae Baharuddin & P.C. Boyce, Acta Phytotax. Geobot. 60(3): 163 (2010). Distribution: Peninsular Malaysia (Perak). Notes: Locally endemic. Homalomena brevispatha Engl. Bot. Tidsskr. 24: 274 (1902). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Notes: Provisionally accepted. Homalomena confusa Furtado, Gard. Bull. Straits Settlem. 10: 189 (1939). Homalomena angustifolia var. parvula Ridl., Fl. Malay Penins. 5: 109 (1925).

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Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena corneri Furtado, Gard. Bull. Straits Settlem. 10: 191 (1939). Distribution: Peninsular Malaysia ( Johor). Notes: Provisionally accepted. Homalomena curtisii Ridl., J. Bot. 40: 34 (1902). Distribution: Peninsular Malaysia (Perak). Notes: Provisionally accepted. Homalomena curvata Engl., Pflanzenr., IV, 23Da: 53 (1912). Distribution: Peninsular Malaysia (Melaka). Notes: Provisionally accepted. Homalomena griffithii (Schott) Hook.f., Fl. Brit. India 6: 534 (1893). Chamaecladon griffithii Schott, Bonplandia (Hannover) 6: 369 (1858). Chamaecladon obliquatum Schott, Bonplandia (Hannover) 6: 369 (1858). Homalomena obliquata (Schott) Hook.f. Fl. Brit. India 6: 534 (1893). Homalomena griffithii var. obliquata (Schott) Ridl. J. Straits Branch Roy. Asiat. Soc. 44: 177 (1905). Homalomena griffithii forma obliquata (Schott) Furtado Gard. Bull. Straits Settlem. 10: 196 (1939). Chamaecladon ovatum Schott, Bonplandia (Hannover) 7: 30 (1859). Homalomena ovata (Schott) Hook.f. Fl. Brit. India 6: 536 1893. Homalomena griffithii var. ovata (Schott) Engl. Pflanzenr., IV, 23Da: 44 (1912). Homalomena trapezifolia Hook.f., Fl. Brit. India 6: 534 (1893). Homalomena kingii Hook.f., Fl. Brit. India 6: 535 (1893). Homalomena griffithii forma kingii (Hook.f.) Furtado, Gard. Bull. Straits Settlem. 10: 196 (1939).

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Homalomena deltoidea Hook.f., Fl. Brit. India 6: 536 (1893). Homalomena elliptica Hook.f., Fl. Brit. India 6: 536 (1893). Homalomena falcata Ridl. J. Bot. 40: 35 (1902). Homalomena griffithii forma falcatoidea Furtado, Gard. Bull. Straits Settlem. 10: 197 (1939). Homalomena multinervia Ridl., J. Bot. 40: 36 (1902). Homalomena elliptica var. paucinervia Furtado, Gard. Bull. Straits Settlem. 10: 193 (1939). Homalomena griffithii var. acuminata Ridl. J. Straits Branch Roy. Asiat. Soc. 44: 177 (1905). Homalomena griffithii forma acuminata (Ridl.) Furtado, Gard. Bull. Straits Settlem. 10: 197 (1939). Homalomena grabowskii Engl. Pflanzenr., IV, 23Da: 45 (1912) Homalomena hayupensis Engl., Bot. Jahrb. Syst. 48: 93 (1912). Homalomena griffithii forma eugriffithii Furtado, Gard. Bull. Straits Settlem. 10: 195 (1939), nom. illeg. Homalomena griffithii forma sigmoidea Furtado, Gard. Bull. Straits Settlem. 10: 196 (1939). Homalomena griffithii var. sumatrana Furtado, Gard. Bull. Straits Settlem. 10: 198 (1939). Homalomena habokoana Alderw. Bull. Jard. Bot. Buitenzorg, III, 4: 183 (1922). Distribution: Southern Peninsular Thailand, Sumatera, Peninsular Malaysia, and Borneo. Homalomena hendersonii Furtado, Gard. Bull. Straits Settlem. 10: 198 (1939). Distribution: Peninsular Malaysia (Kelantan). Notes: Provisionally accepted. Homalomena humilis ( Jack) Hook.f., Fl. Brit. India 6: 533 (1893). Calla humilis Jack, Malayan Misc. 1(1): 22 (1820). Chamaecladon humile ( Jack) Miq., Fl. Ned. Ind. 3: 213 (1856).

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Calla angustifolia Jack, Malayan Misc. 1(1): 24 (1820). Chamaecladon angustifolium ( Jack) Schott, Bonplandia (Hannover) 6: 369 (1858). Homalomena angustifolia ( Jack) Hook.f., Fl. Brit. India 6: 533 (1893). Aglaonema pygmaeum Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 161 (1842). Chamaecladon pygmaeum (Hassk.) Engl., Monogr. Phan. 2: 345 (1879). Homalomena pygmaea (Hassk.) Engl., Bot. Jahrb. Syst. 25: 18 (1898). Aglaonema pygmaeum var. majus Hassk., Syst. Verz.: 76 (1854). Homalomena humilis var. major (Hassk.) Furtado, Gard. Bull. Straits Settlem. 10: 200 (1939). Chamaecladon lanceolatum Miq., Bot. Zeitung (Berlin) 14: 564 (1856). Chamaecladon purpurascens Schott, Bonplandia (Hannover) 6: 369 (1858). Homalomena pygmaea var. purpurascens (Schott) Engl., Bot. Jahrb. Syst. 25: 18 (1898). Homalomena purpurascens (Schott) Hook.f., Fl. Brit. India 6: 535 (1893). Homalomena pumila var. purpurascens (Schott) Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 176 (1906). Chamaecladon pygmaeum var. latifolium Engl., Malesia 1: 283 (1882). Homalomena pumila var. latifolia (Engl.) Ridl. J. Straits Branch Roy. Asiat. Soc. 44: 177 (1905). Homalomena pygmaea var. latifolia (Engl.) Engl. Pflanzenr., IV, 23Da: 36 (1912). Homalomena pumila Hook.f. Fl. Brit. India 6: 533 (1893). Homalomena humilis var. pumila (Hook.f.) Furtado Gard. Bull. Straits Settlem. 10: 203 (1939). Homalomena velutina Scort. ex Hook.f., Fl. Brit. India 6: 534 (1893). Homalomena humilis var. velutina (Scort.ex Hook.f.) Ridl., Materials Fl. Mal. Peninsular 3: 25 (1907). Homalomena propinqua Ridl., J. Bot. 40: 35 (1902), nom. illeg. Homalomena undulatifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 57: 111 (1911). Homalomena humilis var. undulata Ridl. Fl. Malay Penins. 5: 105 (1925). Homalomena angustifolia var. ophirensis Ridl. ex Engl. Pflanzenr., IV, 23Da: 31 (1912). Homalomena humilis var. ophirensis (Ridl. ex Engl.) Furtado, Gard. Bull. Straits Settlem. 10: 203 (1939).

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Homalomena pygmaea var. pumila Engl., Pflanzenr., IV, 23Da: 36 (1912). Homalomena coccinea Alderw. Bull. Jard. Bot. Buitenzorg, III, 4: 180 (1922). Homalomena humilis var. major subvar. coccinea (Alderw.) Furtdao, Gard. Bull. Straits Settlem. 10: 201 (1939). Homalomena johorensis Engl. Pflanzenr., IV, 23Da: 37 (1912). Homalomena pusilla Alderw. Bull. Jard. Bot. Buitenzorg, III, 4: 175 (1922). Homalomena rubrovaginata Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 331 (1922). Homalomena rubrovaginata var. subpurpurea Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 332 (1922). Homalomena pygmaea forma longipes Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 333 (1922). Homalomena ellipticifolia Furtado, Gard. Bull. Straits Settlem. 10: 193 (1939). Homalomena repens Ridl., Fl. Malay Penins. 5: 106 (1925). Homalomena humilis var. parvula Ridl., Fl. Malay Penins. 5: 109 (1925). Homalomena humilis var. ovatifolia M. Hotta, Acta Phytotax. Geobot. 22: 153 (1967). Distribution: Sumatera, Peninsular Malaysia, Java, Borneo, and Sulawesi. Homalomena kiahii Furtado, Gard. Bull. Straits Settlem. 10: 207 (1939). Homalomena kiahii f. maculata Furtado, Gard. Bull. Straits Settlem. 10: 207 (1939). Distribution: Peninsular Malaysia (Kelantan). Notes: Provisionally accepted.

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Homalomena lancifolia Hook.f., Fl. Brit. India 6: 533 (1893). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena major Griff., Not. Pl. Asiat. 3: 153 (1851). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena minor Griff., Not. Pl. Asiat. 3: 153 (1851). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena montana Furtado, Gard. Bull. Straits Settlem. 10: 210 (1939). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena nutans Hook.f., Fl. Brit. India 6: 535 (1893). Distribution: Peninsular Malaysia (Kelantan). Notes: Provisionally accepted. Homalomena pineodora Sulaiman & P.C. Boyce, Gard. Bull. Singapore 57: 8 (2005). Distribution: Peninsular Malaysia (Perak). Endemic. Homalomena pontederifolia Griff. ex Hook.f., Fl. Brit. India 6: 533 (1893). Distribution: Southern Peninsular Thailand to Peninsular Malaysia.

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Homalomena rostrata Griff., Not. Pl. Asiat. 3: 154 (1851) Cyrtocladon sanguinolentum Griff., Not. Pl. Asiat. 3: 147 (1851); Chamaecladon sanguinolentum (Griff.) Schott, Prodr. Syst. Aroid.: 316 (1860). Homalomena sagittifolia Jungh. ex Schott, Prodr. Syst. Aroid.: 311 (1860). Homalomena miqueliana Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 126 (1863). Homalomena propinqua Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 280 (1864). Homalomena beccariana Engl., Bull. Soc. Tosc. Ortic. 4: 296 (1879). Homalomena paludosa Hook.f., Fl. Brit. India 6: 531 (1893). Homalomena sagittifolia var. pontederiifolia Ridl., J Straits Branch Roy. Asiat. Soc. 44: 172 (1905). Homalomena ridleyana Engl., Bot. Jahrb. Syst. 37: 123 (1905). Homalomena teysmannii Engl., Pflanzenr., IV, 23Da: 68 (1912). Homalomena raapii Engl. Pflanzenr., IV, 23Da: 73(1912). Homalomena triangularis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 181 (1922). Homalomena sagittifolia var. sumatrana Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 192 (1922). Homalomena ensiformis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 335 (1922). Homalomena miqueliana var. truella Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 336 (1922). Homalomena sagittifolia var. angustifolia Furtado, Gard. Bull. Straits Settlem. 10: 228 (1939). Distribution: Southern Peninsular Thailand, Sumatera, Peninsular Malaysia, Borneo, and Maluku (Pulau Ceram).

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Homalomena scortechinii Hook.f., Fl. Brit. India 6: 534 (1893).

Homalomena crassa Ridl., J. Bot. 40: 36 (1902).

Distribution: Peninsular Malaysia. Endemic. Homalomena singaporensis Regel, Index Seminum (LE) 1869: 18 (1869). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Homalomena truncata (Schott) Hook.f., Fl. Brit. India 6: 535 (1893). Chamaecladon truncatum Schott, Bonplandia (Hannover) 6: 369 (1858) Distribution: Southern Myamnar, South Peninsular Thailand, and Peninsular Malaysia. Notes: Provisionally accepted. Homalomena wallichii Schott, Bonplandia (Hannover) 7: 30 (1859). Distribution: Peninsular Malaysia. Notes: Provisionally accepted. Landoltia Les & D.J. Crawford Landoltia Les & D.J. Crawford, Novon 9: 532 (1999, publ. 2000). Taxonomy: One species, widespread in the tropics and subtropics of both hemispheres. Most recent account: Landolt (1986); Les & Crawford (1999, publ. 2000) Ecology: Slow-moving or static water in old monsoon drains and road-side ditches, rare. Notes: Landoltia was separated from Spirodela primarily on the basis of molecular studies of the duckweeds aimed at providing additional data for undertaking taxonomic studies in a group with so few macromorphological data.

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In comparison with Spirodela, Landoltia plants exhibit a reduction in overall dimensions, number of roots and veins per plant, and have scattered brown epidermal pigment cells (visible on dead plants of Landoltia punctata), that are lacking in Spirodela. Landoltia punctata (G.Mey.) Les & D.J. Crawford, Novon 9: 532 (1999 publ. 2000). Lemna punctata G.Mey., Prim. Fl. Esseq.: 262 (1818). Spirodela punctata (G.Mey.) C.H. Thomps., Rep. (Annual) Missouri Bot. Gard. 1898: 28 (1898). Lemna melanorrhiza F. Muell. ex Kurz, J. Bot. 5: 115 (1867). Spirodela oligorrhiza var. melanorrhiza (F. Muell. ex Kurz) Hegelm., Lemnac.: 148 (1868). Lemna oligorrhiza Kurz, J. Linn. Soc., Bot. 9: 267 (1867). Spirodela oligorrhiza (Kurz) Hegelm., Lemnac.: 147 (1868). Lemna pleiorrhiza F. Muell. ex Kurz, J. Bot. 5: 115 (1867). Spirodela oligorrhiza var. pleiorrhiza (F. Muell. ex Kurz) Hegelm., Lemnac.: 149 (1868). Lemna javanica F.A. Bauer ex Hegelm., Lemnac.: 150 (1868). Lemna pusilla Hegelm., Lemnac.: 147 (1868). Spirodela oligorrhiza var. javanica Hegelm., Lemnac.: 150 (1868). Spirodela oligorrhiza var. pusilla Hegelm., Lemnac.: 149 (1868). Distribution: Throughout the tropics and subtropics. Lasia Lour. Lasia Lour., Fl. Cochinch.: 81 (1790). (Plate 17)

Lasius Hassk., Cat. Hort. Bot. Bogor.: 59 (1844).

Taxonomy: Two species; one (L. spinosa) widespread in tropical Asia from India to New Guinea, the other (L. concinna) endemic on Borneo. Most recent account: Hay (1988).

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Plate 17. Lasia Lour. A-D. Lasia spinosa (L.) Thwaites. The greatly elongated spathe (B), very short spadix with perigoniate flowers (C), and spinulous fruits (D) are diagnostic. Spathe colour varies from rich purple (as here) to dirty yellow. Leaf shape is also highly variable with large, vigorous plants always with the leaf blade pinnatipartite to pinnatisect, whereas smaller, weaker plants tend to have hastate leaves.

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Ecology: Plants of lowland freshwater swamp forest margins, sometimes persisting in seasonally flooded pasture and padi where they are often maintained as a vegetable – paku longkan – with the young shoots being picked and fried with belacan (shrimp paste). Notes: Lasia is a genus of clumping or colonial often stoloniferous herbs with a thick prickly creeping or unarmed erect stem. The prickly petioles carry either sagittate to hastate-sagittate (the latter mainly in juveniles) to deeply simply to 4× pinnatifid leaves. The spathe is notably thick and somewhat spongy and either twisted and opening basally (L. spinosa (L.) Twaites) or straight and opening terminally (L. concinna Alderw.). Lasia spinosa is widespread and often common in Peninsular Malaysia, as well as being variable in habit, from erect and clump-forming to somewhat decumbent and colonial via the production of long stolons, with leaves varying from simple hastate to pinnatifid. By comparison L. concinna appears to be restricted to the Kupuas river valley in Kalimantan (Indonesian Borneo) and with the widely separated populations remarkably morphologically stable. Lasia spinosa (L.) Thwaites, Enum. Pl. Zeyl.: 336 (1864). Dracontium spinosum L., Sp. Pl.: 967 (1753). Pothos spinosus (L.) Buch.-Ham. ex Wall., Numer. List: 4447 C (1831). Lasia aculeata Lour., Fl. Cochinch.: 81 (1790). Pothos heterophyllus Roxb., Fl. Ind. 1: 457 (1820). Lasia heterophylla (Roxb.) Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). Pothos lasia Roxb., Fl. Ind. 1: 458 (1820). Lasia loureiroi Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832), nom. illeg. Lasia roxburghii Griff., Not. Pl. Asiat. 3: 155 (1851). Lasia hermannii Schott, Bonplandia (Hannover) 5: 125 (1857). Lasia jenkinsii Schott, Bonplandia (Hannover) 5: 125 (1857). Lasia zollingeri Schott, Bonplandia (Hannover) 5: 125 (1857). Lasia desciscens Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 127 (1863).

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Lasia crassifolia f. angustisecta Engl., Exsicc. (Arac.): 194 (1883). Lasia crassifolia f. latisecta Engl., Exsicc. (Arac.): 194 (1883). Lasia crassifolia Engl., Bot. Jahrb. Syst. 25: 15 (1898). Distribution: Throughout tropical and subtropical Asia.

Lemna L. Lemna L., Sp. Pl.: 970 (1753). (Plate 18) Lenticularia Ség., Pl. Veron. 3: 129 (1754). Lenticula P.Micheli ex Adans., Fam. Pl. 2: 471 (1763). Hydrophace Hallier, Hist. Stirp. Helv. 3: 68 (1768). Telmatophace Schleid., Linnaea 13: 391 (1839). Staurogeton Rchb., Deut. Bot. Herb.-Buch: 33 (1841). Thelmatophace Godr., Fl. Lorraine 3: 18 (1844). Lenticularia P.Micheli ex Montandon, Guide Bot.: 308 (1868). Taxonomy: Thirteen species, widespread in the tropics, subtropics and warm temperate regions of both hemispheres. Three species in Peninsular Malaysia. Most recent account: Landolt (1986). Ecology: Slow-moving or static water in old monsoon drains and road-side ditches Lemna aequinoctialis Welw., Apont.: 578 (1859). Lemna angolensis Welw., J. Bot. 3: 112 (1865). Lemna perpusilla var. trinervis Austin in A. Gray, Manual, ed. 5: 479 (1867). Lemna trinervis (Austin) Small, Fl. S.E. U.S.: 230 (1903). Lemna paucicostata Hegelm., Lemnac.: 138 (1868).

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Plate 18. Lemnoideae. A. Lemna aequnoctialis Welw. B. Spirodela polyrrhiza (L.) Schleid. Note the multiple roots emerging from each plant body and the reddish colouration. C. Minute Wolffia globosa (Roxb.) Hartog & Plas surrounding Spirodela polyrrhiza. D. Lemna minor L.

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Lemna paucicostata var. membranacea Hegelm., Lemnac.: 141 (1868). Lemna minima Blatt. & Hallb., J. Indian Bot. 2: 50 (1921), nom. illeg. Lemna blatteri McCann, J. Bombay Nat. Hist. Soc. 42: 153 (1942). Lemna eleanorae McCann, J. Bombay Nat. Hist. Soc. 42: 154 (1942). Lemna leiboensis M.G. Liu & C.H. Hou, J. Southwest. Agric. Coll. (Chongqing) 1983(4): 58 (1983). Lemna aoukikusa T. Beppu & Murata, Acta Phytotax. Geobot. 36: 55 (1985). Lemna aoukikusa subsp. hokurikuensis T. Beppu & Murata, Acta Phytotax. Geobot. 36: 56 (1985). Distribution: Throughout the tropics and subtropics. Lemna minor L., Sp. Pl.: 970 (1753). Lenticula minor (L.) Scop., Fl. Carniol., ed. 2, 2: 213 (1772). Hydrophace minor (L.) Bubani, Fl. Pyren. 4: 23 (1901). Lenticula palustris Garsault, Fig. Pl. Méd.: t. 336 (1764), opus utique oppr. Lemna palustris Haenke ex Mert. & W.D.J. Koch in J.C. Röhling, Deutschl. Fl., ed. 3, 1: 295 (1823). Lenticula vulgaris Lam., Fl. Franç. 2: 189 (1779). Lemna vulgaris (Lam.) Lam., Encycl. 3: 464 (1792). Lemna minima Thuill. ex P.Beauv., J. Phys. Chim. Hist. Nat. Élément 82: 113 (1816). Lemna cyclostasa Elliott ex Chev., Fl. Gén. Env. Paris 2: 256 (1827). Lenticula cyclostasa (Elliott ex Chev.) Kurz, J. Linn. Soc., Bot. 9: 266 (1867). Lemna minor var. minima Chev., Fl. Gén. Env. Paris 2: 256 (1827). Lenticula minima (Chev.) Kurz, J. Linn. Soc., Bot. 9: 266 (1867). Lemna obcordata Bojer, Hortus Maurit.: 358 (1837), nom. inval. Lemna conjugata Willd. ex Schleid., Linnaea 13: 391 (1839), nom. inval. Lemna ovata A.Br. ex C.Krauss, Flora 28: 344 (1845). Lemna minor var. oxymitra Hegelm., Lemnac.: 143 (1868).

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Lemna monorhiza Montandon, Guide Bot.: 308 (1868). Lenticularia monorhiza Montandon, Guide Bot.: 308 (1868). Lemna rwandensis De Sloover, Bull. Jard. Bot. Natl. Belg. 43: 366 (1973). Distribution: Cosmopolitan. Lemna tenera Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40(2): 78 (1871). Staurogeton tener (Kurz) Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 84(7): 114 (1999). Distribution: Peninsular Malysia, through Indo-China to northern Australia.

Piptospatha N.E.Br. Piptospatha N.E.Br., Gard. Chron., n.s., 1879(1): 138 (1879). (Plate 19) Rhynchopyle Engl., Bot. Jahrb. Syst. 1: 183 (1881). Gamogyne N.E.Br., J. Bot. 20: 195 (1882). Taxonomy: Eight species, two in Peninsular Malaysia and southern Thailand, six endemic on Borneo. Most recent account: Ridley (1925). Ecology: All are obligate rheophytes along streams and by waterfalls in lowland to lower montane perhumid to everwet tropical broadleaf forest. Most species are obligate to specific geologies, notably granite in Peninsular Malaysia. Notes: Piptospatha is defined by the shedding spathe limb, erect splash cup, parietal placentation, fruits either fused into a syncarpium (most) or free but cohering (P. insignis N.E.Br.), and the presence of micropylar appendage on the seed. The generally pink, nodding inflorescences are diagnostic although similar inflorescences are found in Bornean Ooia S.Y. Wong and P.C. Boyce. Piptospatha perakensis (Engl.) Ridl., Fl. Malay Penins. 5: 114 (1925). Piptospatha elongata var. perakensis Engl., Pflanzenr., IV, 23Da: 125 (1912). Rhynchopyle perakensis (Engl.) Ridl., J. Bot. 51: 202 (1913).

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Plate 19. Piptospatha N.E.Br. A & B. Piptospatha perakensis (Engl.) Ridl. The plain leaves with conspicuous primary lateral veins, and white inflorescences smelling powerfully of jasmine (methyl jasmonate?), are immediately diagnostic. C & D. Piptospatha ridleyi N.E.Br. ex Hook.f.. Endemic to the Peninsula. The blotched leaves are distinctive (although plain leaves also occur in some populations) as is the pink and green striped spathe. A–D © Ooi Im Hin. Used with permission.

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Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Piptospatha ridleyi N.E.Br. ex Hook.f., Bot. Mag. 121: t. 7410 (1894). Distribution: Peninsular Malaysia ( Johor, Pahang). Endemic.

Pistia L. Pistia L., Sp. Pl.: 963 (1753). (Plate 20) Kodda-Pail Adans., Fam. Pl. 2: 25 (1763). Zala Lour., Fl. Cochinch.: 405 (1790). Apiospermum Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 351 (1853). Limnonesis Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 24 (1853). Taxonomy: One species occurring throughout the tropics and subtropics. Most recent account: Bogner & Hay (2000). Ecology: Floating aquatic on slow moving or still water, often in roadside ditches and old monsoon drains, and often pestilential. Notes: The spathe is highly modified in Pistia, externally pubescent and glabrous within, with a central constriction and the lower spathe margins connate with each other and adnate with ovary wall thus forming tube, and free margins between tube and limb folded between stigma and staminate flower, forming a partition between an upper male partial chamber and a lower female one. The spadic is mostly adnate to spathe, with only the apical staminate flower zone free. The pistillate flower zone bears a single pistil at the base and has a thin, green, pouch-shaped flap just below spathe partition, while the staminate flower zone is subtended by a thin, marginally lobed, green, annular flap, composed of a basally naked spadix axis supporting a single whorl of 2–8 flowers. Pistia stratiotes L., Sp. Pl.: 963 (1753).

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Plate 20. Pistia. L. A & B. Pistia stratiotes L. Immediately recognizable as a species, but not obviously as an aroid. The inflorescences appear deep between the leaves (B).

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Zala asiatica Lour., Fl. Cochinch.: 405 (1790). Pistia spathulata Michx., Fl. Bor.-Amer. 2: 162 (1803). Pistia stratiotes var. spathulata (Michx.) Engl. in C.F. P.von Martius & auct. suc. (eds.), Fl. Bras. 3(2): 214 (1878). Pistia crispata Blume, Rumphia 1: 78 (1836). Pistia leprieuri Blume, Rumphia 1: 79 (1836). Pistia linguiformis Blume, Rumphia 1: 79 (1836). Pistia minor Blume, Rumphia 1: 78 (1836). Pistia occidentalis Blume, Rumphia 1: 79 (1836). Pistia aegyptiaca Schleid., Allg. Gartenzeitung 6: 19 (1838). Pistia commutata Schleid., Allg. Gartenzeitung 6: 20 (1838). Limnonesis commutata (Schleid.) Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 24 (1853). Pistia obcordata Schleid., Allg. Gartenzeitung 6: 20 (1838). Apiospermum obcordatum (Schleid.) Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 351 (1853). Pistia stratiotes var. obcordata (Schleid.) Engl. in C.F.P. von Martius & auct. suc. (eds.), Fl. Bras. 3(2): 214 (1878). Pistia horkeliana Miq., Linnaea 18: 81 (1845). Pistia africana C. Presl, Epimel. Bot.: 240 (1851). Pistia amazonica C. Presl, Epimel. Bot.: 240 (1851). Pistia weigeltiana C. Presl, Epimel. Bot.: 240 (1851). Pistia turpinii K. Koch, Bot. Zeitung (Berlin) 10: 577 (1852). Limnonesis friedrichsthaliana Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 24 (1853). Pistia aethiopica Fenzl ex Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 354 (1853). Pistia brasiliensis Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 356 (1853). Pistia cumingii Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 354 (1853).

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Pistia gardneri Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 356 (1853). Pistia natalensis Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 354 (1853). Pistia schleideniana Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 356 (1853). Pistia texensis Klotzsch, Abh. Königl. Akad. Wiss. Berlin 1853: 356 (1853). Pistia stratiotes var. linguiformis Engl. in C.F. P. von Martius & auct. suc. (eds.). Distribution: Throughout the tropics and subtropics.

Podolasia N.E.Br. Podolasia N.E.Br., Gard. Chron., n.s., 1882(2): 70 (1882). (Plate 21) Taxonomy: One species occurring disjunctly in Sumatera, Peninsular Malaysia, and Borneo. Most recent account: Hay (1988). Ecology: Lowland peat swamp forest, usually on slightly raised peat deposits (podzols) but still subject to seasonal inundation. Notes: The richly coloured spathe and spadix becoming bicoloured as staminate anthesis progresses are diagnostic. The spadix begins white, and remains white through pistillate anthesis. At the onset of staminate anthesis (which proceeds from the tip to the base of the spadix) the individual flowers turn to deep purple as staminate anthesis proceeds. By the end of anthesis and throughout fruit development, the spadix is deep purple. Podolasia may be distinguished vegetatively by the short, distinct, unarmed internodes, the persistent leaf bases (often decaying into tough fibres), though which adventitious roots emerge, and by the downward-pointing petiole prickles. The fruits are also characteristic, being relatively very large (their length considerably exceeding the diameter of the spadix), individually distinct berries exserted well clear of the tepals. Podolasia stipitata is very sporadic in distribution throughout its range, although it is frequently locally abundant.

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Plate 21. Podolasia N.E.Br. A–B. Podolasia stipitata N.E.Br. A. Flowering and fruiting plant in habitat. B. Inflorescence at staminate anthesis. Note the terminal portion of the spadix is differently coloured, this zone comprises flowers that are post anthesis and will proceed down the spadix until post anthesis the entire spadix is purple. C. Detail of the spadix base near to completion of anthesis. Note that only a small area of white spadix remains. D. Infructescence nearing maturity. Note that the fruits are almost wholly exserted from between the tepals. When ripe the berries are scarlet.

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Podolasia stipitata N.E.Br., Gard. Chron., n.s., 1882(2): 70 (1882). Lasia stipitata N.E.Br., Gard. Chron., n.s., 1882(2): 70 (1882), pro syn. Cyrtosperma angustilobum Engl., Pflanzenr., IV, 23C: 21 (1911). Distribution: Sumatera, Peninsular Malaysia, and Borneo.

Pothos L. Pothos L., Sp. Pl.: 968 (1753). (Plate 22) Tapanava Adans., Fam. Pl. 2: 470 (1763). Batis Blanco, Fl. Filip.: 791 (1837). Goniurus C.Presl, Epimel. Bot.: 244 (1851). Taxonomy: Approximately 70 species distributed from Madagascar through to India, the subtropical eastern Himalayas, throughout subtropical and tropical Asia into western Oceania and tropical eastern Australia. Nine species in Peninsular Malaysia, of which one is endemic. Most recent account: Boyce & Hay (2001). Ecology: Primary to disturbed secondary lowland to upper hill perhumid evergreen broadleaf forest, often on slopes or ridges, less often in poorly drained valley bottoms. Some species (i.e., P. kingii Hook.f., P. leptostachyrus Schott, P. macrocephalus Scort. ex Hook.f., and P. ovatifolius Merr.) are limestone obligates. Notes: Pothos is divided into two subgenera – Pothos and Allopothos, with Allopothos further divided into two informal supergroups – Allopothos and Goniurus. Subgenus Pothos is distinguishable by the blade-like petiole, lacking an obvious sheath with the whole leaf resembling that of many Citrus. Subgenus Allopothos has a distinct petiole, sheathing for much of its length. The Allopothos supergroup has a spadix with congested flowers while the flowers of the Goniurus supergroup are scattered, with portions of the spadix axis clearly visible between individual flowers. Species of the Goniurus supergroup (e.g., P. curtsiii Hook.f., and notably Bornean P. oliganthus P.C. Boyce & A. Hay – see Boyce & Hay, 2001) are of interest as they appear to link the genera Pothos and Bornean Pedicellarum. All three supergroups are present in Peninsular Malaysia.

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Plate 22. Pothos L. A. Pothos macrocephalus Scort. ex Hook.f. is unmistakeable in Peninsula Malaysia by the large size and bright colours of the inflorescences. It is most similar to species from Java (Pothos junghuhnii de Vriese) and S. Vietnam (P. gigantipes Buchet ex P.C. Boyce). B. Pothos leptostachyus Schott. A widespread species that appears to be restricted to limestone throughout its range. C–D. Pothos scandens. The strongly curving stipe is immediately diagnostic. The red fruits of the majority of Pothos (D) are mostly likely dispersed by birds, although there are no direct observations of this. A © Royal Forest Dept. Bangkok. Used with permission.

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Pothos barberianus Schott, Aroideae: 24 (1856). Distribution: Sumatera, Peninsular Malaysia, and Borneo. Pothos curtisii Hook.f., Fl. Brit. India 6: 554 (1893). Pothos kunstleri Hook.f., Fl. Brit. India 6: 554 (1893). Pothos latifolius Hook.f., Fl. Brit. India 6: 554 (1893), nom. illeg. Pothos peninsularis Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 381 (1920). Pothos maingayi Hook.f., Fl. Brit. India 6: 554 (1893). Distribution: Southern Peninsular Thailand, Sumatera, and Peninsular Malaysia. Pothos kingii Hook.f., Fl. Brit. India 6: 553 (1893). Pothos ellipticus Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 48 (1904), nom. illeg. Pothos ridleyanus Furtado, Gard. Bull. Straits Settlem. 8: 150 (1935). Pothos grandispathus Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 48 (1904). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Pothos lancifolius Hook.f., Fl. Brit. India 6: 554 (1893). Pothos penicilliger Gagnep., Notul. Syst. (Paris) 9: 135 (1941). Distribution: South Vietnam and Peninsular Malaysia. Pothos leptostachyus Schott, Prodr. Syst. Aroid.: 571 (1860). Pothos lorispathus Ridl., J. Straits Branch Roy. Asiat. Soc. 86: 310 (1922). Distribution: Southern Peninsular Thailand, Sumatera, Peninsular Malaysia, and Borneo.

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Pothos macrocephalus Scort. ex Hook.f., Fl. Brit. India 6: 553 (1893).

Southern Peninsular Thailand to Peninsular Malaysia and Sumatera.

Pothos ovatifolius Engl., Pflanzenr., IV, 23B: 40 (1905). Pothos merrillii K. Krause, Bot. Jahrb. Syst. 45: 657 (1911). Pothos ovatifolius var. simalurensis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 337 (1922). Distribution: Sumatera, Peninsular Malaysia, Borneo, and the Philippines. Pothos salicifolius Ridl. ex Burkill & Holttum, Gard. Bull. Straits Settlem. 3: 88 (1923). Distribution: Peninsular Malaysia. Endemic. Pothos scandens L., Sp. Pl.: 968 (1753). Batis hermaphrodita Blanco, Fl. Filip.: 791 (1837). Pothos hermaphroditus (Blanco) Merr., Sp. Blancoan.: 90 (1918). Podospadix angustifolia Raf., Fl. Tellur. 4: 124 (1838). Pothos angustifolius (Raf.) C. Presl, Epimel. Bot.: 243 (1851). Tapanava indica Raf., Fl. Tellur. 4: 14 (1838). Tapanava rheedei Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 165 (1842). Pothos leptospadix de Vriese in F.A.W. Miquel, Pl. Jungh.: 105 (1851). Pothos longifolius C. Presl, Epimel. Bot.: 242 (1851). Pothos microphyllus C. Presl, Epimel. Bot.: 243 (1851). Pothos roxburghii de Vriese in F.A.W. Miquel, Pl. Jungh.: 103 (1851). Pothos longipedunculatus Engl., Pflanzenr., IV, 23B: 27 (1905), nom. illeg. Pothos scandens var. sumatranus de Vriese in F.A.W. Miquel, Pl. Jungh.: 103 (1851). Pothos scandens var. zeylanicus de Vriese in F.A.W. Miquel, Pl. Jungh.: 103 (1851).

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Pothos chapelieri Schott, Oesterr. Bot. Wochenbl. 5: 19 (1855). Pothos zollingerianus Schott, Oesterr. Bot. Wochenbl. 5: 19 (1855). Pothos scandens var. zollingerianus (Schott) Engl., Pflanzenr., IV, 23B: 26 (1905). Pothos angustifolius Reinw. ex Miq., Bot. Zeitung (Berlin) 14: 561 (1856), nom. illeg. Pothos cognatus Schott, Aroideae: 22 (1856). Pothos scandens var. cognatus (Schott) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 84 (1879). Pothos exiguiflorus Schott, Aroideae: 21 (1856). Pothos horsfieldii Miq., Fl. Ned. Ind. 3: 178 (1856). Pothos decipiens Schott, Bonplandia (Hannover) 7: 165 (1859). Pothos fallax Schott, Prodr. Syst. Aroid.: 560 (1860). Pothos zollingeri Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 85 (1879), orth. var. Pothos scandens f. angustior Engl., Bot. Tidsskr. 24: 272 (1902). Pothos scandens var. helferianus Engl., Pflanzenr., IV, 23B: 26 (1905). Distribution: Madagascar through to Philippines, as far north as western China (southern Sichuan).

Rhaphidophora Hassk. Rhaphidophora Hassk., Flora 25(2 Beibl. 1): 11 (1842). (Plate 23) Raphidophora Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 168 (1842), orth. var. Afrorhaphidophora Engl. in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam., Nachtr. 3: 31 (1906). Taxonomy: About 100 species occurring from wet tropical Africa, throughout tropical south and South East Asia, perhumid and everwet subtropical and tropical Australia, and into western Oceania with extensions into the subtropical

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Plate 23. Rhaphiodophora Hassk. A & B. Rhaphidophora korthalsii Schott. A species with adult (A) and juvenile (B) growth that look completely different. While the adult pant has large leaves (sometime more than 2 m long including the petiole) with the leaf blade pinnatisect, the juvenile stage produces leaves that overlap in the manner or roof shingles. C & D. Rhaphidophora lobbii Schott. C. Inflorescence at late pistillate anthesis. Note that that the spathe is beginning to fall. D. Ripe infructescence with the stylar plates starting to slough away. This infructescence type is termed a monsterocarp.

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Himalaya, southern China, and the southernmost islands of Japan. In Peninsular Malaysia there are 18 species of which three are endemic. Most recent account: Boyce (1999); Othman & Boyce (2010). Ecology: Usually in well drained subtropical and tropical perhumid to everwet broadleaf forest at low to mid-montane elevations, less often in peat swamp (R. lobbii Schott) or freshwater swamp tropical forest (R. minor Hook.f.). Notes: Revisionary work for Peninsular Malaysia (Boyce, 1999) recorded 15 species in the Peninsula, with two endemic. Since then one species has been resurrected (R. burkillina Ridl.) and there are two new records (hitherto Bornean endemic R. latevaginata M.Hotta and R. megasperma Engl.). Thus Peninsular Malaysia 18 species comprising three endemic species, two species otherwise only in Sumatera, two species known also from southern Thailand and 11 species with distributions extending variously to Borneo, the Philippines and western Oceania. However, much remains to be done, notably with the squarestemmed Hongkongensis Group, which undoubtedly comprises a considerable number of undescribed species. Rhaphidophora angustata Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 128 (1863). Scindapsus pteropodus Teijsm. & Binn., Natuurw. Tijdschr. Ned.-Indië 27: 23 (1864). Rhaphidophora pteropoda (Teijsm. & Binn.) Engl., Bull. Soc. Tosc. Ortic. 4: 268 (1879). Rhaphidophora laetevirens Ridl., J. Bot. 40: 37 (1902). Distribution: Sumatera and Peninsular Malaysia. Rhaphidophora beccarii (Engl.) Engl., Bot. Jahrb. Syst. 1: 181 (1881). Epipremnum beccarii Engl., Bull. Soc. Tosc. Ortic. 4: 269 (1879). Rhaphidophora borneensis Engl., Bot. Jahrb. Syst. 7(15): 1 (1886). Rhaphidophora fluminea Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 186 (1905). Distribution: Southern Peninsular Thailand to Peninsular Malaysia, Sumatera, and Borneo.

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Rhaphidophora burkilliana Ridl., Fl. Malay Penins. 5: 121 (1925). Distribution: Peninsular Malaysia (Selangor). Endemic. Rhaphidophora corneri P.C.Boyce, Gard. Bull. Singapore 51: 205 (1999). Distribution: Peninsular Malaysia (Terengganu). Endemic. Rhaphidophora crassifolia Hook.f., Fl. Brit. India 6: 543 (1893). Distribution: Southern Peninsular Thailand to Peninsular Malaysia, and Borneo (Sarawak). Rhaphidophora falcata Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 198 (1905). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Rhaphidophora foraminifera (Engl.) Engl., Pflanzenr., IV, 23B: 45 (1908). Epipremnum foraminiferum Engl., Bot. Jahrb. Syst. 25: 11 (1898). Distribution: Sumatera, Peninsular Malaysia, and Borneo. Rhaphidophora korthalsii Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 129 (1863). Rhaphidophora maxima Engl., Bull. Soc. Tosc. Ortic. 4: 269 (1879). Pothos celatocaulis N.E.Br., Gard. Chron., n.s., 1880(1): 200 (1880). Rhaphidophora celatocaulis (N.E.Br.) Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 382 (1920). Scindapsus anomalus Carrière, Rev. Hort. 1884: 536 (1884), prov. syn. Pothos bifarius Wall. ex Hook.f., Fl. Brit. India 6: 555 (1893). Rhaphidophora copelandii Engl., Bot. Jahrb. Syst. 37: 115 (1905). Rhaphidophora grandis Ridl., J. Straits Branch Roy. Asiat. Soc. 49: 51 (1908), nom. illeg. Rhaphidophora ridleyi Merr., J. Straits Branch Roy. Asiat. Soc. Sped. ed.: 90 (1921).

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Rhaphidophora grandifolia K. Krause, Bot. Jahrb. Syst. 44(101): 11 (1910). Rhaphidophora trinervia Elmer, Leafl. Philipp. Bot. 8: 3073 (1919). Rhaphidophora latifolia Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 341 (1922). Rhaphidophora palawanensis Merr., Philipp. J. Sci. 26: 451 (1925). Rhaphidophora trukensis Hosok., J. Jap. Bot. 13: 195 (1937). Epipremnum multicephalum Elmer, Leafl. Philipp. Bot. 10: 3624 (1938), no latin descr. Distribution: Southern Thailand through to western Oceania. Rhaphidophora latevaginata M. Hotta, Acta Phytotax. Geobot. 22: 4 (1966) Distribution: Peninsular Malaysia and Borneo. Rhaphidophora lobbii Schott, Prodr. Syst. Aroid.: 379 (1860). Scindapsus lobbii (Schott) Ender, Index Aroid.: 74 (1864). Distribution: Peninsular Thailand through to West and central Malesia. Rhaphidophora maingayi Hook.f., Fl. Brit. India 6: 543 (1893). Rhaphidophora apiculata Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 383 (1920), nom. illeg. Distribution: Southern Peninsular Thailand to Peninsular Malaysia, and Sumatera. Rhaphidophora megasperma Engl., Bot. Jahrb. Syst. 25: 8 (1881). Scindapsus havilandii Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 184 (1905). Rhaphidophora jaculiformis Alderw., Bull. Jard. Bot. Buitenzorg III, 4: 197 (1922). Rhaphidophora subfalcata M. Hotta, Acta Phytotax. Geobot. 22: 6 (1966). Distribution: Peninsular Malaysia and Borneo.

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Rhaphidophora minor Hook.f., Fl. Brit. India 6: 544 (1893). Rhaphidophora celebica K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem 11: 331 (1932). Distribution: Southern Peninsular Thailand through to West and central Malesia. Rhaphidophora montana (Blume) Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 128 (1863). Calla montana Blume, Catalogus: 62 (1823). Scindapsus montanus (Blume) Kunth, Enum. Pl. 3: 64 (1841). Anadendrum montanum (Blume) Schott, Bonplandia (Hannover) 5: 45 (1857). Scindapsus angulatus Miq., Fl. Ned. Ind. 3: 219 (1856). Rhaphidophora angulata (Miq.) Schott, Prodr. Syst. Aroid.: 379 (1860). Rhaphidophora fallax Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 128 (1863). Distribution: Sumatera and Peninsular Malaysia. Rhaphidophora nicolsonii P.C. Boyce, Gard. Bull. Singapore 51: 236 (1999). Distribution: Peninsular Malaysia (Pahang). Endemic. Rhaphidophora puberula Engl., Bot. Jahrb. Syst. 1: 180 (1881). Rhaphidophora gracilipes Hook.f., Fl. Brit. India 6: 545 (1893). Rhaphidophora kunstleri Hook.f., Fl. Brit. India 6: 546 (1893). Rhaphidophora scortechinii Hook.f., Fl. Brit. India 6: 545 (1893). Rhaphidophora batoeensis Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23B: 27 (1908). Rhaphidophora hallieri Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 385 (1920). Rhaphidophora pilosula Alderw., Bull. Jard. Bot. Buitenzorg, III, 1: 386 (1920). Rhaphidophora scaberula Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 196 (1922).

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Distribution: Sumatera, Peninsular Malaysia, Nusa Tenggara, and Borneo (Sabah). Rhaphidophora sylvestris (Blume) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 239 (1879). Calla sylvestris Blume, Catalogus: 62 (1823). Scindapsus sylvestris (Blume) Kunth, Enum. Pl. 3: 64 (1841). Scindapsus angustifolius Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 164 (1842), nom. illeg. Rhaphidophora angustifolia Schott, Bonplandia (Hannover) 5: 45 (1857), nom. illeg. Scindapsus lingulatus Hassk., Cat. Hort. Bot. Bogor.: 58 (1844). Rhaphidophora lingulata (Hassk.) Schott, Bonplandia (Hannover) 5: 45 (1857). Monstera lingulata K. Koch ex Endler, Index Aroid.: 74 (1864), nom. inval. Scindapsus aruensis Engl., Bull. Soc. Tosc. Ortic. 4: 270 (1879). Rhaphidophora wrayi Hook.f., Fl. Brit. India 6: 544 (1893). Rhaphidophora gratissima Becc., For. Borneo: 604 (1902). Rhaphidophora nigrescens Ridl., J. Straits Branch Roy. Asiat. Soc. 44: 185 (1905). Rhaphidophora motleyana Engl. & K. Krause in H.G.A. Engler, Pflanzenr., IV, 23B: 25 (1908). Scindapsus fragilis Elmer, Leafl. Philipp. Bot. 10: 3702 (1939), no latin descr. Distribution: Sumatera through to New Guinea (Kepulauan Aru). Rhaphidophora tetrasperma Hook.f., Fl. Brit. India 6: 548 (1893). Distribution: Southern Peninsular Thailand and Peninsular Malaysia.

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Schismatoglottis Zoll. & Moritzi Schismatoglottis Zoll. & Moritzi in H. Zollinger, Syst. Verz.: 83 (1854). (Plate 24)

Apatemone Schott, Gen. Aroid.: t. 57 (1858).

Colobogynium Schott, Oesterr. Bot. Z. 15: 34 (1865).

Nebrownia Kuntze, Revis. Gen. Pl. 2: 742 (1891).

Taxonomy: In excess of 150 species, ranging from Sumatera to New Guinea and as far north as southwestern China, generally in wetter areas. Five species in Peninsular Malaysia, one or perhaps two endemic. Most recent account: Hay & Yuzammi (2000). Ecology: Schismatoglottis are variously terrestrial, often on steep slopes, or lithophytic, rheophytic or rarely chasmophytic (the last three not in Peninsular Malaysia), in lowland to lower montane perhumid to everwet evergreen forest. Notes: The Schismatoglottideae is a diverse group of terrestrial, rheophytic, lithophytic or chasmophytic rainforest herbs centred on Borneo. Schismatoglottis Zoll. & Moritzi is by far the largest genus, extending throughout Malesia to western Oceania, north through much of Indochina and into South West China and Taiwan. The remaining genera are relatively small to monospecific and with the exception of Hestia, Apoballis, and Piptospatha are endemic on Borneo. Schismatoglottis is defined by a constricted spathe, staminate flowers lacking thecae horns, seeds without a micropylar appendage, and parietal placentation. Growth ranges from stoloniferous, colonial helophytes to clumping or solitary rheophytes and lithophytes, with stems variously elongated and creeping to erect or even weakly climbing, to clumping and congested. Schismatoglottis brevicuspis Hook.f., Fl. Brit. India 6: 537 (1893). Distribution: Sumatera and Peninsular Malaysia. Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi in H. Zollinger, Syst. Verz.: 83 (1854). Calla calyptrata Roxb., Fl. Ind. ed. 1832, 3: 514 (1832). Homalomena calyptrata (Roxb.) Kunth, Enum. Pl. 3: 57 (1841). Zantedeschia calyptrata (Roxb.) K. Koch, Index Seminum (B) 1854 (App.): 9 (1855).

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Plate 24. Schismatoglottis Zoll. & Moritzi A & B. Schismatoglottis calyptrata (Roxb.) Zoll & Moritzi. A colonial species with underground stolons (A). As typical for most Schismatoglottis the spathe limb is shed during anthesis. In this species the limb is shed while still fresh. C & D. Schismatoglottis scortechinii Hook.f. A species forming loose clumps by creeping stems. In this group of species the spathe limb degrades, often liquefying and later drying on the spadix. E. Schismatoglottis brevicuspis Hook.f. An unusual species with long, creeping stems that root as they proceed. It is the only species in Peninsular Malaysia of a group otherwise restricted to Borneo.

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Colocasia humilis Hassk., Flora 25(2 Beibl. 1): 10 (1842). Colocasia humilis Hassk., Cat. Hort. Bot. Bogor.: 56 (1844), nom. illeg. Schismatoglottis longipes Miq., Fl. Ned. Ind. 3: 214 (1856). Schismatoglottis picta Schott, Oesterr. Bot. Z. 13: 317 (1863). Schismatoglottis riparia Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 281 (1864). Colocasia neoguineensis Linden ex André, Ill. Hort. 27: 68 (1880). Schismatoglottis neoguineensis (Linden ex André) N.E.Br., Gard. Chron., n.s., 1885(2): 776 (1885). Alocasia neoguineensis (Linden ex André) Sieber & Voss, Vilm. Ill. Blumengärtn., ed. 3, 1: 1170 (1895), prov. syn. Schismatoglottis variegata N.E.Br., Gard. Chron., n.s., 1885(2): 776 (1885). Schismatoglottis bitaeniata Engl., Bot. Jahrb. Syst. 37: 124 (1905). Schismatoglottis cordifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 57: 112 (1911). Schismatoglottis acutangula Engl., Pflanzenr., IV, 23Da: 110 (1912). Schismatoglottis calyptrata f. minor Engl., Pflanzenr., IV, 23Da: 116 (1912). Schismatoglottis calyptrata var. ornata Ridl. ex Engl., Pflanzenr., IV, 23Da: 116 (1912). Schismatoglottis djamuensis Engl., Bot. Jahrb. Syst. 49: 99 (1912). Schismatoglottis emarginata Engl., Pflanzenr., IV, 23Da: 93 (1912). Schismatoglottis hellwigiana Engl., Nova Guinea 8: 806 (1912). Schismatoglottis hellwigiana var. subcordata Engl., Pflanzenr., IV, 23Da: 102 (1912). Schismatoglottis nieuwenhuisii Engl., Bot. Jahrb. Syst. 48: 95 (1912). Schismatoglottis tenuifolia Engl., Nova Guinea 8: 807 (1912). Schismatoglottis klossii Ridl., Trans. Linn. Soc. London, Bot. 9: 239 (1916).

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Schismatoglottis dorensis Gibbs, Fl. Arfak Mts.: 201 (1917). Schismatoglottis angustifolia Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 344 (1922). Schismatoglottis calyptratoides Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 213 (1922). Schismatoglottis maculata Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 218 (1922). Schismatoglottis parvifolia Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 342 (1922). Schismatoglottis potamophila Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 343 (1922). Schismatoglottis pseudocalyptrata Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 212 (1922). Schismatoglottis ruttenii Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 211 (1922). Schismatoglottis sublaxiflora Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 222 (1922). Schismatoglottis vanvuurenii Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 221 (1922). Distribution: China (Guangxi) to Vanuatu. Schismatoglottis penangensis Engl., Pflanzenr., IV, 23Da: 88 (1912). Distribution: Peninsular Malaysia (Penang). Notes: Provisionally accepted. Schismatoglottis scortechinii Hook.f., Fl. Brit. India 6: 537 (1893). Schismatoglottis marginata Ridl., J. Bot. 40: 36 (1902), nom. illeg. Schismatoglottis kingii Engl., Nova Guinea 8: 806 (1912). Distribution: Peninsular Malaysia. Endemic.

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Schismatoglottis wallichii Hook.f., Fl. Brit. India 6: 537 (1893). Schismatoglottis wallichii var. oblongata Hook.f., Fl. Brit. India 6: 537 (1893). Schismatoglottis wallichii f. oblongata (Hook.f.) Engl., Pflanzenr., IV, 23Da: 100 (1912). Schismatoglottis wallichii var. fasciata Ridl., Mat. Fl. Malay. Penins. 3: 33 (1907). Schismatoglottis longicuspis Engl., Pflanzenr., IV, 23Da: 100 (1912). Schismatoglottis conversa Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 344 (1922). Distribution: Sumatera, Peninsular Malaysia, and Borneo.

Scindapsus Schott Scindapsus Schott in H.W. Schott & S.L. Endlicher, Melet. Bot.: 21 (1832). (Plate 25) Cuscuaria Schott, Bonplandia (Hannover) 5: 45 (1857). Taxonomy: As many as 60 species, with about half yet to be described, throughout tropical Asia from North East India though New Guinea to eastern Australia (Queensland). Seven species in Peninsular Malaysia but most poorly understood. Most recent account: Othman, Boyce & Chan (2010). Ecology: Mainly well-drained perhumid to everwet broadleaf forest at low to mid elevations, although S. scortechinii Hook.f. forms a distinctive feature of montane forest. Notes: There is no modern account for Scindapsus and many of the existing species are known from fragmentary collections. The plants are variable in appearance, including colossal high-climbing primary hemiepiphytes (e.g., S. perakensis Hook.f.), almost shrubby secondary hemiepiphytes (notably the S. coriaceus complex – in which many of the novel species occur), and the perching litter-trapping epiphytes of the S. beccarii complex.

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Plate 25. Scindapsus Schott A. Scindapsus pictus Hassk. The juvenile stage with distinctive shingling leaves. Adult flowering stems from curtains hanging from the canopy with sickle-shaped, unmarked leaves. B. Scindapsus lucens Bogner & P.C. Boyce. Closely related to S. pictus, but readily distinguished but the pale grey leaves with a conspicuous bullate surface, and by flowering on stems of similar morphology to the juvenile growth. C. Scindapsus scortechinii. A highland species with stiff, glossy green leaves. It is related to a group of species otherwise restricted to Borneo, and which in the main have yet to be formally described.

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Scindapsus beccarii Engl., Bot. Jahrb. Syst. 1: 182 (1881). Distribution: Sumatera and Peninsular Malaysia. Scindapsus hederaceus Miq., Fl. Ned. Ind. 3: 185 (1856). Pothos hederaceus Zoll. & Moritzi in H. Zollinger, Syst. Verz.: 83 (1854), nom. illeg. Scindapsus pothoides Miq., Fl. Ned. Ind. 3: 184 (1856). Scindapsus inquinatus Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 283 (1864). Scindapsus poilanei Gagnep., Notul. Syst. (Paris) 9: 139 (1941). Distribution: Southern Indo-China to Thailand, Sumatera, Peninsular Malaysia, Java, Borneo, and the Philippines. Scindapsus lucens Bogner & P.C. Boyce, Kew Bull. 49: 789 (1994). Distribution: Sumatera, Peninsular Malaysia. Scindapsus perakensis Hook.f., Fl. Brit. India 6: 542 (1893). Scindapsus longipetiolatus Ridl., Bull. Misc. Inform. Kew 1926: 93 (1926). Distribution: Sumatera, Peninsular Malaysia, and Java(?) Scindapsus pictus Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9: 164 (1842). Scindapsus pothoides Schott, Prodr. Syst. Aroid.: 394 (1860), nom. illeg. Pothos argyraeus Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 255 (1879), prov. syn. Scindapsus argyraeus Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 255 (1879). Scindapsus pictus var. argyraeus (Engl.) Engl., Bot. Jahrb. Syst. 25: 13 (1898). Pothos argenteus W. Bull, Cat. 1887: 11 (1887). Scindapsus pictus var. oblongifolius Engl., Bot. Jahrb. Syst. 25: 13 (1898).

The Checklist

Distribution: Sumatera, Peninsular Malaysia, Java, Borneo, Sulawesi, and the Philippines. Scindapsus scortechinii Hook.f., Fl. Brit. India 6: 541 (1893). Distribution: Southern Peninsular Thailand to Peninsular Malaysia. Scindapsus treubii Engl., Bot. Jahrb. Syst. 25: 13 (1898). Pothos enderianus N.E.Br., Gard. Chron., n.s., 1884(1): 711 (1884). Distribution: Peninsular Malaysia, Java, and Borneo.

Spirodela Schleid. Spirodela Schleid., Linnaea 13: 391 (1839). (Plate 18) Taxonomy: Three species: one cosmopolitan, one from South America, and one restricted to southern central China. One species occurring on Peninsular Malaysia. Most recent account: Landolt (1986). Ecology: Slow-moving or static water in old monsoon drains and road-side ditches, rare. Spirodela polyrrhiza (L.) Schleid., Linnaea 13: 392 (1839). Lemna polyrrhiza L., Sp. Pl.: 970 (1753). Lenticula polyrrhiza (L.) Lam., Fl. Franc. 2: 189 (1779). Telmatophace polyrrhiza (L.) Godr., Fl. Lorraine 3: 18 (1844). Lemna orbicularis Kit. ex Schult., Oestr. Fl., ed. 2, 1: 64 (1814). Telmatophace orbicularis (Kit. ex Schult.) Schur, Verh. Mitth. Siebenbürg. Vereins Naturwiss. Hermannstadt 4: 70 (1853). Lemna thermalis P.Beauv. ex Nutt., Gen. N. Amer. Pl. 1: 19 (1818). Lemna orbiculata Roxb., Fl. Ind. ed. 1832, 3: 565 (1832). Lemna bannatica Waldst. & Kit. ex Schleid., Linnaea 13: 392 (1839).

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Lemna major Griff., Not. Pl. Asiat. 3: 216 (1851). Lemna transsilvanica Schur, Enum. Pl. Transsilv.: 635 (1866), prov. syn. Lemna polyrrhiza var. concolor Kurz, J. Bot. 5: 115 (1867). Lemna umbonata A.Br. ex Hegelm., Lemnac.: 156 (1868). Spirodela atropurpurea Montandon, Guide Bot.: 309 (1868). Lemna maxima Blatt. & Hallb., J. Indian Bot. 2: 49 (1921). Spirodela maxima (Blatt. & Hallb.) McCann, J. Bombay Nat. Hist. Soc. 43: 158 (1942). Spirodela polyrrhiza var. masonii Daubs, Illinois Biol. Monogr. 34: 13 (1965). Distribution: Cosmopolitan.

Typhonium Schott Typhonium Schott, Wiener Z. Kunst 3: 72 (1829). Desmesia Raf., Fl. Tellur. 3: 63 (1837). Heterostalis (Schott) Schott, Oesterr. Bot. Wochenbl. 7: 261 (1857). Taxonomy: Perhaps more than 100 species occurring from India through much of subtropical and tropical Asia into southern Japan. Species diversity is greatest in areas with strongly seasonal (wet-dry) climates. Two of the five Peninsular Malaysian species are non-indigenous or possibly so, and occur only in association with human habitation where their presence in all possibility is the result of introduction as medicinal plants. Most recent account: Ridley (1925). Ecology: Indigenous species occur in seasonally dry evergreen to semideciduous forest on limestone. Typhonium filiforme Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 45 (1904). Distribution: Southern Thailand to Peninsular Malaysia.

The Checklist

Typhonium flagelliforme (Lodd.) Blume, Rumphia 1: 134 (1837). Arum flagelliforme Lodd., Bot. Cab. 4: t. 396 (1819). Heterostalis flagelliformis (Lodd.) Schott, Oesterr. Bot. Wochenbl. 7: 261 (1857). Arum divaricatum L., Sp. Pl.: 966 (1753). Arum cuspidatum Blume, Catalogus: 101 (1823). Typhonium cuspidatum (Blume) Decne., Nouv. Ann. Mus. Hist. Nat. 3: 367 (1834). Typhonium cuspidatum var. ptychiurum Blume, Rumphia 1: 134 (1837). Arum ptychiurum Zipp. ex Kunth, Enum. Pl. 3: 26 (1841). Typhonium sylvaticum Voigt, Hort. Suburb. Calcutt.: 686 (1845). Arum angulatum Griff., Not. Pl. Asiat. 3: 143 (1851). Arum flagelliferum Griff., Not. Pl. Asiat. 3: 144 (1851). Typhonium flagelliferum Griff., Not. Pl. Asiat. 3: 144 (1851). Typhonium hastiferum Miq., Fl. Ned. Ind. 3: 194 (1856). Typhonium reinwardtianum de Vriese & Miq. ex Miq., Fl. Ned. Ind. 3: 195 (1856). Typhonium flagelliforme var. angustissimum Ridl., J. Straits Branch Roy. Asiat. Soc. 59: 218 (1911). Typhonium incurvatum Blatt. & McCann, J. Bombay Nat. Hist. Soc. 35: 22 (1931). Distribution: North East India through Burma to South West China and south through Malesia to northern Australia. Notes: The natural distributional range is unclear as T. flagelliforme is a weedy species much utilized for medicine (keladi tikus). Typhonium fultum Ridl., J. Straits Branch Roy. Asiat. Soc. 41: 45 (1904). Distribution: Southern Thailand to Peninsular Malaysia.

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Typhonium roxburghii Schott, Aroideae 1: 12 (1853). Typhonium divaricatum var. roxburghii (Schott) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 612 (1879). Arum diversifolium Blume, Catalogus: 102 (1823). Typhonium divaricatum Blume, Nouv. Ann. Mus. Hist. Nat. 3: 367 (1834), nom. illeg. Dracunculus divaricatus Raf., Fl. Tellur. 3: 65 (1837). Typhonium divaricatum var. robustum Blume, Rumphia 1: 132 (1837). Typhonium javanicum Miq., Fl. Ned. Ind. 3: 193 (1856). Typhonium mottleyanum Schott, Prodr. Syst. Aroid.: 106 (1860). Typhonium divaricatum var. mottleyanum (Schott) Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 612 (1879). Typhonium schottii Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 67: 303 (1898). Typhonium divaricatum var. schottii (Prain) Engl., Pflanzenr., IV, 23F: 118 (1920). Typhonium amboinense Blatt. & McCann, J. Bombay Nat. Hist. Soc. 3: 23 (1931). Distribution: Widespread throughout tropical and subtropical Asia, from India to Taiwan and south to Java, Borneo, and east to New Guinea. Notes: Much of the range may be artificial through feralization, although some seemingly natural populations occur in the western part of the Peninsula. Typhonium trilobatum (L.) Schott, Wiener Z. Kunst 3: 72 (1829). Arum trilobatum L., Sp. Pl.: 965 (1753). Dracunculus trilobatus (L.) Raf., Fl. Tellur. 3: 65 (1837). Arum pumilum Lam., Encycl. 3: 8 (1789). Arum orixense Roxb. ex Andrews, Bot. Repos. 5: t. 356 (1804). Typhonium orixense (Roxb. ex Andrews) Schott, Wiener Z. Kunst 3: 72 (1829). Desmesia orixensis (Roxb. ex Andrews) Raf., Fl. Tellur. 3: 63 (1837). Arum auriculatum Sims, Bot. Mag. 49: t. 2324 (1822).

The Checklist

Arisaema pumilum Blume, Rumphia 1: 107 (1836). Typhonium triste Griff., Not. Pl. Asiat. 3: 145 (1851). Typhonium siamense Engl. in A.L.P. de Candolle & A.C.P. de Candolle, Monogr. Phan. 2: 615 (1879). Distribution: Widespread from India through to South China and south as far as Singapore. Notes: How much of the more southerly distributional range in the Peninsular is natural is unclear; T. trilobatum is a weedy species and formely much utilized for medicine.

Wolffia Horkel ex Schleid. Wolffia Horkel ex Schleid., Beitr. Bot. 1: 233 (1844), nom. cons. (Plate 18) Horkelia Rchb. ex Bartl., Ord. Nat. Pl.: 76 (1830). Grantia Griff. ex Voigt, Hort. Suburb. Calcutt.: 692 (1845). Bruniera Franch., Billotia 1: 25 (1864). Taxonomy: Eleven species, widespread in the tropics, subtropics and warm temperate regions of both hemispheres. Two species in Peninsular Malaysia. Most recent account: Landolt (1986). Ecology: Slow-moving or static water in old monsoon drains and road-side ditches, rare. Wolffia angusta Landolt, Veröff. Geobot. Inst. ETH Stiftung Rübel Zürich 70: 29 (1980). Distribution: Peninsular Malaysia to Australia.

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Wolffia globosa (Roxb.) Hartog & Plas, Blumea 18: 367 (1970). Lemna globosa Roxb., Fl. Ind. ed. 1832, 3: 565 (1832). Grantia globosa (Roxb.) Griff., Hort. Suburb. Calcutt.: 692 (1845). Wolffia schleidenii Miq., Ned. Kruidk. Arch. 3: 428 (1855). Telmatophace cylindracea Welw. ex Hegelm., Lemnac.: 123 (1868). Distribution: Tropical and subtropical Asia.

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. 1941. Alocasia macrorrhiza and its varieties. Gard. Bull. Singapore 11: 244– 257. Griffith, W. 1851a. Notulae ad Plantas Asiaticas, vol. 3. Calcutta: Bishop's College Press. . 1851b. Icones Plantarum Asiaticarum, vol. 3. Calcutta: Bishop's College Press. Hay, A. 1988. Cyrtosperma (Araceae) and its Old World allies. Blumea 33: 427–469. . 1996. A new Bornean species of Colocasia Schott (Araceae: Colocasieae),with a synopsis of the genus in Malesia and Australia. Sandakania 7: 31–48. . 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia and Sulawesi. Gard. Bull. Singapore 50: 221–334. Hay, A., Boyce, P.C., Bogner, J., Hetterscheid, W.L.A., Jacobsen, N. & Murata, J. 1995. Checklist & Botanical Bibliography of the Aroids of Malesia, Australia, and the tropical western Pacific. Blumea, Suppl. 8: 1–210. Hay, A. & Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I – Schismatoglottis. Telopea, 9(1): 1–177. Hetterscheid, W.L.A. 1994. Notes on the genus Amorphophallus (Araceae): 2. New species from tropical Asia. Blumea 39: 237–281. Hetterscheid, W.L.A. & Van der Ham, R.W.J.M. 2001. Notes on the genus Amorphophallus (Araceae): 11. New and obsolete species from East Malesia and continental Southeast Asia. Blumea 46: 253–282. Hooker, J.D. 1893. Araceae. In J.D. Hooker (ed.), The Flora of British India 6: 490–556. London: Reeve & Co. Hotta, M. 1985. New species of the genus Homalomena (Araceae) from Sumatra and a short note on the genus Furtadoa. Gardens' Bulletin Singapore 38(1): 43–54. Jack, W. 1820. Descriptions of Malayan plants (No. 1). Malayan Miscellanies, vol.1, no. 1. Bencoolen: Sumatran Mission Press. Jacobsen, N. 1985. Cryptocoryne cordata Griffith. Aqua-Planta 2–85: 15. . 1987. Cryptocoryne purpurea Ridley. Aqua-Planta 2–87: 61–62. Jacobsen, N. & Bogner, J. 1986. Die Cryptocorynen der Malaiischen Halbinsel (1 Teil). Aqua-Planta 11: 135–139. . 1987. Die Cryptocorynen der Malaiischen Halbinsel (2–4 Teil). Aqua-Planta 12: 13–20, 56–60, 96–103. Kartini, S. & Mashhor, M. 1996a. Note on the ecology of Cryptocoryne ciliata. Malayan Nature Journal 50: 165–166. . 1996b. Differential temporal patterns of insect visitors on Cryptocoryne ciliata. Journal of Malaysian Applied Biology 25(2): 95–100. Krause, K. 1908. Araceae-Calloideae. In A. Engler (ed.), Das Pflanzenreich 37(IV. 23B): 140–155. Berlin: W. Engelmann. . 1913. Philodendroideae-Philodendreae-Philodendrinae. In: A. Engler (ed.), Das Pflanzenreich 60(IV. 23Db): 1–143. Berlin: W. Engelmann.

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Landolt, E. 1986. Biosystematic investigations in the family of duckweeds (Lemnaceae), vol. 2. Veröff. Geobot. Inst. ETH Stiftung Rübel Zürich 95. Les, H.L. & Crawford, D.J. 1999 – publ. 2000. Landoltia (Lemnaceae), a new genus of duckweeds. Novon 9: 530–533. Mashhor, M. 1987. The major aquatic plants of Peninsular Malaysia. Aquatics 6: 17–19. . 1997. Noxious floating weeds of Malaysia. Hydrobiologia 360: 121–125. Mashhor, M. & Masnadi, M. 1994. Cryptocoryne elliptica, an endangered amphibious plant in Pondok Tanjung Forest Reserve. Aquatic Botany 47: 91–96. Mashhor, M. & Sulaiman, B. 1997. Notes on Araceae of Pondok Tanjung Forest Reserve, Peninsular Malaysia. Tropical Biodiversity 4: 157–161. Mashhor, M., Zulfigar, Y. & Ali, A. 1990. Notes on the distribution of Cryptocoryne affinis in Sungai Kincin. Malayan Nature Journal 43(3): 256–258. Mayo, S.J., Bogner, J. & Boyce, P.C. 1997. The Genera of Araceae. Kew, UK: Royal Botanic Gardens. Nicolson, D.H. 1964. A taxonomic revision of the genus Aglaonema (Araceae). PhD thesis, Cornell University, New York. . 1968a. The genus Xenophya Schott (Araceae). Blumea 16: 115–118. . 1968b. The genus Spathiphyllum in the east Malesian and west Pacific Islands. Blumea 16: 119–121. . 1968c. A revision of Amydrium (Araceae). Blumea 16: 123–127. . 1969. A revision of the genus Aglaonema (Araceae). Smithsonian Contr. Bot. 1: 1–69. . 1975. Lectotypification of genera of Araceae. Taxon 24: 467–468. . 1984. Suprageneric names attributable to Araceae. Taxon 33(4): 680–690. . 1987. Derivation of aroid generic names. Aroideana 10(3): 15–25. Othman, A.S. 1997. Molecular systematics of the tropical plant genus Cryptocoryne Fischer ex Wylder (Araceae). PhD thesis, University of St Andrews, Scotland. Othman, A.S. & Boyce, P.C. 2010. Studies on Monstereae (Araceae) of Peninsular Malaysia II: Rhaphidophora latevaginata, newly recorded for West Malaysia. Gardens' Bulletin Singapore 62(1): 1–8. Othman, A.S., Boyce, P.C. & Chan, L.K. 2010. Studies on Monstereae (Araceae) of Peninsular Malaysia III: Scindapsus lucens, a new record for Malaysia, and a key to Peninsular Malaysian Scindapsus. Gardens' Bulletin Singapore 62(1): 9–15. Othman, A.S., Jacobsen, N. & Mashhor Mansor. 2009. Cryptocoryne of PeninsularMalaysia. Pulau Pinang: Penerbit Universiti Sains Malaysia. Othman, A.S. & Sim, C.H. 2005. Genetic variation of Cryptocorye minima Ridl. (Araceae) of Selarong River and Jelutung River in Perak. Jurnal Biosains 16(1): 1–12. Othman, A.S., Yu, C.H., Chin, C.L. & Mansor, M. 2007. Genetic variation within the aquatic plant species Cryptocoryne × purpurea in Tasik Bera, Malaysia as revealed by RAPD analysis. Jurnal Biosains 18(2): 99–107.

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Raunkiaer, C.C. 1934. The Life Forms of Plants and Statistical Plant Geography. Oxford: Oxford University Press. Ridley, H.N. 1907. Materials for a Flora of the Malay Peninsula, pt.3. Singapore: Government Printer. . 1925. Flora of the Malay Peninsula, vol. 5. London: Reeve & Co. Roxburgh, W. 1820. Flora Indica. (Carey/Wallich ed.). Serampore: Mission Press. . 1832. Flora Indica. (Ed. 2, Carey). Calcutta: Thacker & Co. Schimper, A.F.W. 1903. Plant-geography Upon a Physiological Basis. Translation of Pflanzen-geographie auf physiologischer Grundlage by William Rogers Fisher, Percy Groom & Isaac Bayley Balfour. William Rogers Fisher, ed. Clarendon Press. Schott, H.W. 1829a. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(3) 88: 752. . 1829b. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(3) 94: 779–780. . 1829c. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(3) 97: 803. . 1829d. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(3) 100: 828. . 1829e. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(3) 108: 892. . 1830. Für Liebhaber der Botanik. Wiener Zeitschr. Kunst, Literatur, Theater und Mode 1829(4) 127: 1028. . 1832. Aroidearum synopsis. In H.W. Schott & S.L. Endlicher (eds.) Meletemata Botanica, 15–22. Vienna: C.Gerold. . 1856. Synopsis Aroidearum. Vienna: Mechitarists' Press. . 1858. Genera Aroidearum exposita. Vienna: Hölzel. . 1860. Prodromus Systematis Aroidearum. Vienna: Mechitarists' Press. Sulaiman, B. & Mashhor, M. 2001. Endemic riverine Araceae of Peninsular Malaysia. In Proceedings of the fourth International Flora Malesiana Symposium 1998, The Cornerstone of Biodiversity, L.G. Saw, L.S.L. Chua and K.C. Khoo (eds.). Kuala Lumpur. Whitmore, T.C. 1975. Tropical Rain Forests of the Far East. Oxford: Clarendon Press. . 1984. Tropical Rain Forests of the Far East. 2nd edn. Oxford: Clarendon Press. . 1990. An Introduction to Tropical Rain Forests. New York: Oxford University Press. . 1998. An Introduction to Tropical Rain Forests. 2nd edn. New York: Oxford University Press. Wong, S.Y. & Boyce, P.C. 2010. Studies on Schismatoglottideae (Araceae) of Borneo IX: A new genus, Hestia, and resurrection of Apoballis. Botanical Studies (Taipei) 51: 249–255.

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Index Adelonema, 80 Afrorhaphidophora, 109 Aglaodorum, 15, 18, 32, 33 Aglaodorum griffithii, 3, 13, 32, 33 Aglaonema, 6, 12, 13, 18, 21, 32, 34 Aglaonema alpinum, 38 angustifolium, 37 angustifolium, var. undulatum, 37 birmanicum, 37 borneense, 37 brevispathum, 19 brevivaginatum, 37 costatum, 13, 34 f. foxii, 34 var. foxii, 34 var. maculatum, 34 elongatum, 37 emarginatum, 37 fallax, 37 flemingianum, 34 grande, 37 integrifolium, 36 latius, 37 longicuspidatum, 36 malaccense, 36 minus, 34 var. nanum, 34 var. scortechinii, 34 nanum, 34 nebulosum, 5, 34, 35 f. nanum, 34

nicobaricum, 37 nieuwenhuisii, 38 nitidum, 35, 36 f. cinereum, 36 f. curtisii, 36 oblanceolatum, 36 f. maximum, 36 oblongifolium, 36 var. curtisii, 36 obovatum, 34 palustre, 32 pictum, var. nanum, 34 var. scortechinii, 34 pierreanum, 37 princeps, 36 propinquum, 37 pumilum, 13 pygmaeum, 87 var. majus, 87 schottianum, 36 f. angustifolium, 37 var. brownii, 37 var. genuinum, 36 var. malaccense, 36 var. winkleri, 37 scortechinii, 34 siamense, 37 simplex, 36 f . angustifolium, 37 f. inaequale, 38 135

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Index

f. nicobaricum, 37 f. nieuwenhuisii, 38 f. typicum, 36 subarborescens, 38 subfalcatum, 37 tenuipes, 37 Aglaonemateae, 13 Allopythion, 44 Alocasia, 6, 10, 18, 20, 21, 38, 39, 40, 57 Alocasia amabilis, 41 cochinchensis, 41 cordifolia, 42 crassinervia, 44 curtisii, 41 cuspidata, 41 denudata, 41 var. elongata, 41 dussii, 63 eminens, 41 grandis, 43 illustris, 63 indica, 42 var. diversifolia, 43 var. heterophylla, 43 var. metallica, 42 var. rubra, 42 var. typica, 42 var. variegata, 43 inornata, 40 korthalsii, 40 leoniae, 41 longifolia, 41 longiloba, 39, 40 lowii, 41 var. picta, 41 var. veitchii, 40 lucianii, 41 macrorrhizos, 5, 12, 42, 43 margaritae, 43 marginata, 43 metallica, 42 neoguineensis, 118 nobilis, 40 ovalifolia, 43 pallida, 42 perakensis, 12, 43 plumbea, 43

puber, 43 pucciana, 41 putzeysii, 41 rapiformis, 42 reginae, 5 reversa, 5 ridleyi, 5 singaporensis, 41 sinuata, 5 spectabilis, 41 thibantiana, 41 uhinkii, 43 variegata, 43 veitchii, 40 Ambrosina, 10, 16 ciliaris, 66 ciliata, 66 Amorphophallus, 10, 13, 14, 16, 18, 19, 20, 22, 44, 45, 46 Amorphophallus bintangensis, 50 boyceanus, 46, 47 bufo, 45, 47 campanulatus, 49 f. darnleyensis, 49 var. blumei, 49 carneus, 45, 47 chatty, 49 decurrens, 49 dixenii, 49 dubius, 49 elatus, 46, 47 elegans, 46, 47 excentricus, 46, 47 giganteus, 48 gigantiflorus, 49 haematospadix, 45, 48 loerzingii, 50 longituberosus, 48 malaccensis, 49 manta, 45, 48 microappendiculatus, 49 microspadix, 50 minor, 48 paeoniifolius, 48 var. campanulatus, 49 panomensis, 50 perakensis, 50

Index

prainii, 50 rex, 49 sativus, 49 siamensis, 48 sparsiflorus, 50 viridis, 48 virosus, 49 Amydrium, 6, 50, 51 Amydrium humile, 13, 52, 72 medium, 19, 51 Anadendrum, 52, 53, 72, 74 Anadendrum angustifolium, 54 latifolium, 54 malaianum, 54 marginatum, 54 medium, 52 microstachyum, 54 montanum, 114 Anthelia, 72 Anthurium, 13, 14 Anubias, 14 Apatemone, 116 Apiospermum, 100 Apiospermum obcordatum, 102 Apoballis, 18, 54, 55 Apoballis acuminatissima, 55 brevipes, 55, 56 mutata, 55, 56 Areae, 20, 22 Aridarum, 21 Ariopsis, 19, 20, 21 Arisacontis, 69 Arisacontis chamissonis, 69 Arisaema, 9, 13, 16, 18, 20, 21, 57, 58 Arisaema anomalum, 58, 59 cuspidatum, 60 fallax, 59 filiforme, 59 f . pedatisectum, 59 f. ternatum, 59 var. fallax, 59 var. sumatranum, 59 var. typicum, 59 var. verrucosum, 59 fimbriatum, subsp. fimbriatum, 58, 59 fouyou, 64 harmandii, 60

kunstleri, 60 laminatum, 59 makoyanum, 59 pierreanum, 60 pumilum, 127 putii, 59 roxburghii, 58, 60 var. kunstleri, 60 scortechinii, 4, 60 stictopodum, 59 wrayi, 5, 58, 60 Arisarum, 9 Aroideae, 15 Aron colocasium, 62 Arum, 9, 16 Arum angulatum, 125 auriculatum, 126 campanulatum, 48 chinense, 62 ciliatum, 66 colocasia, 62 colocasioides, 62 cordifolium, 42 cuspidatum, 60, 125 decurrens, 49 divaricatum, 125 diversifolium, 126 esculentum, 62 filiforme, 59 flagelliferum, 125 flagelliforme, 125 indicum, 42 integrifolium, 36 lividum, 62 macrorrhizon, 42 mucronatum, 42 nymphaeifolium, 62 orixense, 126 peltatum, 62 peregrinum, 42 phalliferum, 49 ptychiurum, 125 pumilum, 126 rapiforme, 42 rumphii, 48, 49 trilobatum, 126

137

138

Index

Baharuddin Sulaiman, 6 Batis, 105 Batis hermaphrodita, 108 Biarum, 9, 16 Bogner, Josef, 6 Boyce, Peter, 6 Brachyspatha, 44 Brown, Nicholas Edward, 2, 4–5, Bruniera, 127 Caladium acre, 62 colocasia, 62 colocasioides, 62 esculentum, 62 giganteum, 64 glycyrrhizum, 62 indicum, 43 lowii, 41 macrorrhizon, 42 metallicum, 43 nymphaeifolium, 62 odoratum, 42 plumbeum, 43 princeps, 36 simplex, 36 veitchii, 40 violaceum, 62, 63 Calla, 10 Calla angustifolia, 87 badian, 42 calyptrata, 116 gaby, 62 humilis, 86 maxima, 42 montana, 114 nitida, 36 sylvestris, 115 virosa, 62 Candarum, 44 Candarum hookeri, 48 roxburghii, 48 rumphii, 48 Chamaecladon, 80 Chamaecladon angustifolium, 87 griffithii, 85 humile, 86

lanceolatum, 87 obliquatum, 85 ovatum, 85 purpurascens, 87 pygmaeum, 87 var. latifolium, 87 sanguinolentum, 90 Colobogynium, 116 Colocasia, 6, 19, 21, 40, 60 Colocasia acris, 62 aegyptiaca, 63 antiquorum, f. acuatica, 63 f. eguimo, 63 f. oyasetage, 63 f. purpurea, 63 f. yamamotoi, 64 var. multifolia, 63 var. patens, 63 var. rosea, 63 var. rupicola, 63 var. stolonifera, 63 boryi, 42 colocasia, 62 esculenta, 12, 60, 61, 62 f. ebiimo, 64 f. rotundifolia, 64 var. aquatilis, var. rupicola, 63 var. stolonifera, 63 euchlora, 63 gigantea, 40, 61, 64 gracilis, 63 himalensis, 63 humilis, 118 indica, 42 var. rubra, 42 macrorrhizos, 42 mucronata, 42 neocaledonica, 63 neoguineensis, 118 nymphaeifolia, 62 peltata, 62 peregrina, 42 prunipes, 64 pubera, 43 rapiformis, 42

Index

tonoimo, 64 vera, 63 virosa, 62 vulgaris, 62 Conophallus, 44 Conophallus, giganteus, 48 sativus, 49 Corner, Edred John Henry, 5 Corynophallus, 44 Cryptocoryne, 6, 12, 15, 18, 19, 64, 65, 66 Cryptocoryne affinis, 15, 65, 66 subsp. haerteliana, 66 alata, 66 amicorum, 68 blassii, 67 caudata, 68 ciliata, 3, 15, 22, 66 var. latifolia, 66 cordata, 3, 15 var. diderici, 67 cordata × C. nurii [= Cryptocoryne, × timahensis], 68 cordata × C. griffithii [= Cryptocoryne × purpurea], 68 decus-silvae, 67 diderici, 67 drymorrhisa, 66 elata, 66 elliptica, 67 evae, 67 var. recordata, 67 gasseri, 68 griffithii, 3, 67 haerteliana, 66 hejnyi, 68 johorensis, 68 kerrii, 67 longicauda, 68 minima, 68 nurii, 68 × purpurea, 65, 68 × purpurea nothovar. purpurea, 68 schulzei, 65, 68 siamensis, 67 var. ewansii, 67 var. kerrii, 67 spiralis, 15

stonei, 67 × timahensis, 68 zewaldiae, 68 zukalii, 69 Cryptocoryneae, 15, 21 Culcaseae, 14 Curmeria, 80 Curtis, Charles, 4 Cuscuaria, 120 Cyrtocladon, 80 Cyrtocladon sanguinolentum, 90 Cyrtosperma, 2, 15, 69, 70 Cyrtosperma angustilobum, 105 bantamense, 71 chamissonis, 69 cuspidilobum, 71 dubium, 71 edule, 71 intermedium, 71 lasioides, 69 merkusii, 12, 69, 70 var. giganteum, 71 var. intermedium, 71 nadeaudianum, 71 Desmesia, 124 orixensis, 126 Diandriella, 80 Dochafa, 57 Dracontium paeoniifolium, 48 polyphyllum, 49 spinosum, 94 Dracunculus, 10, 16 Dracunculus divaricatus, 126 trilobatus, 126 Dr King’s Collector, 3 Dunalia, 44 Eminium, 10, 16 Engler, Adolf, 3 Ensolenanthe, 38 Epipremnopsis, 50 Epipremnopsis huegelii, 52 media, 52 subcordata, 52 Epipremnum, 72, 73 Epipremnum angustilobum, 76

139

140

Index

beccarii, 111 ceramense, 72 elegans, 76 f. ternatensis, 76 falcifolium, 72 foraminiferum, 112 formosanum, 76 giganteum, 19, 75 glaucicephalum, 76 humile, 52 medium, 52 merrillii, 76 mirabile, 75 f. eperforatum, 76 f. multisectum, 76 multicephalum, 113 pinnatum, 73, 74 f. multisectum, 76 robinsonii, 76 truncatum, 52 Flagellarisaema, 57 Furtado, Caetano Xavier Dos Remedios, 5 Furtadoa, 5, 77, 80, 83 Furtadoa mixta, 77 sumatrana, 77 Gamble, James Sykes, 5 Gamogyne, 98 Goniurus, 105 Gorgonidium, 16 Grantia, 127 Grantia globosa, 128 Griffith, William, 2 Hansalia, 44 Hapale, 77 Hapaline, 18, 20, 77, 78 Hapaline brownii, 78 Hay, Alistair, 6 Helicodiceros, 9 Hestia, 78, 79 Hestia longifolia, 78, 79, 80 Heteroarisaema, 57 Heterostalis, 124 Heterostalis flagelliformis, 125 Hetterscheid, Wilbert, 7

Holochlamys, 14 Homalomena, ix, 2, 5, 10, 12, 13, 14, 18, 19, 20, 21, 80, 81, 82 Homalomena angustifolia, 87 var. ophirensis, 87 var. parvula, 84 argentea, 84 var. purpurascens, 84 asmae, 84 beccariana, 90 brevispatha, 84 calyptrata, 116 coccinea, 88 confusa, 84 corneri, 85 curtisii, 85 curvata, 85 deltoidea, 86 elliptica, 86 var. paucinervia, 86 ellipticifolia, 88 ensiformis, 90 expedita, 15 falcata, 86 grabowskii, 86 griffithii, 3, 85 f. acuminata, 86 f. eugriffithii, 86 f. falcatoidea, 86 f. kingii, 85 f. obliquata, 85 f. sigmoidea, 86 var. acuminata, 86 var. obliquata, 85 var. ovata, 85 var. sumatrana, 86 habokoana, 86 hayupensis, 86 hendersonii, 86 humilis, 82, 86 var. major, 87 var. major, subvar. coccinea, 88 var. ophirensis, 87 var. ovatifolia, var. parvula, 88 var. pumila, 87 var. undulata, 87

Index

var. velutina, 87 insignis, 5 johorensis, 88 kiahii, 88 f. maculata, 88 kingii, 85 lancifolia, 89 major, 89 minor, 89 miqueliana, 90 var. truella, 90 mixta, 77 montana, 89 multinervia, 86 nutans, 89 obliquata, 85 ovata, 85 paludosa, 90 pineodora, 89 pontederifolia, 81, 89 propinqua, 87, 90 pumila, 87 var. latifolia, 87 var. purpurascens, 87 purpurascens, 87 pusilla, 88 pygmaea, 87 f. longipes, 88 var. latifolia, 87 var. pumila, 88 var. purpurascens, 87 raapii, 90 repens, 88 ridleyana, 90 rostrata, 3, 12, 13, 90 rubrovaginata, 88 var. subpurpurea, 88 sagittifolia, 90 var. angustifolia, 90 var. pontederiifolia, 90 var. sumatrana, 90 scortechinii, 4, 91 singaporensis, 91 Supergroup Chamaecladon, 83 Cyrtocladon, 84 Homalomena, 83

teysmannii, 90 trapezifolia, 85 triangularis, 90 truncata, 13, 91 undulatifolia, 87 velutina, 87 wallichii, 2, 91 Homalomeneae, 13 Homalonema, 80 Hooker, Joseph Dalton, 2 Horkelia, 127 Hydrophace, 95 Hydrophace minor, 97 Hydrosme, 44 Hydrosme gigantiflora, 49 longituberosa, 48 Jack, William, 2 Jacobsen, Niels, 6 King, George, 3, 5 Kodda-Pail, 100 Kunda, 44 Kunda verrucosa, 48 Kunstler, Herman H., 3 Lagenandra, 21 Landoltia, 91, 92 Landoltia punctata, 92 Lasia, 15, 18, 22, 92, 93 Lasia aculeata, 94 concinna, 92, 94 crassifolia, 95 f. angustisecta, 95 f. latisecta, 95 desciscens, 94 hermannii, 94 heterophylla, 94 jenkinsii, 94 loureiroi, 94 merkusii, 69 roxburghii, 94 spinosa, 92, 93, 94 stipitata, 105 zollingeri, 94 Lasioideae, 15 Lasius, 92

141

142

Index

Lemna, 95 Lemna aequinoctialis, 95, 96 angolensis, 95 aoukikusa, 97 subsp. hokurikuensis, 97 bannatica, 123 blatteri, 97 conjugata, 97 cyclostasa, 97 eleanorae, 97 globosa, 128 javanica, 92 leiboensis, 97 major, 124 maxima, 124 melanorrhiza, 92 minima, 97 minor, 97 var. minima, 97 var. oxymitra, 97 monorhiza, 98 obcordata, 97 oligorrhiza, 92 orbicularis, 123 orbiculata, 123 ovata, 97 palustris, 97 paucicostata, 95 var. membranacea, 97 perpusilla, var. trinervis, 95 pleiorrhiza, 92 polyrrhiza, 123 var. concolor, 124 punctata, 92 pusilla, 92 rwandensis, 98 tenera, 98 thermalis, 123 transsilvanica, 124 trinervis, 95 umbonata, 124 vulgaris, 97 Lemnaceae, 15 Lemnoideae, 15, 18, 96 Lenticula, 95 Lenticula cyclostasa, 97 minima, 97

minor, 97 palustris, 97 polyrrhiza, 123 vulgaris, 97 Lenticularia, 95 Lenticularia monorhiza, 98 Leucocasia esculenta, 62, 64 Limnonesis, 100 Limnonesis commutata, 102 friedrichsthaliana, 102 Low, Hugh (Sir), 3 Lysichiton, 9 Maingay, Alexander Carroll, 4 Mangonia, 16 Mashhor Mansor, 6 Monstera caudata, 75 dilacerata, 75 gigantea, 75 lingulata, 115 pinnata, 75 Monstereae, 19, 21, 72 Monsteroideae, 14 Montrichardia, 15 Muricauda, 57 Myrioblastus, 64 Nebrownia, 116 Nicolson, Dan H., 6 Orontioideae, 15, 18 Orontium, 9 Panzhuyuia, 38 Pedicellarum, 72, 74, 105 Peltandra, 9 Philodendron, 14, 15, 20 Philodendron dilaceratum, 76 nechodomae, 76 peregrinum, 42 punctatum, 42 Pinellia, 9, 16 Piptospatha, 18, 19, 98 Piptospatha elongata var. perakensis, 98 perakensis, 98, 99 ridleyi, 5, 99, 100 Pistia, 15, 18, 19, 22, 100, 101

Index

Pistia aegyptiaca, 102 aethiopica, 102 africana, 102 amazonica, 102 brasiliensis, 102 commutata, 102 crispata, 102 cumingii, 102 gardneri, 103 horkeliana, 102 leprieuri, 102 linguiformis, 102 minor, 102 natalensis, 103 obcordata, 102 occidentalis, 102 schleideniana, 103 spathulata, 102 stratiotes, 15, 101 var. linguiformis, 103 var. obcordata, 102 var. spathulata, 102 texensis, 103 turpinii, 102 weigeltiana, 102 Plesmonium, 44 Plesmonium nobile, 49 Pleuriarum, 57 Podolasia, 15, 103, 104 Podolasia stipitata, 12, 103, 104, 105 Podospadix angustifolia, 108 Polypodium laciniatum, 75 Porter, George, 2 Potheae, 19 Pothoideae, 14, 72 Pothoidium, 74 Pothos, 6, 10, 18, 19, 72, 74, 105, 106 Pothos angustifolius, 108, 109 argenteus, 122 argyraeus, 122 barberianus, 107 bifarius, 112 caudatus, 75 celatocaulis, 112 chapelieri, 109 cognatus, 109 curtisii, 4, 105, 107

decipiens, 109 decursivus, 75 ellipticus, 107 enderianus, 123 exiguiflorus, 109 fallax, 109 giganteus, 75 gigantipes, 106 grandispathus, 107 hederaceus, 122 hermaphroditus, 108 heterophyllus, 94 horsfieldii, 109 junghuhnii, 106 kingii, 4, 105, 107 kunstleri, 107 lancifolius, 107 lasia, 94 latifolius, 107 leptospadix, 108 leptostachyus, 105, 106, 107 longifolius, 108 longipedunculatus, 108 lorispathus, 107 macrocephalus, 105, 106, 108 maingayi, 107 malaianus, 54 merrillii, 108 microphyllus, 108 monopetalus, 54 oliganthus, 105 ovatifolius, 105, 108 var. simalurensis, 108 penicilliger, 107 peninsularis, 107 pinnatifidus, 75 pinnatus, 75 ridleyanus, 107 roxburghii, 108 salicifolius, 108 scandens, 106, 108 f. angustior, 109 var. cognatus, 109 var. helferianus, 109 var. sumatranus, 108 var. zeylanicus, 108 var. zollingerianus, 109

143

144

Index

spinosus, 94 subgenus Allopothos, 105 subgenus Pothos, 74, 105 Supergroup Allopothos, 105 Supergroup Goniurus, 105 zollingeri, 109 zollingerianus, 109 Prain, David (Sir), 5 Proteinophallus, 44 Pseudodracontium, 44 Pycnospatha, 22 Pythion, 44 campanulatum, 49 Pythonium, 44 Remusatia, 14, 18, 20 Rhaphidophora, 10, 12, 18, 19, 72, 109, 110 Rhaphidophora angulata, 114 angustata, 13, 111 angustifolia, 115 apiculata, 113 batoeensis, 114 beccarii, 12, 111 borneensis, 111 burkilliana, 13, 111, 112 caudata, 75 celatocaulis, 112 celebica, 114 copelandii, 112 corneri, 112 crassifolia, 112 cunninghamii, 76 dilacerata, 75 fallax, 114 fluminea, 111 foraminifera, 112 formosana, 76 gigantea, 75 gracilipes, 114 grandifolia, 113 grandis, 112 gratissima, 115 hallieri, 114 Hongkongensis Group, 111 hookeri, 19 huegelii, 52 humilis, 52

jaculiformis, 113 korthalsiana, 52 korthalsii, 18, 74, 110, 112 kunstleri, 114 laciniata, 75 laetevirens, 111 latevaginata, 111, 113 latifolia, 113 lingulata, 115 lobbii, 12, 110, 113 lovellae, 76 magasperma, 111, 113 maingayi, 4, 113 maxima, 112 merrillii, 76 minor, 12, 114 montana, 114 motleyana, 115 neocaledonica, 76 nicolsonii, 6, 12, 114 nigrescens, 115 palawanensis, 113 pilosula, 114 pinnata, 75 pinnatifida, 75 pteropoda, 111 puberula, 114 ridleyi, 112 scaberula, 114 scortechinii, 114 subfalcata, 113 sylvestris, 115 tetrasperma, 74, 115 trinervia, 113 trukensis, 113 vitiensis, 76 wallichii, 76 wrayi, 115 Rhaphiophallus, 44 Rhynchopyle, 98 Rhynchopyle perakensis, 98 Ridley, Nicholas Henry, 5 Roxburgh, William, 2 Schismatoglottideae, 13, 14, 20 Schismatoglottis, 6, 10, 13, 18, 19, 20, 21, 83, 116, 117

Index

Schismatoglottis acutangula, 118 angustifolia, 119 batoeensis, 56 bitaeniata, 118 brevicuspis, 116, 117 brevipes, 56 brooksii, 56 calyptrata, 116, 117 f. minor, 118 var. ornata, 118 calyptratoides, 119 conversa, 120 cordifolia, 118 djamuensis, 118 dorensis, 119 emarginata, 118 hellwigiana, 118 var. subcordata, 118 kingii, 119 klossii, 118 leptophylla, 56 linguiformis, 56 longicuspis, 120 longifolia, 80 longipes, 118 maculata, 119 marginata, 119 minor, 56 monticola, 56 mutata, 56 neoguineensis, 118 nicolsonii, 6 nieuwenhuisii, 118 parvifolia, 119 penangensis, 119 picta, 118 potamophila, 119 pseudocalyptrata, 119 ridleyana, 56 riparia, 118 rotundifolia, 56 ruttenii, 119 scortechinii, 4, 117, 119 sublaxiflora, 119 tenuifolia, 118 vanvuurenii, 119 variegata, 118

wallichii, 2, 120 f. oblongata, 120 var. fasciata, 120 var. oblongata, 120 Schizocasia, 38 Scindapsus, 10, 12, 14, 18, 72, 120, 121 Scindapsus angulatus, 114 angustifolius, 115 anomalus, 112 argyraeus, 122 aruensis, 115 beccarii, 122 beccarii, Complex, 120 bipinnatifidus, 76 caudatus, 75 coriaceus, Complex, 120 decursivus, 75 dilaceratus, 75 forsteri, 75 fragilis, 115 giganteus, 75 havilandii, 113 hederaceus, 122 huegelii, 52 inquinatus, 122 latifolius, 72 lingulatus, Hassk, 115 lobbii, 113 longipetiolatus, 122 lucens, 121, 122 marginatus, 54 medius, 52 microstachyus, 54 montanus, 114 perakensis, 120, 122 pictus, 14, 121, 122 var. argyraeus, 122 var. oblongifolius, 122 pinnatifidus, 75 pinnatus, 75 poilanei, 122 pothoides, 122 pteropodus, 111 roseus, 72 scortechinii, 4, 12, 121, 123 splendidus, 72 sylvestris, 115

145

146

Index

tonkinensis, 38 treubii, 123 Schott, Heinrich Wilhelm, 3 Scortechini, Rev. Fr. Benedetto, 3 Sofiman Othman, 6 Spathiphylleae, 13 Spathiphyllum, 6 Spirodela, 91, 92, 123 Spirodela atropurpurea, 124 maxima, 124 oligorrhiza, 92 var. javanica, 92 var. melanorrhiza, 92 var. pleiorrhiza, 92 var. pusilla, 92 polyrrhiza, 96, 123 var. masonii, 124 punctata, 92 Spirospatha, 80 Staurogeton, 95 tener, 98 Stenospermation, 14 Steudnera, 20 virosa, 62 Symplocarpus, 9 Synandrospadix, 16 Synantherias, 44 Syngonium, 21

var. robustum, 126 var. roxburghii, 126 var. schottii, 126 flagelliferum, 125 flagelliforme, 12, 15, 125 var. angustissimum, 125 fultum, 125 hastiferum, 125 incurvatum, 125 javanicum, 126 mottleyanum, 126 orixense, 126 reinwardtianum, 125 roxburghii, 126 schottii, 126 siamense, 127 sylvaticum, 125 trilobatum, 126, 127 triste, 127 Typhonodorum, 15, 18

Tapanava, 105 Tapanava indica, 108 rheedei, 108 Tapeinophallus, 44 Telmatophace, 95 Telmatophace cylindracea, 128 orbicularis, 123 polyrrhiza, 123 Thomsonia, 44 Thomsonieae, 20 Tornelia dilacerata, 75 Typhonium, 13, 14, 15, 19, 22, 124 Typhonium amboinense, 126 cuspidatum, 125 var. ptychiurum, 125 divaricatum, 126 var. mottleyanum, 126

Xenophya, 6, 38

Wallich, Nathaniel, 2 Wolffia, 127 Wolffia angusta, 127 globosa, 96, 128 schleidenii, 128 Wong Sin Yeng, 6 Wray, Leonard ( Jr.), 4

Zala, 100 Zala asiatica, 102 Zantedeschia calyptrata, 116 virosa, 62