THE GARDENS’ BULLETIN, SINGAPORE

The Gardens’ Bulletin, Singapore is a peer-reviewed journal publishing original papers and

reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,

floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and

conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.

Dr. K.M. Wong Dr. Jana Leong-Skorniékova

(Editor-in-Chief) (Managing Editor)

Salvec Y.W. Low Christina Soh

(Graphics Editor) (Copy Editor) (Business Manager)

Editorial Advisory Board

Professor Sir Peter Crane Dr. Nigel P. Taylor

Yale University Singapore Botanic Gardens

U.S.A. National Parks Board, Singapore

Dr. Rogier P.J. de Kok Dr. I.M. Turner

Royal Botanic Gardens, Kew Singapore Botanic Gardens

U.K. National Parks Board, Singapore

Dr. W.J. Kress Dr. J.F. Veldkamp

National Museum of Natural History Nationaal Herbarium Nederland

Smithsonian Institution The Netherlands

U.S.A. Dr. Jun Wen

Dr. David J. Middleton National Museum of Natural History

Royal Botanic Garden Edinburgh Smithsonian Institution

U.K. U.S.A.

Dr. Kiat W. Tan Professor Nianhe Xia

Gardens By The Bay South China Institute of Botany

National Parks Board, Singapore PR. China

Published twice yearly by the National Parks Board, Singapore, the annual subscription for the journal

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(continued on inside back cover)

ARNOLD ARBORET::.

~\_ LIBRARY

wow 1 4 2012

The Gardens’ Bulletin

Singapore

VOL. 64(1) 2012 ISSN 0374-7859

Centenary Issue 2012

CONTENTS

K.M. Wong

Menunered years of the Gatdens’ Bulletin, Singapore ....................2..ccccessccsesncesseooecnetes l

Charles Clarke and Ch’ien C. Lee

A revision of Nepenthes (Nepenthaceae) from Gunung Tahan, Peninsular Malaysia .....

Helena Duistermaat

A taxonomic revision of Amischotolype (Commelinaceae) 1n AS1a ..........:::ceeeeeeees 51

H. Kurzweil and S. Lwin

First record of Taeniophyllum (Orchidaceae) in Myanmat .................:ccescceeeeeereeeeees 133

C.Y. Ling and S. Julia

Diversity of the tree flora in Semenggoh Arboretum, Sarawak, Borneo ................. 139

R.R. Mill and M. Whiting

Podocarpus orarius (Podocarpaceae), a new species from the Solomon Islands and a

taxonomic clarification of Podocarpus spathoides from Malaysia ..............::0:000++ 171

Q.B. Nguyen and Jana Leong-Skorniékova

Distichochlamys benenica (Zingiberaceae), a new species from Vietnam .............. 195

Hans-Juergen Tillich and Leonid V. Averyanov

Four new species of Aspidistra Ker-Gawl. (Asparagaceae) from China and Vietnam

with a comment on A. Jongifolia Hook.f. and A. hainanensis W.Y.Chun & F.C.How ...

seg sng Sea osc ge SS SACO DSO a Ene re 201

A.P.J. Ting, S.Y. Wong, J. Jamliah and P.C. Boyce

Phylogenetic study of the Schismatoglottis Nervosa Complex (Araceae:

Schismatoglottideae): sxe. aloe eec eee ees eee soe aee ease ta cee eee 211

I.M. Turner

The plant taxa of H.N. Ridley, 4. The primitive angiosperms (Austrobaileyales,

Canellales, Chloranthales, Laurales, Magnoliales, Nymphaeales and Piperales) .... 221

S.Y. Wong, P.C. Boyce and S.L. Low

Studies on Schismatoglottideae (Araceae) of Borneo XVII: The Schismatoglottis

Hottae Complex, a new informal taxon, and three new species from Sarawak,

Malaysian B Onno: z2..c.2:c0.aecaccenseeeseee tech oo 2s eink e ons ssc Jn aceecoes stecteee scene tage teen meee 257

BOOK REVIEWS

Heritage Trees of Penang. S. Gardner, P. Sidisunthorn & E.M. Lai. 2011. (S.K.

GF: Geko 10) ee ee A ae er ea Ecce oot Quousassoacevoocetes 271

The Wild Orchids of Hong Kong. G. Baretto, PJ. Cribb & S. Gale. 2011. (Hubert

Kaa ZEAL) isc daceg cases centeed eh sehen Souee banisab ds cova leas vcnicees pcan an ee dsaap eee eee eee DAS

Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 2. R. Kiew, R.C.K. Chung,

LiG Saw, E. Soepadmo @i-G. Boyce (eds); 20 MACY. WL Ow)eeece eee 20h

Wild Orchids of Peninsular Malaysia. P 7: Ong, P. O'Byrne, W.S.Y. Yong & L.G. Saw.

2011. (KM. Wong) tazcsei.cestan staasaeess od catia dates tN chur beet aes anes eee eee 279

Date of publication: 15 July 2012

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 64(1): 1-32. 2012 l

A hundred years of the Gardens’ Bulletin, Singapore

K.M. Wong

Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. Historical developments are traced pertaining to the founding and transformation

of the Agricultural Bulletin of the Malay Peninsula, 1891—1900, the Agricultural Bulletin of

the Straits and Federated Malay States, 1901—1911, the first two periodicals of the Singapore

Botanic Gardens, and the Agricultural Bulletin of the Straits and Federated Malay States, Third

Series, which began in 1912. This third series soon continued as the Gardens’ Bulletin, Straits

Settlements when in 1913 it was decided to continue the journal from the Botanic Gardens

with a name change to avoid confusion with an Agricultural Bulletin separately begun for

the Federated Malay States, as their new Department of Agriculture developed and economic

activities around agriculture intensified. After World War II, this continued as the Gardens’

Bulletin, Singapore, which achieved its centenary in 2012. The early focus on the Hevea rubber

crop and industry during the time of H.N. Ridley, its founding editor, and the re-orientation of

the Gardens’ Bulletin into a journal with increased original content in the botanical (especially

taxonomic) sciences from the period of I.H. Burkill, Ridley’s successor, are described.

Historical events, especially the administrative divergence between the Straits Settlements and

the Federated Malay States, the impact of World War II and post-war political development,

the development of administrative organisation within the newly independent Singapore; and

the integration of botanical science over the Malesian botanical region wherein the Malay

Peninsula is located, have contributed to shaping the focus and scope of the Bulletin. The

development phases of the Singapore Botanic Gardens—home of the Bu//etin—as well as the

pivotal roles of its leading botanists, are examined, through stages of scientific transformation

from an essentially “Malayan” perspective largely maintained by a small botanical home team,

to a more regionally relevant research programme, and finally an international outlook that

continues to sustain its Southeast Asian emphasis.

Keywords. Agricultural Bulletin, botanical journal, Federated Malay States, Flora Malesiana,

Gardens’ Bulletin, Hevea rubber, Malaya, Malay Peninsula, Malesia, plant taxonomy, Ridley,

Singapore Botanic Gardens, Straits Settlements

A centenary measures not just antiquity, but also continuity, progress and outlook. Its

arrival makes us take stock of developments and the many phases that would have

naturally accrued. Even though centenaries are now much more commonly observed

than previously, still each arriving occasion never fails to be evocative, for a hundred

years of history yield a great many interactions and events that would have touched

very many aspects of life and society. This year, 2012, the Gardens ' Bulletin, Singapore

turns a hundred years old.

Scientific publishing at the Singapore Botanic Gardens began when the

organisation was part of a larger entity called the Gardens and Forest Department,

2 Gard. Bull. Singapore 64(1) 2012

Straits Settlements. The Straits Settlements were a British Crown Colony made up

of Malacca, Penang, Province Wellesley, Singapore, and Pangkor and the Sembilan

Islands off the Perak (mid-western) coastline of the Malay Peninsula. The Gardens and

Forest Department was created in 1883 under the Superintendent of Gardens, Straits

Settlements, a position occupied by Nathaniel Cantley from 1880 to 1888. Although the

Singapore Botanic Gardens had been incepted in 1859 at its present site in Tanglin by

the Singapore Agri-Horticultural Society, its maintenance was passed onto the Straits

Settlements Government in late 1874, and by 1878 the legislation was approved for the

Gardens to be officially managed by the colonial government (Burkill 1918a, b). Its

main role then was to serve agricultural and economic development for its territories

as well as the rest of Malaya (Burkill 1983), a geographical designation that referred to

the main part of the Malay Peninsula (including Singapore) which came under British

influence. Cantley was only the second full-time horticultural superintendent recruited

from the Royal Botanic Gardens, Kew, following James Murton (1875—1880).

Prelude as an agricultural bulletin

In 1888, Henry Nicholas Ridley became the first Director of the Singapore Botanic

Gardens and was intricately linked to the development of the Gardens’ periodicals that

followed. In fact, two other series preceded the present Bulletin. The first was called

the Agricultural Bulletin of the Malay Peninsula, with No. | (pages 1-17) issued in

April 1891 and No. 9 (pages 239-289), the last, issued in May 1900. The front page

for each of the first five issues (up to May 1896) declared its prospectus:

It is proposed to publish from time to time, as occasion may serve, Bulletins on subjects

connected with Agriculture and Horticulture in the Malay Peninsula. It is hoped

that planters will send to the Director of the Botanic Gardens, Singapore, notes and

observations on the cultivations of the various crops under their care. Observations on

insect and fungus-pests are specially requested, and these should always be accompanied

by specimens of the injurious insect or fungi either alive, or preserved in spirits, except

in the case of butterflies and moths which should be sent dry in envelopes.

Although most of the issues carried brief articles, the coverage included diseases

of coffee, nutmeg and clove trees, pests and injurious fungi. There were interesting

essays on crops of potentially wider cultivation, such as gambir, patchouli, sago, sugar

cane, lemon grass oil, citronella oil, fibre plants, dye plants, spices, vegetables, and,

of course, Para rubber. Ridley edited and wrote most of the material and was himself

only occasionally credited at the end of an article from issue No. 5 (1896) onwards,

with Charles Curtis (Assistant Superintendent of Gardens and Forests in Penang,

1884-1903) contributing two articles on the cultivation of pot plants and sugar cane

plants developing from seed, and A.L. Butler (of the Selangor Museum), an account

of an extensive bee-hawk moth caterpillar attack on a coffee plantation. The brevity

of the issues was not surprising, as Ridley’s time was also taken up for developing

the Gardens, organising an improved herbarium and specimen exchange with, and

contributions from, Kew, Calcutta, the British Museum and collectors of the Malayan

Centenary of Gardens’ Bulletin, Singapore 3

flora in general, besides himself conducting an active field survey and collecting

programme into Malayan forests and starting many horticultural investigations.

The issuance of an agricultural bulletin made sense with the emphasis given

to economic botany. The purview of the Singapore Botanic Gardens was not only to

assist in introducing crops of economic benefit and making studies for enhancing their

cultivation, but also to help explore and document what was then a very poorly known

flora of Malaya. As it would turn out, Ridley’s well-documented accomplishments

were as incredible in consequence as they were wide in scope. He not only pioneered

the cultivation and excision tapping techniques that helped to encourage the planting

of Hevea as an economic crop (Fig. 1) (Wycherley 1959, Brockway 2000), but was

probably the best known explorer and scribe of the Malayan flora (Ridley 1922-1925).

A distinct Department of Forests, Straits Settlements (S.S.) and Federated

Malay States (F.M.S.), headquartered in Kuala Lumpur, was set up in 1901 after the

urging of both Ridley and H.C. Hill (1900a, b) of the Indian Forest Service, the latter

commissioned to report on forest administration in these territories. (The F.M.S. were

formed by the states of Perak, Selangor, Negri Sembilan and Pahang, each of which had

a British Resident.) Responsibility for forest administration was thus separated from the

Gardens’ function, with Alfred M. Burn-Murdoch transferred from the Burma Forest

Service to be Chief Forest Officer of the S.S. and F.M.S. in 1901, this post becoming

the Conservator of Forests from 1904 (Wong 1987). The Singapore Botanic Gardens

continued to be the centre for disseminating information on agriculture (Burkill 1983),

so the beginning of a new and more focussed bulletin was commensurate with this

role. This second series of the bulletin was the Agricultural Bulletin of the Straits and

Federated Malay States, with Vol. 1, No. 1 issued in October 1901 and carrying an

introduction by Walter Fox, Acting Director of the Singapore Botanic Gardens when

Ridley was away on leave:

... having regard to the large and increasing number of Agriculturists and others taking an

interest in Agriculture, the time has now come when something more than the Bulletin

mentioned above is required... It is now proposed to publish a monthly Bulletin, which

shall incorporate the old one, and as far as possible enlarge its scope by making it the

medium for the exchange and record of Planters’ experiences in all that pertains to their

interests ...1n a word make it a Planters’ Paper for Planters.

The march of the bulletins

Ridley continued to serve as editor. The monthly issues in this second series

carried a few longer articles but had a good measure of attention given to potential,

experimental or emerging crops, and commodity prices in London and Singapore. The

products listed reflected market interest then and included cocaine, then used as a local

anaesthetic and becoming more commonly known as a performance enhancer (Karch

1998). Mention on page 35 in No. | of Vol. 1 (October 1901) stated: “The exports of

crude cocaine from Peru during 1900 amounted to 16,479 Ibs., valued at $563,625. Of

this quantity the United States received 1,016 lbs. direct.” Ridley’s article on Timbers

4 Gard. Bull. Singapore 64(1) 2012

ne pie i. nn ERY Eee

Fig. 1. Henry Ridley (left) with a rubber-tapping experiment. From Gardens Bulletin, Straits

Settlements 1(8), facing p. 263.

Centenary of Gardens’ Bulletin, Singapore 5

of the Malay Peninsula anchored the inaugural number and subsequent instalments up

to its conclusion in No. 8 (May 1902). His other contribution made in instalments was

Fruits of the Malay Peninsula, Wild or Cultivated (Vol. 1 No. 10: 371-381, printed

as July 1902 but hand-corrected to August), which continued into Nos. 11, 12 and 13.

Shorter articles included such as The World’ Tea and Coffee Consumption (No. 3:

115-117), Notes on Gutta Percha Trees by Charles Curtis (No. 6: 220-223), 4 New

Instrument for Tapping Rubber Trees (No. 6: 230-231), New Rubber in Saigon (No.

7: 274), Rubber vines in French Indo-China (No. 9: 333-335) and The Cultivation

of Orchids for Amateurs by Curtis (No. 14: 586-588). When authorship was not

indicated, these were usually abstracts from other reports, notices or short notes; a

number by Ridley himself were indicated with the initials “H.N.R.” at the end, such as

his essay on Volatile Oils (No. 9: 335-342). The topics represented a healthy interest

in all things possible to grow or transact in a tropical environment, and centred on

commercial hevea and other types of rubber (with Gutta Rambong or Ficus elastica,

the India Rubber, already mentioned in No. 5 as an interesting potential source of

commercial rubber being tried in Malacca, and which later brought Ridley there to

view trials set up by local businessman Tan Chay Yan).

The lack of a very large diversity of contributing authors probably led to E.B.

Skinner’s The United Planters’ Association, F- M.S. Report for 1901 (published in the

Bulletin No. 10: 393-400) stating that the Agricultural Bulletin should benefit from

an identified “band of contributors” that included Charles Curtis (Superintendent

of Gardens & Forests, Penang & Province Wellesley), Robert Derry (then Assistant

Superintendent of Forests, partly in Malacca, partly in Perak; subsequently also a

Curator at the Botanic Gardens in Singapore), Leonard Wray (Curator of the Perak

State Museum), Stanley Arden (an official of the Agricultural Department in Selangor),

the Chief Forest Officer (Colony & F.M.S.) and E.V. Carey (Chairman of the Planters’

Association). Aside from mentioning complaints on typographical errors, the report

also recorded that the Governments of Colony and the F.M.S. “each promised a grant

of $300 per annum towards the expenses of publishing the Bulletin.”

Timeliness of the issues in this second series was generally well observed. As

Vol. 1 began in October 1901, it went into No. 14 in December 1902; thereafter, from

Vol. 2 onwards, each volume began with No. | in January and ended with No. 12 in

December of a calendar year. In all, ten volumes were published, ending with Vol. 10,

No. 12 in December 1911. Ridley was the editor in the main, although Volumes 5—7

were credited as being edited by H.N. Ridley (Director of Botanic Gardens, S.S.) and

J.B. Carruthers (Director of Agriculture & Government Botanist, F.M.S.). Nonetheless,

the editorial in Vol. 5 No. 1 (January 1906) still used the singular form:

The Editor would always be glad to receive correspondence or notes on Agricultural or

Horticultural subjects...He would call the attention of planters and others to the fact that

when the Bulletin was started in its present form, many were the promises of support in

these matters. During the past year hardly a note has been sent for the Bulletin...

Following this lament, most of the writing was still Ridley’s. Carruthers left the F.M.S.

for a position in Trinidad in March 1909, returning the editorship solely to Ridley.

This Bulletin series recorded many notable snippets of information, such as

6 Gard. Bull. Singapore 64(1) 2012

Ridley’s Malay Drugs (Vol. 5 No. 6: 193-206, No. 7: 245-254, No. 8: 269-282) and

others like Fall of Hail in Ulu Langat, Pandan Hats, Note on the Method of Preparing

Dragons Blood, A Nest of Termes malayanus with many Queens, etc. He had many

agriculturally inclined articles too, among which were: Pine-Apples (Vol. 3: 1-6) and

Pineapple Cultivation (Vol. 3: 37-40), the latter heralding an increasing interest in the

crop that was eventually, in the 1970s and 1980s, to become a significant feature of

the Johor agricultural landscape in south Peninsular Malaysia, before giving way to

oil palm. As for the beginnings of commercial interest in oil palm, Ridley (1908: 4)

noted that “There was a demand for seeds of the Oil Palm (E/aeis guineensis) due to an

article in the ‘Agricultural Bulletin’ of this year [1907]” pointing out the value of this

plant in cultivation. Articles such as A remarkably prolific coconut (Vol. 8 No. 7, July

1909) (Fig. 2) provided incomparable interest in the unusual, yet possible, obtainment

of extreme productivity, of incredible attraction to the agriculturist’s essentially

insatiable expectations.

And Ridley did entertain all manner of agricultural correspondence, including

offering advice on ridding soil of rhinoceros beetle larvae (Vol. 3 No. 1: 18-19). The

Director’s interest in his Gardens was not forgotten, and there were also articles of

practical value there, such as The Palm Collection of the Botanic Gardens, Singapore

(Vol. 3 No. 7: 249-266), which recorded 236 species in 90 genera, probably the finest

in Malaya. Later (March 1910), there was occasion for disappointment, when Ridley

reported The Abolition of the Botanic Gardens of Penang (Vol. 9 No. 3: 97-105): plans

were afoot to convert the Gardens in Penang into a reservoir, which, however, did not

materialise.

Undeniably, there was an overwhelming interest in rubber in the Agricultural

Bulletin of that time. In May 1906 (Vol. 5 No. 5), The United Planters’ Association

Report for 1905 declared for rubber thus—’The triumphant progress of this part of

our Agricultural Industry has continued unslacked. The fame of the F.M.S. as a rubber

producing country is spreading far and wide...” Many articles appeared that were

concerned with the properties, cultivation and tapping of rubber, industrial processing

and attendant machinery, and trade in the commodity. For the 2nd International

Exhibition of Rubber and Allied Trades 1911 in London (24 June to 11 July, 1911), a

major event, it was noted that “Mr Ridley promised to be responsible for the supply

of stumps to show the methods of tapping” and “...to write a pamphlet dealing with

the history of rubber in Malaya” (Vol. 10 No. 1: 11-12). In March 1911 (Vol. 10

No. 3), Ridley even describes the operation of his trial Rubber Smoking House: “...1

will first describe the smoking house in the Botanic Gardens, which has proved quite

satisfactory and economical. The building is 55% feet long and 19 feet wide, oblong in

shape, and made of ordinary planking with a high roof...”

So Mr Ridley was evidently kept rather busy. He was also editor of the Journal

of the Royal Asiatic Society, Straits Branch, from 1889 to 1911, as well as secretary of

the Society, and published many taxonomic papers in that journal as the Agricultural

Bulletin was so devoted to economic botany. The Agricultural Bulletin included a

lighter side with snippets that must have involved some straight-faced moments,

such as with The Mosquito Plant, Ocimum viride (Vol. 3 No. 1: 24), in which Ridley

Centenary of Gardens’ Bulletin, Singapore

> 3.

‘

AR Aas, =)

Fig. 2. A photo by W.J. Gallagher, Government Mycologist, F.M.S., showing “...an exceedingly

prolific coconut tree grown on Klanang Estate, Jugra in Selangor...only eleven years old and

the total number of nuts on the tree...was more than three hundred and sixty.” From H.N.

Ridley (1909) A remarkably prolific coconut, Agricultural Bulletin of the Straits and Federated

Malay States 8(7), facing p. 318.

8 Gard. Bull. Singapore 64(1) 2012

discusses his own simple observations:

The mosquitoes in fact quite ignored the mosquito plant and took no notice of it at

all...A writer...rubbed his face and hands with the juice of the leaves. This he found

effectively kept the mosquitoes off, but he found next day that he had developed a rash...

and eventually for five days his face and hands were as if badly scalded, and he came to

the conclusion that the evil was preferable to the remedy...Editor.

One or two inconsistencies were also not unexpected. In Ridley’s Curious root-

development of Albizzia, Vol. 7 No. 4 (erroneously printed as Vol. 6 No. 7), April 1908,

the article starts by describing roots of A/eurites moluccana ascending an oil-palm

tree, but ends by referring to the tree as an A/bizzia as in the title of this short note.

Albizia (or ‘Albizzia’) moluccana was the name used then for the leguminous tree

we now know as Falcataria moluccana (with Albizia falcataria and Paraserianthes

falcataria to add to a colourful synonymy that keeps foresters in awe), and Aleurites

moluccana is the candlenut tree known for its seed oil. Clearly, even the taxonomist

must watch their very busy moments.

In April 1909, Vol. 8 No. 4: 169 carried Ridley’s obituary note on Sir George

King, late Director of the Botanic Gardens at Calcutta and Director of the Botanical

Survey of India, who retired in 1898:

... Sir George King was also the Author of...Materials for a Flora of the Malay Peninsula,

which is as yet unfinished...It was originally intended that he in collaboration with Sir

Joseph Hooker should publish the complete Flora of the Malay Peninsula, but his death

has prevented this from being carried out.

Ridley (1907) had, himself, treated the monocots in three volumes (‘Parts’), although

the rest of the Materials was never completed, comprising 25 instalments that

appeared in the Journal of the Asiatic Society of Bengal between 1889 and 1915, with

a 26th instalment only in 1936 (Ng & Jacobs 1983). Thus around this time, Ridley had

already been active collecting botanical specimens (with many cited in the Materials)

as well as compiling a flora. As it turned out, the Materials was to pave the way

for Ridley’s five volumes of The Flora of the Malay Peninsula (Ridley 1922-1925),

which, although it does include a number of astute insights into the classification of

Malayan plants, is considered by some scholars as having some apparently hastily

written and poorly compiled parts. Given King’s uncompleted account (continued to

some extent by J.S. Gamble), Ridley’s work pressures, the imperatives of having the

rich Malayan flora surveyed as a taxonomic package, and the fact that he could only

settle in to compile his Flora at Kew following retirement, a rapid pace of completion

was probably not to be compromised. Even then, the onset of the Ist World War around

1914 had disrupted life. Whatever the criticisms—and awareness of many weaknesses

came as soon as the Flora appeared: see Holttum (1959)—Ridley’s completion of a

primary documentation of the Malayan flora was a feat in itself and an accomplishment

in discipline. He would be admired for a great many things (Purseglove 1955a, b, c).

Purseglove (1959) also commented that Ridley “belongs to that great pre-specialised

age of scientific natural history and he collected and studied many animals, as well as

distributing specimens and writing about them.”

Malaya entered a great rubber boom in 1910. Advertisements apparently made

their debut in Vol. 8 No. 12 (December 1909), when four items were carried, from

Centenary of Gardens’ Bulletin, Singapore 9

Howarth Erskine Ltd., Singapore, “makers of rubber machinery”; The Borneo Co.,

Ltd.; the Journal d’Agriculture Tropicale; and the Sun Life Assurance Company of

Canada. These were vignettes of what the region was getting busy with at the time:

rubber and agriculture, development and business. In the last two volumes 9 and 10

of this second Agricultural Bulletin (1910 and 1911), advertisements had gained in

number conspicuously. This was a notable change, a kind of culmination to a very

regimented bulletin production that highlighted economic successes through the

agricultural programme, more possibilities with potentially profitable crops, and a

necessary brevity pervading throughout because of the need to bring issues out very

frequently.

The Third Series: birth of the Gardens’ Bulletin

Ridley retired officially on March Ist, 1912 but had gone on leave by January 18th

(Burkill 1913). The Agricultural Bulletin of the Straits and Federated Malay States,

Third Series, was introduced in this year, somewhat awkwardly, by its No. | comprising

a 66-page Index for articles and topics of the past Bulletins from 1891 to 1911. In the

Index, ‘OS’ stood for the “Old Series’ and ‘1’ to ‘10° for volume numbers of the “New”

(i.e.. Second) Series. Then No. 2 began anew with Page 1. The numbers 1-5 appeared

monthly for January to May 1912 and had lead pages declaring Founded by H.N.

Ridley, C.M.G., M.A., F-R.S., & c., in 1891, and edited by him up to 1911. The Assistant

Curator of the Garden, J.W. Anderson, compiled Nos. 1, 2 and 5, and the Curator, R.

Derry, compiled Nos. 3 and 4. But not long afterwards, the new Director of Gardens,

Isaac Henry Burkill (1913) (Fig. 3) recorded that “In consequence of want of staff the

‘Agricultural Bulletin’ was suspended after five numbers had been issued.”

In the Planters’ Association of Malaya Fifth Annual Report for the year

ending 31st March, 1912 published in the Bulletin (3rd Series) Vol. 1 No. 4: 137-142,

an appreciation of Ridley as well as new arrangements for the Bulletin itself were

recorded:

MR. RIDLEY.—This gentleman, who was truly the Father of the Rubber Industry in this

Peninsula, has retired and left the East, after having devoted many years of his life to

benefitting the planting industry...

AGRICULTURAL BULLETIN.—It is a great satisfaction to hear that arrangements

have been made for this paper to be edited and published in Kuala Lumpur, as it is one

of great use to the Planting Community.

With the development of commercial activities increasing in much of Malaya,

organisation to serve the F.M.S. better was being put into effect. The Singapore Botanic

Gardens had been an instrument of the S.S. Government for supporting its agricultural

development since 1875 and had practically been concerned with the whole of Malaya,

but in 1905 the F.M.S. had set up a Department of Agriculture headquartered in its

capital, Kuala Lumpur. Agriculture was enjoying a boom. It was only after I.H. Burkill

arrived as the new Director on October 17th, 1912, that new arrangements for the

publication of the journal were ironed out. This Bu//etin would continue its No. 6

Gard. Bull. Singapore 64(1) 2012

Fig. 3. The second Director, I.H. Burkill, who consolidated the Gardens ’ Bulletin as a botanical

journal. (From the Singapore Botanic Gardens archives)

Centenary of Gardens’ Bulletin, Singapore 1]

under a new name, based in Singapore, while the Agricu/tural Bulletin of the F M.S.

would be a different journal, with Kuala Lumpur as its centre of organisation.

Under these circumstances, No. 6 of the Bulletin in Singapore appeared as

The Gardens’ Bulletin, Straits Settlements “into which is incorporated all that has been

published as the Third Series of the Agricultural Bulletin of the Straits and Federated

Malay States” (Fig. 4). The Editor’s Note at the start of Vol. 1 No. 6 (issued December

15, 1913) makes the clarification thus—

Since 1891 the Botanic Gardens have had a publication; at first it was an occasional

publication, then in 1902 it became a monthly, by a joint agreement with the Governments

of the Straits Settlements, and Federated Malay States and the United Planters’

Association of Malaya. Now, again, the period of the agreement having terminated, it

will be occasional...the title has been changed to avoid confusion with the Agricultural

Bulletin of the Federated Malay States. It is thought that the title “Gardens’ Bulletin,

Straits Settlements” is satisfactorily descriptive, distinctive and conveniently short... The

five parts published as the third series of the Agricultural Bulletin, Straits and Federated

Malay States, become the first five parts of the Gardens’ Bulletin... There will be more

original matter...but no market reports and no proceedings of meetings...

Having taken over from his predecessor, Burkill’s writing featured

conspicuously in the pages of the Bulletin from 1913 into the 1920s. The immediate

issue, No. 6, opened with his note on The Coconut Beetles, Oryctes rhinoceros and

Rhynchophorus ferrugineus. Original material came to feature more prominently,

mostly emphasising economic botany, including large summaries like The treatment to

which the Para Rubber trees of the Botanic Gardens, Singapore, have been subjected

(Vol. 1 No. 8: 247-295) and, conspicuously, Burkill’s own research into yams, e.g.,

Experimental cultivation of the Greater Yam Dioscorea alata (Vol. 1 Nos. 9, 11-

12; Vol. 2 No. 2), Some cultivated Yams from Africa, and elsewhere (Vol. 2 Nos. 3,

12), A progress report on the cultivation of the greater yam, Dioscorea alata—in

the Botanic Gardens, Singapore (Vol. 2 No. 4), Yields of the lesser vam and of some

African yams (Vol. 2 No. 5), and A list of Oriental vernacular names of the genus

Dioscorea (Vol. 3 Nos. 4-6). Instalments on The Oil Palm (Elaeis guineensis) in the

East by E. Mathieu appeared in Vol. 2 No. 7: 217-230 and No. 8: 265-275. Articles

on lima bean acclimatisation trials, roselle and castor oil cultivation by E. Mathieu,

on races of the coconut palm by Ahmed Bin Haji Omar, and the betel palm Areca

catechu by F. Flippance were included. More notes and data were published on agri-

horticultural pests (Locusts in Malacca by P.C. Cowley-Brown & I.H. Burkill, Vol. 1

No. 10; Catochrysops pandava, a butterfly destructive to Cycads by Burkill, Vol. 2 No.

1; and various notes on beetle pests of the coconut by Professor C.F. Baker, Vol. 2 No.

1) and fungi by Baker (Host Index for fungi, Vol. 2 No. 1; Hevea versus fungi, Vol. 2

No. 4) and T.F. Chipp (The fungus flora of Hevea brasiliensis, Vol. 2 No. 6; A Host

Index of Fungi of the Malay Peninsula, Vol. 2 Nos. 7, 8; A list of the fungi of the Malay

Peninsula, Vol. 2 Nos. 9, 10 & 11 combined).

Early on in his tenure as Director, Burkill gathered summaries of the

development of the Botanic Gardens. He himself compiled The establishment of the

Botanic Gardens, Singapore (Burkill 1918a) and The second phase in the history of

the Botanic Gardens, Singapore (Burkill 1918b). Flippance contributed 4 Guide to the

12. Gard. Bull. Singapore 64(1) 2012

JEals

—

GARDENS’ BULEE TIN

STRAITS SETTLEMENTS.

INTO WHICH IS INCORPORATED ALL THAT HAS BEEN PUBLISHED

AS THE THIRD SERIES OF THE AGRICULTURAL BULLETIN

OF STRAITS AND FEDERATED MALAY STATES.

The Gardens’ Bulletin will be published as material becomes:

available. Its price is fifty cents a copy, post free, or in

advance for a volume of twelve numbers, post free,

Five dollars in the Straits and Federated Malay States

Nine and a half rupees in India and Ceylon,

Thirteen shillings in Europe.

Subscriptions paid to the third series of the Agricultural

Bulletin, Straits and F. M.S. are counted as subscriptions to it.

ee [ay

SINGAPORE:

THE STRAITS TIMES PRESS, LIMITED,

Fig. 4. The title page of the December 1913 issue No. 6 announcing the journal’s name change

to Gardens’ Bulletin, Straits Settlements.

Centenary of Gardens’ Bulletin, Singapore 13

Palm Collection in the Botanic Gardens (Vol. 2 No. 6: 177-186 and No. 7: 246-258).

Burkill’s interest in more organised platforms for the continuing botanical survey of

the Malayan flora is seen in his Fragments of Malayan Geographic Botany No. 1.

Enumeration of Pahang Plants collected by the late A.M. Burn-Murdoch (Vol. | No.

9): The as-yet botanically unexplored parts of the Malay Peninsula (Vol. 3 Nos. 1-3);

and the classic guide to Botanical Collectors, Collections and Collecting Places in

the Malay Peninsula published in Vol. 4 Nos. 4 & 5 after retirement (Burkill 1927),

which organised its content using a system of grid-square references systematically

enumerating the collections known to have been made in particular localities in each

state of Malaya.

A special interest in orchids was also noticeable. There were Malayan

orchid notes by Burkill and Mohamed Haniff (Vol. 1 No. 10, July 1916), the latter a

botanically experienced Overseer in charge of the Waterfall Gardens in Penang and

under the Director in Singapore; a paper by Burkill discussing evidence produced to

show that Pigeon Orchid (Dendrobium crumenatum) flowers about 8 days after heavy

rain (Vol. 1, Nos. 11-12, March 1917), following Rutgers and Went’s observations of

its gregarious flowering in the Annales du Jardin Botanique de Buitenzorg (29: 129-

160) the previous year; a general article on the flowering of orchids by Burkill (Vol. 2

No. 2); and notes on Southeast Asian orchids by Burkill (Vol. 2 No. 12, August 1921).

These and the earlier orchid collections and notes by Ridley were precursor to the

interest in Malayan and British North Borneo orchids taken up by the Malayan rubber

planter C.E. Carr into the 1930s (Carr had been a frequent associate at the Singapore

Botanic Gardens, visiting to study orchids and contributing specimens; he died in New

Guinea while returning from an expedition in 1936) (Holttum 1959).

Burkill also put out articles on biological and ecological perspectives. These

included Some notes on the pollination of flowers in the Botanic Gardens, Singapore,

and in other parts of the Malay Peninsula (Vol. 2 No. 5: 165-176, September 1919);

and his The fertility of branched coconut palms (Vol. 3 Nos.1-3) was followed by

C.X. Furtado’s 4 study of the coconut flower and its relation to fruit production and

Branched coconut palms and their fertility (Vol. 3 Nos. 7-8). Early ecological studies

included The composition of a piece of well-drained Singapore secondary jungle thirty

years old (Vol. 2 No. 5: 145-157, September 1919) that enumerated trees by species

with height and girth data: and Forests and their retention of rain water (Vol. 2 No.

12: 419-421). Compiled together with Richard Eric Holttum, his Assistant Director

who was appointed in 1922, A botanical reconnaisance upon the Main Range of the

Peninsula at Fraser Hill (Vol. 3 Nos. 1-3, August 1923) would be an indispensable

record of not just seed plants, but also ferns, lycophytes and mosses; this and the

included survey of the largest tree sizes on forested ridges by the Semangkok pass

(later to inspire the keeping of the so-called Big Tree Plot as a Virgin Jungle Reserve

by the Forestry Department: Wyatt-Smith 1950) thus comprise an excellent baseline

study of the flora of Fraser Hill. Holttum continued this foray into floristics, as with The

vegetation ef Gunung Belumut in Johore (Vol. 3 Nos. 7-8). But Burkill's study with

Murray Ross Henderson of The flowering plants of Taiping, in the Malay Peninsula

(Vol. 3 Nos. 7-12) was considered by them as “the first local Flora for any area within

14 Gard. Bull. Singapore 64(1) 2012

the Federated Malay States” and only the third after Charles Curtis’s Catalogue of the

Flowering Plants and Ferns found growing wild in the Island of Penang (Curtis 1894)

and Ridley’s Flora of Singapore (Ridley 1900).

Thus a conspicuous diversity of topics and authorship, both agri-horticultural

and related to the natural history and botany of Malaya, much of it original

contribution, began building up. The bulletin that Ridley founded had been developed

into a fully scientific journal by I.H. Burkill, his successor. The earlier preoccupation

with agri-horticultural news and developments and commodity reports, when the only

agricultural bulletin serving Malaya was co-sponsored by a planters’ association and

required Ridley to maintain that coverage, was now largely replaced by scientific

notes, reports and papers on a wider range of subjects. The need to sustain monthly

issues, and obligations to advertisers who no doubt expected consistent and regular

circulation, had been relieved by the calmer, more research-based attitude with the

change to the Gardens’ Bulletin, Straits Settlements. Perhaps also the development of

the F.M.S. Department of Agriculture in the beginning 20th century, with a growing

specialisation in technical subjects, was as important as the reformulated objectives of

the new Gardens’ Bulletin in bringing greater opportunities for organised research and

scientific publishing for both the agricultural and botanical sciences. The agricultural

component had all but disappeared from the Gardens’ Bulletin by 1924.

Colonial reorganisation, economic slump, War,

and another name change

In 1918, F.W. Foxworthy, who had served with the Bureau of Science in Manila, was

appointed the first Forest Research Officer of the F.M.S. and S.S., stimulating further

growth of the fledgling Forest Herbarium in Kuala Lumpur (Wong 1987). Around

this time also, other developments pressed for centralisation of botanical research

in Kuala Lumpur but this did not materialise, as discussed by Humphrey Morrison

Burkill (1983), late Director of the Singapore Botanic Gardens, and summarised here.

A proposal to the Colonial Office in London for recruiting a Systematic Botanist in

the Museum Department in Kuala Lumpur was made in 1920 by Sir George Maxwell,

Chief Secretary of the F.M.S. and in 1921, M.R. Henderson was appointed to this

post. As the development of both agriculture and forestry became more organised and

important in Malaya, the colonial government decided to centralise botanical research

in Kuala Lumpur to better support these areas. A meeting in December 1923 attended

by G.E.S. Cubitt (Conservator of Forests, S.S. and F.M.S.), A.S. Haynes (Secretary

for Agriculture, S.S. and F.M.S.) and I.H. Burkill discussed arrangements to move

the Singapore Herbarium and the research of the Singapore Botanic Gardens to new

premises in Kuala Lumpur, to absorb the botanical research of the Museum Department

and its Systematic Botanist into the new botanical department, and to develop the

Public Gardens in Kuala Lumpur as a botanic garden with the existing ones at

Singapore and Penang as branch gardens. In the F.M.S., administrative dithering led to

a delay in implementation and Maxwell only agreed to the new botanical department

Centenary of Gardens’ Bulletin, Singapore 15

in Kuala Lumpur in 1926. The rubber slumps of 1921 and 1924 preceded a more

serious worldwide slump in 1929 and brought adverse consequences for government

spending in colonial Malaya. The Museum Department’s Systematic Botanist post was

abolished and by I.H. Burkill’s intervention, Henderson was brought to the Botanic

Gardens, Singapore as Curator of the Herbarium in 1924. No further planning for the

development of the Kuala Lumpur Public Gardens appears to have been undertaken,

and the move to bring Singapore-based botanical resources to Kuala Lumpur was

abandoned (Burkill 1983).

The effort to provide more specialist services in forestry research in Malaya,

meanwhile, was assisted when in 1925 the Regent of Selangor approved the

allocation of 800 acres at Kepong, near Kuala Lumpur, for the establishment of the

Forest Research Institute (Watson 1950, Menon 1969). Its nursery and experimental

plantations were begun on site in 1926, and the main building was constructed in

1929. Parallel to these developments, Ridley’s Flora of the Malay Peninsula, in five

volumes (Ridley 1922-1925), was published, having occupied his main attention

based at the Kew Gardens following retirement. I.H. Burkill, the second Director who

so ably crafted a more focussed scientific direction for the Singapore Botanic Gardens,

retired in February 1925, and concentrated on writing and compiling 4 Dictionary of

the Economic Products of the Malay Peninsula (Burkill 1935), which would appear

a decade following Ridley’s Flora (Furtado & Holttum 1960). Even so, in the years

that followed, Burkill’s notes and papers continued to appear in the Gardens’ Bulletin,

including more notes on yams and various other plants, with taxonomic notes and

revisions, and even ethnobotanical notes on The Chinese Mustards in the Malay

Peninsula and Cosmos in the East (Vol. 5 Nos. 3-6, June 1930).

R.E. Holttum, Assistant Director to I.H. Burkill, became Director in 1925 and

there was some expectation that his work on ferns would lead the preparation of a

cryptogam flora for Malaya to complement Ridley’s seed plant flora. Indeed, work

by Holttum and the Danish Carl Christensen on ferns began appearing during this

period, including an account of Mt Kinabalu ferns in Vol. 7 Part 3, June 1934. But in

the tropics it is seldom possible to stay on one study and not notice the many other

manifestations of nature and life. Holttum’s classic studies of plant phenology, On

periodic leaf-change and flowering of trees in Singapore (Vol. 5 Nos. 7 & 8, June

1931; Vol. 11 Part 2, November 1940) and The flowering of Tembusu trees (Fagraea

fragrans Roxb.) in Singapore 1928-1935 (Vol. 9 Part 1, December 1935) were thus

inevitable.

Also, new work in any part of the flora being re-examined was accruing

additions and changes easily. This was certainly true with specialist work on the

Dipterocarpaceae, the predominant big-tree family in Malayan forests, by C.F.

Symington, who joined the Forest Research Institute in 1929 (his Notes on Malayan

Dipterocarpaceae | to V in the Bulletin spanning 1933 to 1939). It also applies to the

research on palms and aroids by C.X. Furtado (who joined the Singapore Botanic

Gardens as Field Assistant in 1923, then later was Botanist) (Alphonso 1980); new

discoveries or revisions of orchids by C.E. Carr; and the research on ferns and orchids

by Holttum; and their other colleagues. Henderson also compiled The Flowering

16 Gard. Bull. Singapore 64(1) 2012

Plants of Kuala Lumpur, in the Malay Peninsula (Vol. 4 Nos. 6-10, January 1928)

and specially conducted surveys of the Malayan limestone flora. As botanical work

intensified and got the attention of more specialists, not least because Ridley’s Flora

provided an accessible and organised overview, the Malayan flora was fast gaining

additional interest. Henderson, who re-arranged the Herbarium following Ridley’s

Flora, had Additions to the Flora of the Malay Peninsula in Vol. 4 Nos. 2 & 3 (March

1927) and Nos. 11 & 12 (January 1929) (with Furtado), and further additional listings

in Vol. 5 Nos. 3-6 (June 1930) and Vol. 7 Part 2 (May 1933). E.D. Merrill, too, had

Additions and corrections to Ridley’ Flora of the Malay Peninsula (Vol. 8 Part 2,

January 1935).

E.J.H. Corner, who arrived in Singapore as Assistant Director of the Botanic

Gardens in 1929 and a mycologist, would not be able to resist an interest in the

palms and trees of Malaya. His Notes on the systematy and distribution of Malayan

phanerogams and taxonomic work on Ficus began appearing in the Gardens’ Bulletin

Vol. 10 (1939) and continued 1960-65 from the University of Cambridge. His debut

publication in the Bulletin was The identification of the Brown-root fungus (Vol. 5 No.

12, June 1932). In this paper, Corner’s special acumen for graphical representations

showed clearly in his diagrams of microscopic structures. His later work would show a

special talent for watercolour illustration of mushrooms to represent subtle differences

in tone, as well as the classic line-drawings of plant parts, trees and other figures made

famous through his Wayside Trees of Malaya (Corner 1940), and even landscapes

(Corner 1965, Mandalam 2011).

The Gardens’ Bulletin was fast taking on a distinct systematic slant. Furtado

added an interest in botanical nomenclatural matters, fuelled by his interaction with

Professor H. Harms from Berlin, editor-in-chief of the International Rules of Botanical

Nomenclature, ed. 3. There was an apparent burst of productivity over Vol. 5 (August

1929-June 1932) and Vol. 6 (1929-1930). The overlap is unusual and is likely due to

the availability for the latter volume of three sizeable specialist accounts on medicinal

plant use coordinated by I.H. Burkill after settling into his retirement: David Hooper’s

On Chinese Medicine: Drugs of Chinese pharmacies in Malaya, Burkill & Mohamed

Haniff’s Malay Village Medicine, and J.D. Gimlette’s edition of an 1886 translation

of The Medical Book of Malayan Medicine with botanical determinations by Burkill.

Still, the frequency of volumes was essentially occasional. Volume 7, for example, had

three parts, one each in 1932, 1933 and 1934. Furtado was on half-pay for a year from

April 1933 to May 1934, travelling through Europe to work on the palm collections

of key institutions. He was mainly at the Berlin Botanical Garden in conjunction

with Professor M. Burret (then the leading palm specialist, who assembled the type

specimens of Martius from Munich and other material for Furtado’s studies); and then

London, Vienna, Florence and Paris (Johnson & Tay 1999).

Vol. 9 Part 1 (December 1935) was dedicated to Ridley for his 80th birthday

(Fig. 5): “...Few men have accomplished so much in 23 years of tropical service, and

few have been able to complete their work after retirement as Mr. Ridley has completed

it.” B.J. Eaton, Director of the Rubber Research Institute of Malaya, observed that

“In addition to continuing his publications on the flora of Malaya, he [Ridley] is still

Centenary of Gardens’ Bulletin, Singapore 7)

a member of the Technical Sub Committee of the London Advisory Committee for

Rubber Research (Ceylon & Malaya) and thus continues at the age of 80 to maintain

his interest in an industry of which he may be said to have witnessed the birth.”

The “War volume”, Vol. 11, is of special significance. It had Parts 1-4,

spanning May 1939 to September 1947. World War II arrived with the Japanese

military in Singapore in 1942, so the first three parts (Part 1, May 1939; Part 2,

November 1940; Part 3, August 1941) were still issued as the Gardens’ Bulletin,

Straits Settlements. Following the Japanese surrender and the return of Allied forces to

Singapore, the political push for a Malayan Union (comprising Penang and Malacca

from the former S.S., the F.M.S. and other states of Malaya, but excluding Singapore)

was consolidating. It was only in September 1947 that Part 4 was issued to complete

the volume, under the new name Gardens’ Bulletin, Singapore. An insert, taking the

place of pages 261-262, before the start of Part 4 proper on page 263, announced the

consequence of the new political imperative:

CHANGE OF TITLE

The Colony of the Straits Settlements has

ceased to exist. The title of this Bulletin

is now therefore changed, but the present

issue is a direct continuation of the former

Gardens’ Bulletin, Straits Settlements, of

which the last issue was Vol. XI, part 3,

published 30 August 1941.

Part 4 of Vol. 11 opened with a crisp record of events at The Singapore Botanic

Gardens during 1941-46. At the time of the War, J.L. Pestana (Laboratory Assistant),

J.C. Nauen and G.H. Addison (Horticultural Officers) joined the defence forces,

were taken prisoner and sent to the Siam-Burma railway, where Nauen died in 1943.

Henderson was evacuated from Singapore and later worked in the National Botanic

Garden at Kirstenbosch in South Africa. Holttum, Corner and Furtado remained in

Singapore during the Japanese occupation. The Herbarium and most of the Gardens

remained undamaged, as Professor Hidezo Tanakadate of the Tohoku Imperial

University assumed control of the Gardens and Holttum was retained in executive

charge. In December 1942, Kwan Koriba, who had served as Professor of Botany

at the Imperial University of Kyoto, was made Director of the Singapore Botanic

Gardens. [Koriba was interested in physiological and ecological factors influencing

plant morphology, especially flower and leaf disposition, and had done some work

on orchids previously, although this was not highlighted in Singapore then (Arditti

1989).] Following Japanese surrender, the Gardens were placed under British Military

Administration between September 1945 and March 1946, and in May 1946, Holttum

returned to duty as Director, a post he held until 1949, when he became the inaugural

Professor of Botany in the University of Malaya. Henderson became his Assistant

Director. After war ended, Corner left for Latin America in 1947 on UNESCO service

(thence to a lectureship in taxonomy at the Botany School in Cambridge University,

becoming professor in 1965).

18 Gard. Bull. Singapore 64(1) 2012

Fig. 5. Ridley close to his 80th birthday. Gardens’ Bulletin, Straits Settlements Vol. 9 Part 1,

facing page |.

Centenary of Gardens’ Bulletin, Singapore 19

Koriba had returned to Japan in 1946 and retired to publish two books, then

became the President of Hirosaki University in 1954. Koriba’s research undertaken

while at Singapore, On the periodicity of tree growth in the tropics, was published in

the Gardens’ Bulletin, Singapore Vol. 17 Part 1, November 1958, following his death

in December 1957. In an obituary note, Holttum (1958b) wrote:

At Singapore...[Koriba] was dependent on the military organization for funds and

supplies of all kinds, and took every opportunity of securing such amenities as were

possible for the gardens staff. On several occasions he took energetic action to prevent

encroachment on the Nature Reserves of Singapore...He was also greatly concerned that

the herbarium and library at the Gardens should be maintained intact. To his single-

minded devotion to botanical science the Singapore Botanic Gardens owes much...

In the author’s preface to A Revised Flora of Malaya, Vol. I. Orchids of Malaya

(Holttum 1953), Holttum recorded:

The main part of the work of the preparation of this book was carried out during the

Japanese occupation of Singapore... The fact that I was able to undertake the work in the

years 1943-1944 was due to the courtesy of Dr. Kwan Koriba, who was sent...to take

charge of the Botanic Gardens here. I wish to express my most grateful thanks to Dr.

Koriba, for the courtesy with which he allowed me complete freedom to continue my

studies, and for much personal kindness during that period.

The sanction of Koriba’s authority and that of his superiors, and the wealth of research

materials already at the Botanic Gardens, had permitted some intensive preparation for

botanical accounts that included The Zingiberaceae of the Malay Peninsula (Holttum

1950) and Ferns of Malaya, which appeared as Vol. 2 of the Revised Flora series

(Holttum 1954).

The botanical spirit was refired at the Gardens. There was a great deal more

to understand about the flora of Malaya and the region, and plenty to discover. To this

renewal James Sinclair arrived at the Singapore Botanic Gardens in 1948, as Curator

of the Herbarium, a post that was re-titled Keeper of the Herbarium in 1955 and

Botanist (Keeper of the Herbarium) in 1960. The research programme when Sinclair

arrived was to prepare a revised Flora of Malaya, for which he was asked to research

the Annonaceae.

There is no telling how well a centralised botanical research facility of the

S.S. and F.M.S. in Kuala Lumpur would have withstood the War, had earlier colonial

plans been realised for bringing Singapore-based resources to Kuala Lumpur, given

the looting and other damage inflicted on the Forest Research Institute during the

Occupation (Wong 1987). In contrast, the research facilities of the Singapore Botanic

Gardens were reasonably well preserved (Corner 1946, 1981; Holttum 1958b). On

hindsight, it does seem fortunate that the merger never took place, as otherwise the

overall damage to herbarium resources could have been even greater. And, of course,

the Gardens’ Bulletin, Singapore may never have seen its naming as such and its

continuity into the present time.

20 Gard. Bull. Singapore 64(1) 2012

The 1950s and 1960s: two centenaries and increasing regional focus

Ridley’s hundredth birthday was celebrated by the Botanic Gardens on 10th

December, 1955. According to the Director John William Purseglove (1959), there

was “an exhibition of Ridleyana and current work, while the Gardens were floodlit

for one week. A special brochure for private circulation was produced for the occasion

[Purseglove 1955c].” Ridley wrote “It is a great delight to me to have lived to see the

Gardens, the best tropical Gardens in the world,” and that his hundredth birthday was

one of the most enjoyable days of his life (Purseglove 1959).

In the Bu//etin issue marking a hundred years of the Singapore Botanic Gardens

(Vol. 17 Part 2, December 1959), messages and reflections from former Directors

I.H. Burkill (1912-1925), R.E. Holttum (1925-1949), J.W. Purseglove (1954-1957),

and the then Director H.M. Burkill (who first was Assistant Director from 1954),

the son of I.H., the father then 89, were presented alongside those from many other

botanical figures (Purseglove’s was reprinted from a 1957 article). It would appear

to be something of a feat that four Directors had their writings in the same issue,

including both Burkills. H.N. Ridley, the first Director, had died just three years before

in 1956, aged 101. And M.R. Henderson, Director during 1949—54 and having retired,

had severed most professional contacts and gone to live in the Scottish hills (Burkill

1983)):

Although the Gardens were a hundred years old, Purseglove (1959) saw it fit

to reiterate their essential features, aware of the risk of repeated opportunities for their

erosion. He took the trouble to emphasise—

...botanic gardens, to merit the name, are gardens maintained for the scientific study of

the plants. As soon as this vital function is neglected botanic gardens change to public

parks... The Singapore Botanic Gardens are the last gardens in the British tropics which

function as a separate and self-contained department and have never been under the

control of any Agricultural Department, Municipality or University... The southern end

of the Gardens was probably abandoned gambier land, a haunt of tigers, while the north

was still virgin tropical evergreen rain forest, 11 acres of which are still preserved...a

most valuable asset in the centre of a great city.

This article by Purseglove (1959) was brought up to date by a footnote inserted

by H.M. Burkill, who recorded thus:

The penultimate constitutional step in the introduction of full internal autonomy of

Singapore (effected in May 1959) was operative from April 1955. As befits a country in

charge of its own affairs, the Singapore Government adopted a policy of “malayanisation”

of the public service from January Ist, 1957. J.W. Ewart retired in March 1957 in

accordance with this policy, and A.G. Alphonso, who had returned in 1956 from a two

year course of training in horticulture at the Royal Botanic Gardens, Kew, England,

was promoted Curator. J.W. Purseglove retired voluntarily in March 1957, and H.M.

Burkill became Director. H.H. Addison retired in February 1959. Chew Wee Lek was

appointed Botanist in 1956 and went to Cambridge, England, in 1957 for three years to

study for a higher degree. Lam Hin Cheng was appointed Horticultural Assistant in July

1957 and went to the Royal Botanic Gardens, Kew, for a two year course in horticulture.

The new post of Librarian was eventually filled in April 1958, and Tan Kim Ho went

to Melbourne, Australia, in 1959 for training in library management on an Australian

Centenary of Gardens’ Bulletin, Singapore 21

Government award under the Colombo Aid Programme. Attempts to fill the post of

Assistant Director were unavailing, and finally Miss Chang Kiaw Lan was appointed

Botanist (vice Assistant Director) to take up a study of mycology...The last three years,

1957 to this centenary year of 1959 have been a period of transition characterised by an

acute shortage of senior personnel through loss of qualified staff and temporary loss of

the services of officers sent overseas for training... Thus will close a century of expatriate

know-how in the senior botanical and horticultural posts, and there will open the second

century of the Gardens’ history, every bit as promising in the comity of international

botany as the first has been successful, with the majority of the senior posts held by

Malayan personnel.

Following emergence from the war, reorganisation had been intense in the

Botanic Gardens, where “garden work deteriorated generally, as more than half of the

out-door staff (49 men) were sent to work on the Siam-Burma Railway...22 of them

lost their lives” (Purseglove 1959). With senior staff having to attend to all sorts of

duties, it was then that publication lapses were especially evident and the Gardens’

Bulletin was not issued during the years 1948, 1952, 1954 and 1957, even though it

was following an occasional mode.

It was the period after the war that saw an even wider involvement in the

botany of the Southeast Asian region; this would have been reinforced as a necessity

with the consolidation of the Flora Malesiana project encompassing much of the

Malay Archipelago (Steenis 1948):

...the Flora Malesiana should embrace as wide an area as possible...no species can be

properly defined, until it has been examined in all variations induced by the differences

in climate, locality, and soil, which an extensive area affords. Also, the flora of an area

cannot be worked out thoroughly without a knowledge of the botany of the surrounding

countries (these have many plants in common), and so the greater the area encompassed,

the better it will illustrate habits, forms, and variations of the species comprised within

it. For this reason we have extended the limits of our Flora from Sumatra to New Guinea

and from Luzon to Christmas Island, Timor and New Guinea.

Although the work of Ridley had not entirely focussed on Malayan botany in isolation,

and there was in fact a fascination and interest in the plant life of the surrounding

region, the priority then was to complete a Malayan account. A wider, regional

botanical perspective became more visible and important with the work of I.H. Burkill

(who revised the Dioscoreaceae regionally for the Flora Malesiana: Burkill (1951),

in retirement and at age 81, based on earlier monographic work undertaken with Sir

David Prain) (Holttum 1967) and his colleagues.

Holttum’s concentrated experience during wartime internment in working out

the orchids and gingers was reasonably successful only because of the availability of

good living collections and specially collected specimens with good field notes and

flowering parts in spirit; these were not available to the older accounts, causing much

inaccuracy and taxonomic confusion. He was interested to research further monocot

groups in which study was similarly disadvantaged. Bamboos are notoriously difficult

because botanists have tended to name species based on flowering material, most

often produced when the living plants are lacking fresh shoots with more easily seen

features. As bamboos may tend to be in entirely vegetative (non-flowering) states

22 Gard. Bull. Singapore 64(1) 2012

for prolonged periods, the early accounts that did not record adequate features were

essentially useless as identification tools. Incorporating the missing information for an

improved understanding demands specialised collecting and recording of features, a

feat much aided by the existence of special living collections for a region’s bamboo

flora. His revision of Malayan bamboos (Holttum 1958a) had sought perspectives from

the surrounding region, including living bamboos maintained in the more established

Calcutta and Bogor collections. This seminal account in the Bulletin also included

his perspective that the historical migration of peoples through Southeast Asia could

have brought selected clones (especially the larger, more useful Dendrocalamus

and Gigantochloa taxa) to Java and Peninsular Malaysia, where they are apparently

known only in cultivation without any documented wild conspecifics. These are now

recognised as “ancient enduring clones” (Muller 1999) and there is now evidence that

some must have been selected from hybrid swarms that do naturally occur in our

landscape (Muller 1998, Goh et al. 2011). This situation is especially relevant to the

region from India and Myanmar through South and Indo-China, into the Malesian

area. Understanding these basic premises point the way to more careful approaches in

conserving valuable genetic materials selected through the ages.

There are G. robusta clumps planted in the Bogor Botanical Garden in 1844

during the time of the botanist Hasskarl that have remained alive for over 150 years (at

least, they did not die from flowering, if any) and so are good subjects for agriculture

(Wong 2004). Likewise, in discussing Orchids, gingers and bamboos: Pioneer work at

the Singapore Botanic Gardens and its significance for botany and horticulture in the

Bulletin, Holttum (1959) mentions a village bamboo from northern Malaya introduced

to the Singapore Botanic Gardens that has persisted in vegetative state for decades;

this is G. ridleyi, distinctive yet named without flowers (by Holttum to commemorate

Ridley, who introduced it). This bamboo lives even now, over a century since its

introduction, continuing its flowerless state. Holttum (Fig. 6) died in 1990, at the age

of 95.

Steenis (1959) and Lam (1959) have further highlighted the virtues of a

Flora Malesiana approach. Similarly, Corner settled into deeper research with Asian

and Australasian Ficus at Cambridge, beginning on a worldwide perspective in The

Classification of Moraceae (Corner 1962), and later led two Royal Society expeditions

to Mt Kinabalu in 1961 and 1964 (Mandalam 2005). At the same time that Sinclair

worked on Malayan Annonaceae, he reviewed material from India, Burma, Thailand,

Borneo and New Guinea, publishing papers on interesting taxa. After his monograph of

Malayan Myristicaceae (Sinclair 1958a), the taxonomic emphasis in Singapore began

shifting towards closer collaboration with the Flora Malesiana Foundation (Burkill

1968). This was reflected in his Florae Malesianae Precursores XX, XXXI, and XLII

published in the Gardens’ Bulletin, Singapore, on Gymnacranthera (Sinclair 1958b),

Knema (Sinclair 1961), and Myristica (Sinclair 1968, posthumously), respectively.

Sinclair was retired prematurely in 1963 because of the “malayanisation” programme

but re-engaged on contract until 1965, then stayed on at the Botanic Gardens as an

honorary researcher until 1967. He also completed manuscript work on Horsfieldia,

which was only retrieved from the Kew Herbarium following his death in 1968 and

Centenary of Gardens’ Bulletin, Singapore 23

Fig. 6. R.E. Holttum in 1981. (Photo by K.M. Wong)

24 Gard. Bull. Singapore 64(1) 2012

subsequently published in the Bu//etin (Sinclair 1974, 1975). Chew Wee Lek’s doctoral

dissertation work supervised by Corner resulted in the latter’s Florae Malesianae

Precursores XXXIV on Poikilospermum (Chew 1963). The Flora Malesiana effort

was also able to attract the participation of Hsuan Keng, based at the University of

Singapore, resulting in 4 revision of Malesian Labiatae (Keng 1969).

Possibilities for participating in reciprocal research support with other

countries probably became better established during this period, when H.M. Burkill was

Director of the Singapore Botanic Gardens. Chew’s research on Laportea and allied

genera (Chew 1965, 1969a, b) was enhanced by the opportunity in 1964 to visit the

herbaria of Cambridge, Kew, British Museum, Paris, Geneva, Leiden and Utrecht on a

Royal Society Nuffield Foundation Commonwealth Bursary gained through Corner’s

support. This worldwide survey even enabled Chew’s diagnosis of the new Central

American genus Discocnide! On the other hand, revisions of Petraeovitex (Munir

1965) and Symphorema (Munir 1967) by Munir Ahmad Abid from the University of

Sind, Pakistan, were possible with a Colombo Plan Fellowship through a Government

of Singapore award to visit the Singapore Botanic Gardens in 1964 and 1965.

A very broad range of papers—from cryptogamic to seed plant subjects,

covering taxonomy, morphology, evolution, physiology, ecology and conservation, and

including country-specific or more regional geographical scopes—was now carried

in the Gardens’ Bulletin, Singapore. Thus, just about anyone working consistently

with the botany of the Southeast Asian region, and occasionally elsewhere, began to

publish with the journal. Somewhat opposite in direction, there was a “malayanisation”

programme that sought to fill key positions with as many locally domiciled persons

as possible, but meanwhile, the research scope (in tune with the imperatives of

new scientific development) was going towards regionalisation and fast gaining an

international dimension.

The Gardens in a Garden City

Less than 20 years after the admonishment by Purseglove (1959) regarding maintaining

the close relationship between botanic gardens and science, further challenges were

to appear when taxonomic research waned. Chang Kiaw Lan, who obtained her

doctorate in mycology working under Corner at Cambridge, had returned as Botanist

to the Gardens in 1965 but what would be her office in the new Herbarium building

completed in 1964 was still occupied by James Sinclair (Wong 2003). Work on orchid

culture was already underway with Hardial Singh, another Botanist engaged in 1963,

but the development of facilities for Chang’s experimental work in basidiomycete

fungal development also needed to wait. In 1967, as Sinclair left, the Garden City

Campaign was emphasised and Chang was asked to take charge of advisory work under

this programme. Less than a year later she was seconded to the Primary Production

Department, who wanted to begin mushroom cultivation research. Just over a year later,

July 1969, after Chew Wee-Lek signalled there was an “acute shortage of botanists in

the Botanic Gardens” with the retirement of H.M. Burkill, Chang was returned to the

Centenary of Gardens’ Bulletin, Singapore 25

Gardens; Chew succeeded Burkill and Chang became Keeper of the Herbarium in

1970. Unprecedented for a Director of the Gardens to serve just several months, Chew

himself left Singapore in 1971.

In 1973 it was decided to merge the Botanic Gardens with the Parks and

Trees Branch of the Public Works Department to form the Parks and Recreation (P

& R) Division (Anonymous 1974); this became a full department in 1976 (Ministry

of National Development 1977). There was more reorganisation. Geh Siew Yin,

who trained under Hsuan Keng at the University of Singapore and was employed as

Botanist in 1971, was moved to more administrative duties as Assistant Commissioner

of the Garden in 1973. After Geh, no new taxonomists were recruited for another two

decades (Kiew 1999). By 1975, Hardial Singh was also transferred to non-research

duties in P & R and only Chang was left to keep the Herbarium going. All this was an

incredible rate of flux in the Botanic Gardens research outfit, which had, in the first

place, run on only a skeleton staffing of a handful of mainly taxonomic botanists.

H.M. Burkill (1993) deplored this decline and the notion that a botanic gardens of

standing could function effectively without strong research support for its collections,

education and conservation programmes.

Chang (Fig. 7) had a superb command of languages, was an able editor and

prolific correspondent who kept in touch with a great number of botanists worldwide

and their requests for information and assistance (Wong 2003), and ably coordinated the

stay and activities of visiting scientists at the Singapore Herbarium. This encouraged

many botanists internationally to contribute to the Gardens’ Bulletin. There was still

an inflow of manuscripts from former Gardens staff members, including Corner and

Holttum, and a number from Keng, Wee Yeow Chin and A.N. Rao and their students

from the University of Singapore. Corner’s The Freshwater Swamp-Forest of South

Johore and Singapore (Corner 1978) became the first supplementary issue of the

Gardens’ Bulletin, simply enumerated as “Supplement no. 1”. Also at around this

time, Keng’s series on the Annotated list of seed plants of Singapore, which would

become the longest-spanning series published entirely in the Bulletin by a single

author, appeared in 1973 (instalment I); subsequent instalments were published in

1974 (II and III), 1976 (IV), 1978 (V), 1980 (VI), 1982 (VID, 1983 (VII), 1985 (IX),

1986 (X), and 1987 (XI). [Furtado’s series from a different time, Palmae Malesicae

(with 19 instalments spanning 1934-1956) was longer but the first part was published

in the Berlin-based Feddes Repertorium, not the Gardens’ Bulletin.| The Limestone

hill flora of Malaya by S.C. Chin of the University of Malaya, which updated and

extended Henderson’s work previously, appeared in four instalments beginning 1977

and continuing in 1979 (Part II ) and 1983 (Parts III and IV). Chang’s co-editing or

editing, respectively, of the festschrifts for Corner (Vol. 29: Mabberley & Chang 1977)

and Holttum (Vol. 30), came out admirably and on these occasions a special taxonomic

shine returned to the Gardens. At this time, because of a greater research emphasis

by P & R on horticultural matters, there was also an increase in papers reporting

experimental horticultural investigations (both in the laboratory and field).

Abrupt changes came to the management of the Gardens’ Bulletin for Vol.

31 Part 2 (December 1978). Asked to manage other aspects, Chang left the editorship

26 Gard. Bull. Singapore 64(1) 2012

nee

6 ee ee

Paste phi

eee

Fig. 7. Chang Kiaw Lan at her editing desk in December, 1974. (Photographer unknown,

courtesy of Christina Tan)

and was succeeded, for short periods, by Hardial Singh (Vol. 31 Part 2, Vol. 32), Y.S.

Choo (Vol. 33 Parts 1 and 2), and J.F. Maxwell (Vol. 34 Part 1). Then when Geh

chaired the editorial committee in place of the head of department who resigned, she

also had to take over as editor from Vol. 34 Part 2 (December 1981) and brought back

Chang’s expertise to assist with the Bulletin. Chang became managing editor from

1982 until 1987, when she retired, whereupon the editorship was managed by Geh

until Chin See Chung was appointed Keeper of the Herbarium and jointly edited from

1993 (Vol. 45) to 1996 (Vol. 48), when he became Director and Geh relinquished her

role to concentrate on another responsibility. T.W. Foong was also co-editor of the

Bulletin for 1995-1996. The Bulletin had literally gone through thick and thin, through

a difficult period when taxonomic productivity in its own home was at an ebb, and it

was Chang and Geh who plodded on with the work that kept the journal reasonably

well. As it turned out, this perseverance was significant because it brought the Bulletin

through a phase when in-house contribution was much lower than ever before. This

remarkable survival has developed a distinctly international flavour that has kept

up with scholarly expectations in scientific publishing: a significant volume of the

contributions coming from an international field of authors attracted to good standards

in editing, peer review of material, printing quality and timeliness of production.

Besides the broadened array of international contributions, several lines of

work brought out through the Bulletin are of special note for the Malay Peninsula

and for Singapore. Ian Turner, a taxonomist and ecologist and sometime Assistant

Director in charge of Horticulture at the Singapore Botanic Gardens, published A

Centenary of Gardens’ Bulletin, Singapore pay

Catalogue of the Vascular Plants of Malaya (Turner 1995), summarising efforts as

far as possible to present Malayan taxa with their most currently accepted names.

Two series of contributions mainly coordinated by National University of Singapore

botanists became visible, Zhe angiosperm flora of Singapore (Yan et al. 1992a, and

onwards) and Additions to the flora of Singapore (Yan et al. 1992b, and onwards).

A Field Guide to the Grasses of Singapore (Duistermaat 2005) was published as

Supplement to Vol. 57 of the Bulletin, a departure from separate numbering of earlier

supplements. Taxonomic work on orchids was re-emphasised somewhat with J.J.

Vermeulen (2000, and onwards) on the staff for some years, and Peter O’ Byrne having

taken up residence in Singapore. Also, Ruth Kiew’s research on Begonia, besides her

other work that is often reported in the Bu//etin, culminated in a Peninsular Malaysian

account (Kiew 2005).

There was redoubled interest in the biodiversity of Singapore itself. Greater

attention was focussed on the ecology of trees at the Bukit Timah Nature Reserve

(Wong 1987, Swan 1988) and the Central Catchment Nature Reserve (Wong et al.

1994), and a wider stocktaking of biological communities with Rain Forest in the City:

Bukit Timah Nature Reserve, Singapore (Chin et al. 1995). In 1993, a 2-ha permanent

ecological plot was set up in Bukit Timah that joined a long-term network of research

plots in tropical forests around the world coordinated by the Center for Tropical Forest

Science (CTFS) of the Smithsonian Tropical Research Institute, in partnership with the

Nanyang Technological University’s National Institute of Education and the National

Parks Board (LaFrankie et al. 2005). Chan & Corlett (1997) edited a special part on

Biodiversity in the Nature Reserves of Singapore. Turner et al. (1996) continued a

long-standing interest in local ecology in the Bulletin with a new appraisal of the

freshwater swamp forest in Singapore.

The effort to resuscitate the taxonomic sciences at the Singapore Botanic

Gardens only gained in pace when the orchidologist Tan Wee Kiat became Director

of the Singapore Botanic Gardens in 1989. Around this time, the opportunity for

reorganisation identified a National Parks Board as an ideal body for incorporating a

larger emphasis on environmental and biodiversity conservation in its management of

Singapore’s greenscapes and biological resources. With all the reorganisational flux,

the Gardens ’ Bulletin was not issued during 1990, the only time since the Second World

War period when issues missed some years! When Tan became Executive Director

of NParks (as the organisation came also to be known) in 1990 (concurrently still

Director of the Gardens), taxonomic activity was restored. Tay Eng Pin, a taxonomist

and morphologist who trained with Francis Hallé and had served in various capacities

with P & R and then NParks, was asked to take charge of the Keeper’s duties as

Senior Research Officer (Taxonomy) from 1990 until 1993, when he left and Chin,

an ethnobotanist and forest botanist who had studied with D.M. Smith, H.C. Conklin

and Benjamin C. Stone, was appointed Keeper and Senior Research Officer (Plant

Introduction). Tan became Chief Executive Officer of NParks in 1996, and Chin was

made Director of the Gardens. Ruth Kiew (another student of Corner’s) was appointed

Keeper of the Herbarium and concurrently was Editor of the Bulletin, 1997-2006;

succeeded by Benito Tan, 2006-2010, and the present author from 2010.

28 Gard. Bull. Singapore 64(1) 2012

Hindsight a hundred years on

The development of the Gardens’ Bulletin, Singapore can be traced from its roots as

a monthly bulletin. It provided reports of agricultural development and commodities

and occasional notes and articles of interesting crop and wild plants of its immediate

region, which was actively opening up for agricultural and other industries through

colonial development. Its transformation has been mediated by historical events,

from the enlargement of administrative and economic divergence between the Straits

Settlements and the Federated Malay States, to World War II, and the “malayanisation”

period during which the post-war British administration gave way to independence

and much reorganisation.

The Bulletin was reoriented as a scientific periodical during the time of

I.H. Burkill. Scientifically, the scope has diversified from an essentially “Malayan”

perspective painstakingly documented and refined through the primary efforts of a

smal! team of in-house botanical personnel at the Singapore Botanic Gardens, to a more

regionally relevant publication avenue in parallel with the larger floristic insights of

Flora Malesiana, and finally a journal with an international outlook. This development

appears inevitable, as at first, British Malaya was a natural unit largely flanked by Dutch-

held territories across the Malay Archipelago, and as the unfolding science revealed

much more about the region’s floristic limits and extreme biological richness, a greater

regional and then international perspective was necessary for meaningfully continuing

both science and development. The key ingredients that fuel this development have,

at all stages, involved persons with specialised botanical training who had an unusual

appetite for their science and steadfastness of scholarship and purpose. Above all, they

had a great respect for continuing a heritage accrued through the changing fortunes of

boom and ebb, and between prosperity and adversity.

‘We have a vision for our time, but we can be certain that it will not be the last.’

Richard Fortey, Earth (2005)

ACKNOWLEDGEMENTS. I thank Ms Christina Soh for expert assistance with archived

historical material in the Library of the Singapore Botanic Gardens, Y.W. Low for raising the

centenary alarm and assistance in compiling publication data for Gardens’ Bulletin issues, and

Dr. Nigel Taylor, Director of the Botanic Gardens, for encouraging this centenary record and

useful comments. A near-complete set of the Gardens’ Bulletin, Singapore inherited from Dr.

S.C. Chin provided useful insight into the workings of the botanic gardens, herbarium and

journal.

References

Alphonso, A.G. (1980) An obituary—Dr. Caetano Xavier Furtado (1897-1980). Gard. Bull.

Singapore 33: 301-302.

Centenary of Gardens’ Bulletin, Singapore 29

Anonymous (1974) Report of the Botanic Gardens for the year 1973. Gard. Bull. Singapore

27: 221-245.

Arditti, J. (1989) Kwan Koriba: botanist and soldier. Gard. Bull. Singapore 42: 1-17.

Brockway, L.H. (2000) Science and colonial expansion: The role of the British Royal Botanic

Gardens. Amer. Ethnol. 6: 449-465.

Burkill, H.M. (1968) James Sinclair, 1913-1968; An obituary and an appreciation. Gard. Bull.

Singapore 23: i—xxiil.

Burkill, H.M. (1983) Murray Ross Henderson, 1899-1983, and some notes on the administration

of botanical research in Malaysia. J. Malaysian Branch Roy. Asiat. Soc. 56: 87-104.

Burkill, H.M. (1993) The Singapore Botanic Gardens: A reputation to cherish and preserve. In:

Proceedings of the Singapore Botanic Gardens 130th Anniversary Seminar, pp. 10-15.

Singapore: National Parks Board.

Burkill, I.H. (1913) Annual Report on the Botanic Gardens, Singapore and Penang for the year

1912. Singapore: Government Printing Office, Straits Settlements.

Burkill, I.H. (1918a) The establishment of the Botanic Gardens, Singapore. Gard. Bull. Straits

Settlem. 2: 55-63.

Burkill, I.H. (1918b) The second phase in the history of the Botanic Gardens, Singapore. Gard.

Bull. Straits Settlem. 2: 93-108.

Burkill, I.H. (1927) Botanical Collectors, Collections and Collecting Places in the Malay

Peninsula. Gard. Bull. Straits Settlem. 4: 113-202.

Burkill, I.H. (1935) A Dictionary of the Economic Products of the Malay Peninsula. London:

Straits Settlements & Federated Malay States. (Reprinted 1966, Kuala Lumpur: Ministry

of Agriculture & Co-operatives, Malaysia; 3rd printing 1993, Kuala Lumpur: Ministry of

Agriculture, Malaysia.)

Burkill, I.H. (1951) Dioscoreaceae. In: Steenis, C.G.G.J. van (ed) Flora Malesiana Series I,

4(3): 293-335. Batavia: Noordhoff-Kolff N.V.

Chan, L. & Corlett, R.T. (eds) (1997) Biodiversity in the Nature Reserves of Singapore. Gard.

Bull. Singapore 49: 147-159.

Chew, Wee-Lek (1963) Florae Malesianae Precursores—XXXIV. A revision of the genus

Poikilospermum (Urticaceae). Gard. Bull. Singapore 20: 1-103.

Chew, Wee-Lek (1965) Laportea and allied genera (Urticaceae). Gard. Bull. Singapore 21:

195-208.

Chew, Wee-Lek (1969a) Discocnide (Urticaceae). Gard. Bull. Singapore 24: 355-359.

Chew, Wee-Lek (1969b) Nothocnide (Urticaceae) in Malesia. Gard. Bull. Singapore 24: 361—

B73:

Chin, S.C. (1977) The limestone hill flora of Malaya I. Gard. Bull. Singapore 30: 165-219.

Chin, S.C. (1979) The limestone hill flora of Malaya I. Gard. Bull. Singapore 32: 64-203.

Chin, S.C. (1983) The limestone hill flora of Malaya III. Gard. Bull. Singapore 35: 137-190.

Chin, S.C. (1983) The limestone hill flora of Malaya IV. Gard. Bull. Singapore 36: 31-91.

Chin, S.C., Corlett, R.T., Wee, Y.C. & Geh, S.Y. (eds) (1995). Rain Forest in the City: Bukit

Timah Nature Reserve, Singapore. Gard. Bull. Singapore, Supplement no. 3.

Corner, E.J.H. (1940) Wayside Trees of Malaya, 2 vols. Singapore: Government Printing Office.

(2nd ed. 1952, Singapore: Government Printing Office; 3rd ed. 1988, Kuala Lumpur:

Malayan Nature Society.)

Corner, E.J.H. (1946) Japanese men of science in Malaya during the Japanese occupation.

Nature 158: 63.

Corner, E.J.H. (1962) The Classification of Moraceae. Gard. Bull. Singapore 19: 187-252.

Corner, E.J.H. (1965) Mount Kinabalu East. Sabah Soc. J. 4: 170-187.

30 Gard. Bull. Singapore 64(1) 2012

Corner, E.J.H. (1978) The Freshwater Swamp-Forest of South Johore and Singapore. Gard.

Bull. Singapore, Supplement no. |.

Corner, E.J.H. (1981) The Marquis, a tale of Syonan-to. Singapore: Heinemann Asia.

Curtis, C. (1894) Catalogue of the Flowering Plants and Ferns found growing wild in the Island

of Penang. J. Straits Branch Roy. Asiat. Soc. 25: 67-163.

Duistermaat, H. (2005) Field Guide to the Grasses of Singapore (Excluding the Bamboos).

Gard. Bull. Singapore 57 (Supplement): 1-177.

Fortey, R. (2005) Earth. An Intimate History. New York: Vintage Books.

Furtado, C.X. & Holttum, R.E. (1960) I.H. Burkill in Malaya. Gard. Bull. Singapore 17: 350—

356.

Hill, H.C. (1900a) The present system of forest administration in the Federated Malay States

with suggestions for future management. Kuala Lumpur: Government Printer.

Hill, H.C. (1900b) The present system of forest conservancy in the Straits Settlements with

suggestions for future management. Singapore: Government Printer.

Holttum, R.E. (1950) The Zingiberaceae of the Malay Peninsula. Gard. Bull. Singapore 13:

1-250.

Holttum, R.E. (1953) A Revised Flora of Malaya, vol. I. Orchids of Malaya. Singapore:

Government Printing Office. (Reprinted 1960, 2nd ed. 1966.)

Holttum, R.E. (1954) 4A Revised Flora of Malaya, vol. II. Ferns of Malaya. Singapore:

Government Printing Office. (2nd ed. 1957, 3rd ed. 1964.)

Holttum, R.E. (1958a) The bamboos of the Malay Peninsula. Gard. Bull. Singapore 16: 1-135.

Holttum, R.E. (1958b) Kwan Koriba. Gard. Bull. Singapore 17: 339-340.

Holttum, R.E. (1959) Orchids, gingers and bamboos: Pioneer work at the Singapore Botanic

Gardens and its significance for botany and horticulture. Gard. Bull. Singapore 17: 190-

194.

Holttum, R.E. (1967) Isaac Henry Burkill 1870-1965. Gard. Bull. Singapore 22: 67-70.

Johnson, D.V. & Tay, E.P. (1999) C.X. Furtado (1897-1980): Contributions to the study of

palms. Gard. Bull. Singapore 51: 141-150.

Karch, S.B. (1998) A Brief History of Cocaine. Boca Raton, Florida: CRC Press.

Keng, H. (1969) Flora Malesianae Precursores XLVIII. A revision of Malesian Labiatae. Gard.

Bull. Singapore 24: 13-180.

Kiew, R. (1999) The Singapore Botanic Gardens Herbarium—125 years of history. Gard. Bull.

Singapore 51: 151-161.

Kiew, R. (2005) Begonias of Peninsular Malaysia. Kota Kinabalu: Natural History Publications

(Borneo).

LaFrankie, J.V., Davies, S.J., Wang, L.K., Lee, S.K. & Lum, S.K.Y. (2005) Forest Trees of Bukit

Timah. Population Ecology in a Tropical Forest Fragment. Singapore: Simply Green.

Lam, H.J. (1959) A tale of two cities: Singapore and Leiden. Gard. Bull. Singapore 17: 166—

170.

Mabberley, D.J. & Chang, Kiaw Lan (eds) (1977) Tropical Botany. Essays presented to E.J.H.

Corner for his seventieth birthday, 1976. Gard. Bull. Singapore 29: 1-266.

Mandalam, K.R. (2005) Edred John Henry Corner: Leader, First and Second Royal Society

Expeditions to North Borneo. In: Mandalam, K.R., Davison, G.W.H. & Regis, P. (eds)

East Kinabalu, Royal Society Expeditions 40th Anniversary Monograph, pp. 1-3. Kota

Kinabalu: The Sabah Society.

Mandalam, K.R. (2011) The Kotal Route Sketches, Royal Society Kinabalu Expedition 1964.

Original Field Sketches of Prof. John Corner. Kota Kinabalu: The Sabah Society.

Menon, K.D. (1969) A brief history of forest research in Malaya. Malayan Forester 32: 3-29.

Centenary of Gardens’ Bulletin, Singapore 31

Ministry of National Development (1977) Annual Report 1976. Singapore: Ministry of National

Development.

Muller, L. (1998) Flowering and fruiting of a Gigantochloa ridleyi clone, seed germination and

growth. Amer. Bamboo Soc. Newslett. 19(3): 8-11.

Muller, L. (1999) The many reproductive modes of clumping bamboos. Amer. Bamboo Soc.

Newslett. 20(5): 1-4.

Munir Ahmad Abid (1965) A revision of Petraeovitex (Verbenaceae). Gard. Bull. Singapore

Ble 215-257.

Munir Ahmad Abid (1967) A revision of Symphorema. Verbenaceae. Gard. Bull. Singapore 22:

153-171.

Ng, F.S.P. & Jacobs, M. (1983) A guide to King’s “Materials for a Flora of The Malayan

Peninsula”. Gard. Bull. Singapore 36: 177-185.

Purseglove, J.W. (1955a) Mr. H.N. Ridley’s hundredth birthday. Nature 176: 1092-1093.

Purseglove, J.W. (1955b) Ridley, Malaya’s greatest naturalist. Malayan Nat. J. 10(2): 43-55.

Purseglove, J.W. (1955c) The Ridley Centenary. Singapore: Government Printer.

Purseglove, J.W. (1959) History and functions of Botanic Gardens with special reference to

Singapore. Gard. Bull. Singapore 17: 125-154.

Ridley, H.N. (1900) Flora of Singapore. J. Straits Branch Roy. Asiat. Soc. 33: 27-196.

Ridley, H.N. (1907) Materials for a Flora of the Malayan Peninsula. Part 1, pp. 1-233; Part

IT, pp. 1-235; Part III, pp. 1-197. Singapore: Methodist Publishing House.

Ridley, H.N. (1908) Annual Report on the Botanic Gardens, Singapore, for the Year 1907.

Singapore: Government Printing Office.

Ridley, H.N. (1922-1925) The Flora of the Malay Peninsula, vols. 1-5. London: Reeve & Co.

Sinclair, J. (1958) A revision of Malayan Myristicaceae. Gard. Bull. Singapore 16: 205-466.

Sinclair, J. (1958) Florae Malesianae Precursores—XX. The genus Gymnacranthera

(Myristicaceae) in Malaysia. Gard. Bull. Singapore 17: 96-120.

Sinclair, J. (1961) Florae Malesianae Precursores—XXXI, The genus Knema (Myristicaceae)

in Malaysia and outside Malaysia. Gard. Bull. Singapore 18: 102-327.

Sinclair, J. (1968) Florae Malesianae Precursores—XLII, The genus Myristica (Myristicaceae)

in Malesia and outside Malesia. Gard. Bull. Singapore 23: 1-540.

Sinclair, J. (1974) The genus Horsfieldia (Myristicaceae) in and outside Malesia I: H. sabulosa

and H. whitmorei J.Sinclair spp. nov. Gard. Bull. Singapore 27: 133-141.

Sinclair, J. (1975) The genus Horsfieldia (Myristicaceae) in and outside Malesia II. Gard. Bull.

Singapore 28: 1-181.

Steenis, C.G.G.J. van (1948) Flora Malesiana Series I, 4(1). Batavia: Noordhoff-Kolff N.V.

Steenis, C.G.G.J. van (1959) Singapore and Flora Malesiana. Gard. Bull. Singapore 17: 161—

165.

Swan, F.R. (1988) Tree distribution patterns in the Bukit Timah Nature Reserve, Singapore.

Gard. Bull. Singapore 47: 59-81.

Tan, H.T.W., Chua, K.S. & Turner, I.M. (1992a) The Angiosperm Flora of Singapore Project.

Gard. Bull. Singapore 44: 1-2.

Tan, H.T.W., Ali Ibrahim & Chua, K.S. (1992b) Additions to the Flora of Singapore, I. Gard.

Bull. Singapore 44: 127-133.

Turner, I.M. (1995) A Catalogue of the Vascular Plants of Malaya. Gard. Bull. Singapore 47:

1-757.

Turner, I.M., Boo, C.M., Wong, Y.K., Chew, P.T. & Ali Ibrahim (1996) Freshwater swamp

forest in Singapore, with particular reference to that found around the Nee Soon Firing

Ranges. Gard. Bull. Singapore 48: 129-157.

by) Gard. Bull. Singapore 64(1) 2012

Vermeulen, J.J. (2000) Carr’s spirit collection of Kinabalu orchids recovered, with some notes

on Bulbophyllum species present in this collection. Gard. Bull. Singapore 52: 273-284.

Watson, J.G. (1950) Some materials for a forest history of Malaya. Malayan Forester 13: 63—

72:

Wong, K.M. (1987) The herbarium and arboretum of the Forest Research Institute of Malaysia

at Kepong—a historical perspective. Gard. Bull. Singapore 40: 15-30.

Wong, K.M. (2003) Dr Chang Kiaw Lan, 31 July 1927—14 August 2003. Gard. Bull. Singapore

55: 309-315.

Wong, K.M. (2004) Bamboo, The Amazing Grass. A Guide to the Diversity and Study of

Bamboos in Southeast Asia. Kuala Lumpur: International Plant Genetic Resources

Institute (IPGRI) & University of Malaya.

Wong, Y.K. (1987) Ecology of the trees of Bukit Timah Nature Reserve. Gard. Bull. Singapore

40: 45-76.

Wong, Y.K., Chew, P.T. & Ali Ibrahim (1994) The tree communities of the Central Catchment

Nature Reserve, Singapore. Gard. Bull. Singapore 46: 37-63.

Wyatt-Smith, J. (1950) Virgin Jungle Reserves. Malayan Forester 13: 92-94.

Wycherley, P.R. (1959) The Singapore Botanic Gardens and Rubber in Malaya. Gard. Bull.

Singapore 17: 175-186.

ive)

Gardens’ Bulletin Singapore 64(1): 33-49. 2012

A revision of Nepenthes (Nepenthaceae) from Gunung

Tahan, Peninsular Malaysia

Charles Clarke! and Ch’ien C. Lee?

' School of Science, Monash University Sunway Campus,

Jalan Lagoon Selatan, 46150 Bandar Sunway,

Selangor Darul Ehsan, Malaysia

charles.clarke@sci-monash.edu.my

* Peti Surat 2507, 93750 Kuching, Sarawak, Malaysia

ABSTRACT. The Nepenthes from Gunung Tahan in Peninsular Malaysia are revised.

We recognise four species from this mountain: V. alba, N. benstonei, N. gracillima and N.

sanguinea. The reinstatement of N. alba is based on a consistent difference in upper pitcher

colouration between it (typically evenly pale yellowish to ivory white) and N. gracillima

(dark green with purple-brown speckles). Material from Gunung Tahan that was identified in

previous treatments as NV. macfarlanei belongs to N. gracillima and the former species is absent

from Gunung Tahan. Nepenthes alba and N. gracillima are very similar to N. macfarlanei and

further examinations of the relationships among these taxa are warranted.

Keywords. Nepenthes alba, N. benstonei, N. gracillima, N. macfarlanei, Gunung Tahan,

Malaysia

Introduction

Eleven species of Nepenthes (Nepenthaceae) have been recorded from Peninsular

Malaysia, including five montane species, which are generally found in habitats above

1000 m altitude (Cheek & Jebb 2001: Clarke 2001: McPherson 2009). All of the

montane species have been recorded from Gunung Tahan which, at 2187 m above

sea level (asl), is the highest mountain on the Malay Peninsula. Situated in the Timur

Range, Gunung Tahan is isolated from the main Titiwangsa Range of Peninsular

Malaysia by at least 100 km. To date, eight Nepenthes taxa have been recorded from

the mountain, including N. alata Blanco, N. bongso Korth., N. singalana Becc., N.

macfarlanei Hemsl., N. alba Ridl., N. gracillima Ridl., N. sanguinea Lindl. and

N. benstonei C. Clarke. The Nepenthes of Gunung Tahan have been reviewed on a

number of occasions (Ridley 1924; Danser 1928; Kiew 1990; Jebb & Cheek 1997;

Clarke 2001: McPherson 2009), but uncertainty about the correct identity and status

of several taxa persists. Most confusion relates to V. gracillima and its relationships

to N. macfarlanei and N. alba. In this paper, we demonstrate that the source of much

of this confusion arises from a failure by previous researchers to properly delineate N.

gracillima. As a consequence, many collections of this species have been misidentified

as N. macfarlanei. We argue that N. macfarlanei is absent from Gunung Tahan, where

34 Gard. Bull. Singapore 64(1) 2012

it is replaced by N. gracillima, but that these taxa are very closely related, exhibiting

few significant morphological differences. We conclude that only four Nepenthes

species are extant on Gunung Tahan. These are N. alba (also a very close relative

of N. gracillima and N. macfarlanei), N. benstonei, N. gracillima and N. sanguinea.

Our interpretations are based on both detailed field observations and examinations

of herbarium material. As all previous descriptions of N. gracillima have included

material that we consider to belong to other taxa, and a detailed description of N. alba

has never been published, we present revised descriptions of these taxa and a key to

the Nepenthes of Gunung Tahan.

Discovery and description of Nepenthes from Gunung Tahan

The first botanist to describe Nepenthes from Gunung Tahan was H.N. Ridley, who

was Director of the Singapore Botanic Gardens from 1888-1911. He based his

initial identifications and descriptions (Ridley 1908) upon specimens collected by

H.C. Robinson and L. Wray on an expedition to Gunung Tahan made in 1905. The

first species he recorded was identified as N. bongso, a species that was previously

known only from Sumatra. This identification was made on Wray & Robinson 5411

(SING), which comprises a fragment of a climbing stem of a diminutive plant that

bears four upper pitchers and a male inflorescence (Note: Nepenthes are generally

dimorphic, producing two types of pitchers that are usually referred to as “lower” and

“upper”. For a review of plant architecture, see Clarke (2001)). Ridley (1908) also

described a new species, N. gracillima, based upon Wray & Robinson 5309 (SING).

This specimen consists of a fragment of a climbing stem bearing three upper pitchers

and an infructescence, which is badly damaged. The lower pitchers of N. gracillima

were not described by Ridley (1908), nor were any specimens bearing lower pitchers

equated with N. gracillima by him. The descriptions for both taxa are very brief and

lack sufficient detail to make objective comparisons. The pitchers, leaves and stems of

the two specimens are very similar in structure — the primary difference between them

appears to be the colour of the pitchers. Those of Wray & Robinson 5411 were said

to be white, tinted pale green at base and spotted with pink, whereas those of Wray &

Robinson 5309 were stated to be “pale green, tinted in places with dull crimson and

mottled with dull purple”. Ridley (1908) also noted that the “neck and lamina” of the

pitcher lid of N. gracillima was pubescent.

In the same year, Macfarlane (1908) identified Wray & Robinson 5411

as N. singalana (another Sumatran species), but did not explain his interpretation.

Macfarlane (1908) also provided a more detailed description of N. gracillima and

stated that the pitchers were monomorphic (i.e., only one type is produced; thereby

implying that lower pitchers had not been collected because they are not produced).

Of the indumentum of the pitcher lids, he wrote, extus et intus sparse pubescens... (=

“sparsely pubescent inside and out”). At the time of Macfarlane’s revision, the only

other species from Peninsular Malaysia that was known to have hairs on the lower

surface of the pitcher lid (referred to hereafter as “lid hairs”) was N. macfarlanei,

Nepenthes on Peninsular Malaysia ’s Gunung Tahan 35

which had been recorded from a number of mountains in the Titiwangsa range, in

addition to the type locality on Gunung Bubu in Perak. Neither Ridley (1908) nor

Macfarlane (1908) discussed this distinctive, shared characteristic further, nor was any

mention made regarding the presence of lid hairs on Wray & Robinson 5411.

In 1909, Ridley described N. ramispina Ridl., a species that bears strong

similarities to N. gracillima, but which occurs in the main Titiwangsa Range, to

the west of Gunung Tahan (Ridley 1909). This species lacks lid hairs, but in most

other respects, its pitchers are very similar to those of NV. gracillima. Ridley (1909)

distinguished this species from N. gracillima on the basis of the large, branched spurs

at the apices of the pitchers (the spurs of NV. gracillima are simple).

In 1911, Ridley climbed Gunung Tahan and made several further collections,

one of which (Ridley 16097 (SING (1 sheet), K (2 sheets))) he identified as NV. singalana.

The sheet at SING consists of a fragment of a climbing stem bearing two upper pitchers

and two female inflorescences. The leaf structure differs from that of N. gracillima in

two important ways: (a) the leaf blades are subpetiolate instead of sessile, and (b) the

margins of the leaf blades are decurrent along the internodes for up to 1.5 cm. The

sheet at K that is barcoded K000651565 contains a fragment of a climbing stem with

an immature female inflorescence, but no pitchers. The leaf structure of this specimen

is the same as that of the sheet at SING and these clearly belong to the same taxon.

However, the sheet at K barcoded K00065 1564 contains a fragment of a climbing stem

that bears sessile, non-decurrent leaves and small, squat aerial pitchers that resemble

those of Wray & Robinson 5411. Clearly, this is a mixed collection involving two

different taxa, neither of which were equated with N. gracillima by Ridley.

Ridley (1924) eventually corrected his mis-identification of Wray & Robinson

5411, noting that it was not the same as N. bongso (or N. singalana) and instead

described it as a new species, N. alba. Once more, the description was very brief, but

Ridley clearly mentioned the distinctive colour of living plants of N. alba, as follows:

“pitcher...ivory white, sometimes spotted with rose pink in the mouth and lid, rarely

canary yellow.

Danser (1928) revised the genus for the Netherland’s Indies (now Indonesia),

along with a few outliers from adjacent regions, including the Malay Peninsula. With

regard to the Nepenthes of Gunung Tahan, Danser re-determined Ridley 16097 (SING)

as N. alata. Prior to Danser’s revision, N. alata was known only from the Philippines,

but Danser also reduced Nepenthes eustachya Mig. from Sumatra to a synonym of

N. alata, thereby extending its geographical range substantially. This interpretation

also broadened the degree of morphological variation encompassed by N. alata and

as Peninsular Malaysia lies between Sumatra and the Philippines, it is perhaps not

surprising that Danser felt that Ridley 16097 also represented this species. However, no

other collections that can be equated with NV. a/ata have ever been made in Peninsular

Malaysia, casting doubt over the validity of his interpretation of this specimen (Kiew

1990; Clarke 2001; see below).

Danser (1928) also reduced N. ramispina and N. alba to synonyms of N.

gracillima, stating that the differences among them were “of very little importance”.

He noted that herbarium material of N. gracillima was not always easily distinguished

36 Gard. Bull. Singapore 64(1) 2012

from N. macfarlanei and N. sanguinea. Furthermore, he tentatively identified two

specimens (Wray 339 (BO) and Ridley 16174 (SING)) as belonging to a natural hybrid,

N. gracillima x N. macfarlanei. Ridley 16174 was collected from Gunung Tahan, near

Wray’s Camp (the type locality for NV. gracillima) and of these specimens, Danser

(1928) wrote,

The pitchers are too wide for N. gracillima and the underside of the lid in the number

Wray 339 bears the bristles typical of N. macfarlanei. | should prefer to mention this

specimen under the latter species, the pitchers of which vary extraordinarily, when the

inner margin of the peristome were not entire. The other number, Ridley 16174, bears

much more dense and delicate hairs on the underside of the lid, a character that is also

often found in N. sanguinea and N. macfarlanei.

Below, we demonstrate that Ridley 16174 belongs to N. gracillima, and that

Danser was the first to confuse material of N. gracillima with N. macfarlanei, due to

the presence of lid hairs. It has since been demonstrated that N. sanguinea lacks lid

hairs, but that a few hairs may be present on lids of NV. macfarlanei x N. sanguinea

pitchers (Jebb & Cheek 1997; Clarke 2001). Danser’s (1928) revised description of N.

gracillima included material of N. ramispina and N. alba, and made no mention of lid

hairs in this taxon, indicating that he was of the opinion that lid hairs were found only

in N. macfarlanei.

Kiew (1990) was the first since Ridley to combine examinations of herbarium

material from Gunung Tahan with field observations (Macfarlane and Danser never

visited the mountain). She re-determined Ridley 16097 as N. gracillima and explained

that Danser’s (1928) determination was a simple misidentification (Clarke 2001). It

is however intriguing that neither Ridley nor Danser equated this specimen with N.

gracillima; suggesting that the reasons for their misidentifications were not necessarily

straightforward. Kiew (1990) noted that the first Nepenthes to be encountered on

the southern route to Gunung Tahan (from Kuala Tahan) is N. macfarlanei, near

Wray’s Camp at approximately 900 m asl (where Ridley 16174 was collected). She

distinguished it from the other species on the basis of the underside of the pitcher

lid, which is covered in coarse hairs, and by the inner edge of the peristome, which

is toothed. She then discussed N. gracillima, referring to it as the most conspicuous

pitcher plant of the padang, festooning

..every bush with its small, ivory-white pitchers, which from afar look like candles.

Plants begin life producing deep purple leaves and pitchers that are almost black in

colour. Once a climbing stem develops the leaves produced are green and the pitchers

ivory-white with rosy spots in the upper part. It is still possible to find on the same plant

purplish pitchers, which are usually hidden within the crown of the supporting shrub.

Kiew (1990) indicated that Ridley (1915) was of the opinion that the white-

pitchered plants belonged to N. a/ba (although the latter name was not published until

1924), whereas the purple/black-pitchered plants belonged to N. gracillima. However,

she argued that since both pitcher types can be found on the same plant, this was

not the case and all plants belonged to N. gracillima. This concept of N. gracillima

is contingent on how the type specimen (Wray & Robinson 5309) is interpreted: if

the gracile, dark-coloured pitchers of this specimen are considered to be the same as

the lower pitchers of NV. a/ba, then Danser’s (1928) decision to reduce N. alba to a

Nepenthes on Peninsular Malaysia's Gunung Tahan 37

synonym of N. gracillima is justified. However, Wray & Robinson 5309 bears upper

pitchers on a climbing stem—their shape and colouration are not consistent with the

corresponding pitcher type in N. alba.

Kiew’s (1990) observations of N. gracillima on Gunung Tahan also showed that

clear morphological differences exist between this taxon and Ridley’s N. ramispina,

which seems to be confined to the Titiwangsa Range. As a consequence, Jebb & Cheek

(1997) reinstated N. ramispina, thereby ending some of the confusion surrounding

N. gracillima and limiting any persistent uncertainty to populations from the Timur

Range. Jebb & Cheek (1997) retained Danser’s (1928) concept of N. gracillima, with

N. alba as its synonym and, like Kiew (1990), treated Ridley 16097 as belonging to N.

gracillima (they also reinstated N. eustachya, thereby removing much of the confusion

about the status, morphological variation and geographical distribution of N. alata,

which is now known to be endemic to the Philippines (Jebb & Cheek 1997)).

Clarke (2001) adopted Jebb & Cheek’s (1997) interpretations of N. gracillima

and N. ramispina, but rejected Kiew’s (1990) interpretation of Ridley 16097, considering

this specimen to be N. benstonei, a recently-described species from Bukit Bakar in

Kelantan, approximately 120 km to the north of Gunung Tahan (Clarke 1999, 2001).

Clarke (2001) provided a detailed explanation of the differences between N. benstonei

and N. gracillima, noting that the leaf blade structure (in particular, the sub-petiolate

leaf bases and decurrent leaf margins), indumentum and pitcher characteristics of

Ridley 16097 correspond with those of N. benstonei. Clarke’s (2001) treatment of N.

gracillima, N. alba and N. ramispina was limited to herbarium material and followed

that of Jebb & Cheek (1997).

Cheek & Jebb (2001) provided a revised description and illustration of N.

gracillima, based almost entirely on specimens at SING and K that are representative

of Ridley’s concept of this species (e.g., Ridley 16174 (SING, K), Haniff 7890 (SING),

Pannell 1132 (K) and Ridley 16090 (K), rather than N. a/ba. For the first time, specimens

bearing lower pitchers (Pannell 1/32) were included. In terms of size and colour, the

lower pitchers of Pannell 1/32 are similar to those of the upper pitchers on the type

(Wray & Robinson 5309). Both the description and illustration of NV. gracillima (Fig.

8) indicate that the pitcher lids have hairs on the lower surface, yet this feature was not

discussed further.

McPherson (2009) climbed Gunung Tahan via the western route from Sungai

Relau in 2008. He considered N. alba to be distinct from N. gracillima, stating that the

taxa he observed were consistent with Ridley’s original descriptions of them. Several

differences in morphology were used to distinguish these taxa; these are summarised

in Table 1.

McPherson (2009) placed considerable emphasis on variations in pitcher size

and colour (especially the colour of the upper pitchers of NV. a/ba). Although these

characteristics may differ substantially between these taxa, it is worth noting that the

types of both NV. gracillima (Wray & Robinson 5309) and N. alba (Wray & Robinson

5411) bear upper pitchers of roughly equivalent structure and dimensions and are

not atypical for their respective taxa, indicating that the types cannot be reliably or

consistently distinguished on the basis of size or gross morphology: the only apparent

Table 1.

macfarlanei.

Characteristic

Geographical range

Altitudinal range

Habitat

Lid hairs

Tendrils of rosette leaves

Tendrils of leaves on

climbing stems

Size of lower pitchers

Colour of lower pitchers

[on rosettes of mature

plants]

Lid of lower pitchers

Production of

intermediate pitchers

Intermediate pitchers

Upper pitchers

N. alba

Timur Range

1400-2150 m

Terrestrial in highly

stunted, open padang

vegetation

Very fine, up to 1 mm

long

0.8—1.5 times the length

of pitcher height and leaf

length

Equal to or shorter than

pitcher height and leaf

blade length

Up to 12 em tall and 4.5

cm wide, but usually

much smaller

Predominantly purplish

brown, with a lighter

interior and dark purple

or black peristome.

Colouration very

consistent.

Orbiculate or elliptic

Rare

Broadly infundibular

in the lower half, hip

distinct, cylindrical above

Infundibular in the

lower 1/2—2/3, sharply

contracted at the hip,

cylindrical to narrowly

infundibular above

Gard. Bull. Singapore 64(1) 2012

N. gracillima

Timur Range, Gunung

Tapis

900-1700 m

Terrestrial or epiphytic

in closed upper montane

forest

Very fine, up to 2 mm

long

2-5 times longer than

pitcher height

Longer than both leaf

blade length and pitcher

height

Up to 22 cm tall and 7

cm wide, often reaching

these dimensions

Predominantly yellow-

green, with dark red or

purple blotches. Interior

light yellowish-green.

Peristome green or

reddish, often striped

with bands of dark red or

purple.

Elliptic or ovate

Dominant form of pitcher

on climbing stems

Narrowly infundibular

in the lower half,

hip present but not

pronounced, narrowly

infundibular above

Very narrow throughout,

infundibular in the lower

1/3, gradually contracted

above the hip, cylindrical

above

Morphological characteristics that distinguish N. alba, N. gracillima and N.

N. macfarlanei

Titiwangsa Range,

Gunung Bubu

900-2100 m

Terrestrial or epiphytic in

mossy forest

Fine, up to 3 mm long,

except on Gunung Bubu,

where they are coarse, up

to 5 mm long

2-5 times longer than

pitcher height

Longer than both leaf

blade length and pitcher

height

Up to 22 cm tall and 7

cm wide, often reaching

these dimensions

Predominantly yellow-

green, with dark red or

purple blotches. Interior

light yellowish-green.

Peristome red.

Sub-orbiculate

Occasional on climbing

stems in dense forest

Narrowly infundibular

in the lower half, hip

present, infundibular

above

Infundibular from base to

hip, hip located anywhere

from immediately

beneath peristome to

1/3 of the way up from

the base; cylindrical to

slightly infundibular

above the hip

Nepenthes on Peninsular Malaysia's Gunung Tahan

Characteristic

Colour of upper pitchers

Lid of upper pitchers*

Size of intermediate

pitchers on mature plants

Size of upper pitchers on

mature plants

Colour of lower pitchers

on seedling rosettes

Colour of lower pitchers

Colour of the

intermediate pitchers

Colour of the upper

pitchers

N. alba

Conspicuous white

colouration often with red

blotches and flecks

Orbiculate or elliptic

Up to 9 cm tall by 2 cm

wide

Up to 12 cm tall by 3 cm

wide

Reddish-purple

throughout, overlain

with dark purple flecks;

peristome dark pink-

purple

As in the seedling

rosettes

Whitish-green, heavily

overlain with thin,

dark red-brown flecks

throughout

Ivory white throughout,

sometimes overlain

with rose to red specks,

to varying degrees of

intensity

N. gracillima

Uniformly black, strongly

speckled bright green.

Elliptic or ovate

Up to 22 cm tall by 4 cm

wide

Up to 10 cm tall by 1.5

cm wide

Dull green, heavily

overlain with dull red-

brown flecks; peristome

dull brown throughout

Dark to light green,

moderately to heavily

overlain with dark red-

brown flecks throughout;

peristome green with

red-brown stripes or dark

red-brown throughout

As in the rosette pitchers

N. macfarlanei

White with red flecks and

bands ont he peristome,

to green, with dark purple

speckles and peristome

bands.

Sub-orbiculate

Up to 18 cm tall by 5 cm

wide

Up to 20 cm tall by 6 cm

wide

Variable: greenish white.

heavily overlain with

brown, pink or red flecks;

peristome dark red or

pink throughout

As in the seedling

rosettes

As in the rosette pitchers

As in the rosette

pitchers, or ivory white

throughout, usually with

some rose pink flecks

* This characteristic is informative, but we found the lids of N. a/ba pitchers to be orbiculate

and cordate at the base, whereas those of N. gracillima are ovate and not cordate.

differences between them relate to colour (which the collectors’ label annotations on

the type specimens and the species descriptions (Ridley 1908, 1924) clearly record).

Furthermore, there are specimens of N. alba from 2100 m asl on the padang (e.g.,

Strugnell & Sow 42878 (KEP)) that bear heavily speckled pitchers that strongly

resemble N. gracillima. Variation in pitcher size and colour in most Nepenthes species

is considerable and Danser (1928) commented on its lack of taxonomic value (although

40 Gard. Bull. Singapore 64(1) 2012

Clarke (2001) demonstrated that in some taxa, pitcher colour can be both a stable and

informative characteristic).

Several characteristics of N. gracillima that were used by McPherson (2009)

to distinguish it from N. a/ba are not consistent with Ridley’s (1908) or Jebb & Cheek’s

(2001) descriptions of this species: these describe a small, slender plant that is very

similar in most respects to NV. alba, not the robust, large one discussed by McPherson

(2009). In fact, the plant that McPherson treats as N. gracillima bears a very strong

resemblance to N. macfarlanei. McPherson (2009) provided two photographs of N.

gracillima. His Fig. 316 is a photograph of a lower pitcher that is broadly ovoid, with

a peristome that is flattened and expanded towards the rear, and which bears distinct

teeth. The ovate lid is densely covered with simple white hairs — these cause the central

part of the lid to appear out of focus in this photograph, even though both the front and

rear margins are in focus. His Fig. 317 is a photograph of an upper pitcher. Again, the

ovate lid is covered with hairs on the underside, while the peristome is narrower and

mostly cylindrical throughout, but small teeth are still apparent towards the apex of

the orifice at the rear. The pitcher has a distinct hip approximately 1/3 of the way up

from the base and appears in most respects to represent an intermediate pitcher of N.

macfarlanei.

Thus, there is a distinct disparity between the published descriptions of N.

gracillima and McPherson’s (2009) interpretation of it. There are two possible

explanations for this: (a) he confused plants of N. macfarlanei with N. gracillima;

or (b) he correctly identified N. gracillima, but this species is more similar to N.

macfarlanei than anyone has previously realised, possessing some morphological

traits that have generally been considered diagnostic of that species. McPherson

(2009) noted that “significant populations [of N. macfarlanei| occur in.... Taman

Negara’, indicating that he identified some plants that he saw on the mountain as this

species. As a consequence, McPherson’s (2009) arguments for reinstating NV. a/ba have

merit, but NV. gracillima is yet to be clearly defined and delimited with regard to its

putative close relatives. Against this background, we visited Gunung Tahan to study its

Nepenthes flora and attempt to resolve some of the uncertainties about N. benstonei, N.

gracillima, N. macfarlanei and N. alba.

Materials and methods

We conducted detailed examinations of herbarium material at the following herbaria:

Singapore (SING), Forest Research Institute Malaysia (KEP) and Bogor (BO).

Although relevant material at K and L has been seen by one of us (CC), the detailed

examinations required to make a contribution in this paper have not been made,

although digital images of some specimens that are available online were consulted.

We conducted field observations on Gunung Tahan from March 29 to April 1, 2011,

ascending the mountain by the western route, from Sungai Relau. Waypoints and

locations were recorded using a Garmin Dakota 20 GPS receiver (Garmin Corporation,

Kansas), using the WGS84 geodetic system.

Nepenthes on Peninsular Malaysia’s Gunung Tahan 4]

Results

The first Nepenthes we encountered on the western route to Gunung Tahan was N.

benstonei, which is common on steep ridge tops between approximately 800 and 1200

m altitude. This species is clearly abundant and widespread on Gunung Tahan, even

though no other researchers have recorded it there.

From about 1350—1700 m asl, in dense vegetation up to 6-10 m tall along ridge

tops, we encountered a second species, growing as both an epiphyte or terrestrially in

mossy embankments. This taxon resembles N. macfarlanei in virtually all respects,

with the lower surfaces of the pitcher lids being covered with short, simple white hairs.

The lower pitchers of these plants frequently resemble those of Ridley 16174, which

Danser (1928) thought could have been a natural hybrid between N. macfarlanei

and N. gracillima. At about 1500 m asl (N 4.6354°, E 102.2051°), we encountered a

Nepenthes that climbs into the forest canopy (up to 5 m) and produces very slender,

small upper pitchers that match those of the type of V. gracillima. We traced the stems

of this taxon to the ground and found they belong to the same plants that we had

previously thought were N. macfarlanei. We repeated this exercise on many occasions

throughout the range of the plants that resemble N. gracillima and N. macfarlanei

that we saw on Gunung Tahan and determined that they all belonged to the same

taxon. These plants also correspond well with the photographs and description of N.

gracillima provided by McPherson (2009). Several plants at the location produce

pitchers that closely match the large, robust “upper” pitcher of N. gracillima illustrated

in Fig. 317 of McPherson (2009). These are exclusively borne on climbing stems that

are 1-3 m in length. Longer stems produce the smaller, more gracile type of upper

pitchers seen on the type. We observed this growth pattern (and production of two

types of pitchers on climbing stems) to be common to virtually all mature plants of N.

gracillima that we observed on Gunung Tahan.

On the basis of our findings, we concluded that McPherson (2009) did correctly identify

N. gracillima, but that he mis-identified some plants of this species as NV. macfarlanei,

and that the origin of this error can be traced back to the early years of the 20th century.

As a consequence, McPherson’s (2009) concept of V. gracillima overlooks some

important characteristics of this species, particularly the presence of lid hairs and the

production of two types of pitchers on the climbing stems. In distinguishing N. alba

and N. gracillima, he compared the upper pitchers of the former to the intermediate

pitchers of the latter. In most instances, this oversight would be of little consequence,

but in the case of N. alba and N. gracillima, it is the principle reason for the lack of

clarity surrounding N. gracillima.

We tentatively support McPherson’s (2009) decision to reinstate N. alba.

However, McPherson (2009, p.567) is incorrect in stating that:

...With its white upper pitchers, N. alba could be confused with populations of N.

macfarlanei that exhibit similar colouration, such as those from Mount Berincang (or

Brinchang) in the Cameron Highlands. Nepenthes alba can be distinguished from that

species by the lack of simple white hairs on the lower surface of its lid, which are a

consistent feature of V. macfarlanei.

42 Gard. Bull. Singapore 64(1) 2012

Lid hairs are present and prominent on both the lower and upper pitchers of

N. alba, emphasising its close relationship with N. gracillima and N. macfarlanei.

For some reason, these have not been detected or emphasised previously, even

though they are easily seen, both in the field and on herbarium specimens. Table

1 presents the morphological and ecological characteristics that we consider to be

stable and informative for distinguishing N. alba, N. gracillima and N. macfarlanei,

but we contend that these three species are very closely related and exceptions to

these character combinations are likely to occur. Further comments on the features of

individual specimens that assist in delineating the species are provided below.

The montane Nepenthes species of Gunung Tahan

The four montane Nepenthes species from Gunung Tahan that we recognise are

presented below. Revised, comprehensive descriptions for N. alba and N. gracillima

are provided, for the first time in the case of N. alba. Our description of N. gracillima

is the first since Macfarlane (1908) that does not include material of other taxa.

Key to montane Nepenthes of Gunung Tahan

la.. Lower'surface of pitcher lid always lackimp Waits \....¢-c-<.s.0:.01eseeseeree enter eee 2

b. Lower surface of pitcher lid with short, simple white or red hairs ...................... 3

2a. Margins of the leaf blade not decurrent along the internode, margins of rosette

leaves plabbrouss ic isch i255 cee ae eee ee ee N. sanguinea

b. Margins of the leaf blade decurrent along the internode, margins of rosette leaves

lined with short red or wihitte hates: ccserse eee tere ree eee N. benstonei

3a. Lower pitchers funnel shaped above the hip; pitcher lids circular, strongly cordate

at the base; spur < 5 mm long on mature plants; peristome teeth indistinct or

very short, < 0.2 mm; upper pitchers ivory white in colour with pink specks or

occasionally pale yellowac.::22:. ee eee N. alba

b. Lower pitchers only slightly contracted at the hip, cylindrical above; pitcher lids

ovate, not strongly cordate at the base; spur up to 12 mm long on mature plants;

peristome teeth distinct 0.5—5.0 mm; upper pitchers green, heavily marked with

dark purple/browil SpeckS.2).ccinc eset ee ee N. gracillima

Nepenthes alba Ridl.

Nepenthes alba Ridl., Fl. Mal. Pen. 3 (1924) 22; McPherson, Pitcher Plants of the Old

World 1 (2009) 557-567, f. 300-306. TYPE: Wray & Robinson 5411 (lecto SING,

isolecto BO), Peninsular Malaysia, Pahang, G. Tahan, 1500 m, 3 June 1905.

Nepenthes on Peninsular Malaysia's Gunung Tahan 43

Nepenthes singalana auct. non Becc.: Macfarl. in Engl., Pflanzenr. 4, 3 (1908) 47,

partim; Macfarl., J. As. Soc. Beng. 75, 3 (1914) 282. — Nepenthes bongso auct. non

Korth.: Ridl., J. Linn. Soc., Bot. 38 (1908) 320. — Nepenthes gracillima auct. non

Ridl.: Danser, Bull. Jard. Bot. Buitenz. III, 9 (1928) 296; Shivas, Pitcher Plants of

Peninsular Malaysia and Singapore (1984) 31; Kiew, J. Wildlife and National Parks

10 (1990) 36; Jebb & Cheek, Blumea 42 (1997) 43; Cheek & Jebb, Fl. Malesiana 15

(2001) 69; Clarke, Nepenthes of Sumatra and Peninsular Malaysia (2001) 131, partim.

Terrestrial climber to 5 m tall. Stems cylindrical, 3-6 mm diameter. Rosette leaves

coriaceous, sessile, blade narrowly oblong-elliptic, up to 6 cm long x 1.5 cm wide,

apex acute, base cuneate, margins sometimes decurrent for 2—3 mm; tendrils up to 6

cm long, without a curl. Leaves of the short shoots coriaceous, sessile, blade oblong-

elliptic, wp to 8 cm long x 2 cm wide, apex acute, base cuneate, clasping the stem for

3/4 of its circumference, margins not decurrent; tendrils up to 10 cm long, without a

curl. Leaves of the climbing stems coriaceous, sessile, blade oblong to lanceolate, up

to 9 cm long, up to 2 cm wide, apex generally acute, base cuneate, clasping the stem

for 1/2—1/3 its circumference, margins not decurrent; tendrils up to 9 cm long, usually

curled or with a kink in the middle. Longitudinal nerves of all leaf types obscure,

three on each side of the midrib; pennate nerves numerous, reticulate, often indistinct;

tendril insertion simple. Pitchers of seedling rosettes arising gradually from the tendril,

ovate in cross section, infundibular in the lower 1/3—1/2, abruptly contracted at the hip,

which is pronounced; upper parts cylindrical to slightly infundibular throughout; up

to 5 cm tall x 1 cm wide; two wings, up to 2 mm wide, bearing multicellular fringe

elements up to 4 mm long, run from the top to the bottom of the pitcher at the front;

mouth ovate, acute near the lid, oblique, concave; peristome loosely cylindrical at the

front and sides, flattened and slightly expanded towards the rear, 2-3 mm wide, ribs

distinct, 0.2 mm apart, outer edge entire, inner edge with distinct, but minute teeth up

to 0.1 mm long; lid circular, base cordate, no appendages, lower surfaces covered with

simple, white or red hairs, up to 2 mm long, nectar glands large, numerous, round to

elliptic, crater-like, 0.5—0.7 mm diameter; spur simple, or simple with two short, thick

bristles as the apex, up to 3 mm long. Lower pitchers arising abruptly from the tendril,

infundibular in the lower 1/3—1/2, abruptly contracted at the hip, which is pronounced,

slightly infundibular above; up to 9 cm high x 2 cm wide; mouth round, up to 2 cm

long x 2 cm wide, oblique, concave; peristome broadly cylindrical at the front and

sides, expanded towards the rear, up to 5 mm wide; teeth distinct but very short, up

to 0.2 mm long near the apex; lid circular, slightly larger than the mouth up to 2.5

cm long x 2.5 cm wide, strongly cordate at the base; spur simple, up to 6 mm long,

usually bent downwards; other parts as for the rosette pitchers. Upper pitchers arising

abruptly or gradually from the tendril, narrowly infundibular in the lower 1/4—1/3,

with a pronounced hip; upper parts slightly infundibular; up to 12 cm tall x 3 cm wide;

mouth ovate, oblique, not extended into a pronounced neck at the rear; peristome

loosely cylindrical, up to 5 mm wide, slightly wider towards the apex, teeth distinct but

short, up to | mm long; lid circular, strongly cordate at the base, lower surface covered

with short, simple hairs; spur simple, bent downwards, up to 5 mm long; other parts

44 Gard. Bull. Singapore 64(1) 2012

as for the rosette pitchers. Male inflorescence a raceme, peduncle up to 10 cm long,

rachis up to 15 cm long; partial peduncles 2 flowered at the base, 1—2 flowered above,

3-4 mm long, with a filiform bract, up to 5 mm long; tepals elliptic, 3.5 x 2 mm;

androphore 2.5 mm long; anther head 1.5 x 1.5 mm. Female inflorescence a raceme,

peduncle up to 10 cm long, rachis up to 8 cm long; fruit valves 18 x 3 mm approx.

Indumentum of very simple, short hairs, up to 0.05 mm long, in the axils, around the

exterior apex of the pitchers, and inflorescence; simple red or white hairs, up to 1 mm

long, on the lower surface of the lid. Colour of living rosette pitchers dark purple

throughout, overlain with darker purple flecks; intermediate pitchers light greenish-

white, overlain with dark brown-purple flecks; upper pitchers ivory white throughout,

sometimes with variable amounts of rose coloured flecks, sometimes heavily suffused

with red pigment on the outer surfaces.

Observations on Gunung Tahan. Along the western trail to the summit of Gunung

Tahan, March 30-31, 2011. An isolated population occurs at N 4.6355°, E 102.2050°,

1520 m asl. Substantial populations were observed between N 4.6304°, E 102.2142°,

1656 m asl (near Bonsai Camp) and N 4.6288°, E 102.2268°, 1930 m asl (Bukit Botak

Camp).

Comments. This 1s the only Nepenthes that is abundant in the short padang vegetation

on the uppermost parts of Gunung Tahan. The white upper pitchers are distinctive

among the taxa from Gunung Tahan and on the basis of the characteristics in Table 2,

this species can be reliably distinguished from N. gracillima. However, the differences

between the taxa are slight, and there is merit to the argument that both N. gracillima

and N. alba are derived from N. macfarlanei and have diverged due to their isolation

and exploitation of different habitats on Gunung Tahan.

Kiew’s (1990) statement that pitchers on rosettes of N. alba are purple,

then switch to white as the plant climbs into the canopy, is correct. However, when

it switches from producing lower to upper pitchers, the first few upper pitchers are

effectively an intermediate form that is richly coloured with dark and light pigments,

like the upper pitchers of NV. gracillima. These “intermediate” N. a/ba pitchers are well

preserved in Strugnell & Sow 42878 (KEP), but are not readily observed on Gunung

Tahan as few plants produce them at any given time. They are difficult to distinguish

from the upper pitchers of N. gracillima, but they are produced at a different stage in

the plants’ growth cycle — N. gracillima produces these pitchers for extended periods

at the tops of the longest climbing stems, whereas in N. a/ba they are an intermediate

form that is only produced for a very short time; the plant soon switches to producing

the distinctive, ivory white upper pitchers that it is noted for.

Like McPherson (2009), we saw no obvious examples of natural hybrids

between N. alba and N. gracillima. Although natural hybrids of Nepenthes have

received considerable attention from researchers (see Clarke 1997, 2001), they are

generally rare in undisturbed vegetation such as occurs on Gunung Tahan, even in open

areas. The two species appear to be reproductively isolated (possibly by flowering

times, as there are several sites where they grow together, indicating a lack of spatial

Nepenthes on Peninsular Malaysia's Gunung Tahan 45

isolation). Nepenthes hybrids are generally common in sites that have been recently

disturbed, such as landslips or sites of human activities. In such places, plants often

flower at unusual times, causing the breakdown of temporal isolation mechanisms.

Collections from Gunung Tahan examined: Chua 41539 (KEP), Haniff 7890 (SING), 7891

(BO), Holttum 20666 (BO, SING), 20644, fragment bearing smaller pitchers (SING); Kiew

2450 (KEP, SING), 4064 (KEP), Kloss 12227, 12134 (BO), Lim 56340, 56344 (KEP), Ng

020915 (KEP), Ng 1478 (KEP, SING), Ng 020954 (KEP), Noramly Muslim s.n., 4603 second

sheet, s.n. (KEP), Ridley 16097 (K, sheet barcoded K000651564), Wray & Robinson 5411

(SING), Strugnell & Sow 42878 (KEP), Wong & Wvyatt-Smith W58 (KEP).

Nepenthes benstonei C.Clarke

Nepenthes benstonei C. Clarke, Sandakania 13 (1999) 79, f. 1-3; Cheek & Jebb, FI.

Malesiana 15 (2001) 40; Clarke, Nepenthes of Sumatra & Peninsular Malaysia (2001)

108, f. 105-107; McPherson, Pitcher Plants of the Old World I (2009) 568, f. 307—

311. TYPE: Clarke s.n., (holo KEP, iso BO, K, L, SAN, SING), Peninsular Malaysia,

Kelantan, Bukit Bakar, 450-550 m, 24 July 1998.

Nepenthes alata auct. non Blanco: Danser, Bull. Jard. Bot. Buitenz. III, 9 (1928) 258,

partim. — Nepenthes gracillima auct. non Ridl.: Kiew, J. Wildlife and National Parks

10 (1990) 36; Jebb & Cheek, Blumea 42 (1997) 43; Cheek & Jebb, Fl. Malesiana 15

(2001) 69, partim. — Nepenthes sanguinea auct. non Lindl.; Jebb & Cheek, Blumea

42 (1997) 79, quae = N. benstonei C. Clarke et N. sanguinea Lindl.

Description. See Clarke (1999, 2001); Cheek & Jebb (2001).

Observations on Gunung Tahan. N 4.6524°, E 102.1875°, 895 m asl, and surrounding

areas, March 30, 2011.

Comments. N. benstonei is abundant in dense vegetation on steep ridges, at 800—1350

m asl. On the western route to Gunung Tahan, it grows at the edges of open, disturbed

sites, such as landslips, and along the trail itself. Some plants also grow in dense forest;

these are generally inconspicuous and produce few pitchers.

Collections from Gunung Tahan examined. Holttum 20643 (SING), Ridley 16097 (SING, K

(sheet barcoded K000651565 only).

Nepenthes gracillima Rid.

Nepenthes gracillima Ridl., J. Linn. Soc., Bot 38 (1908) 320; Macfarl. in Engl.

Pflanzenr. 4, 3 (1908) 38; J. As. Soc. Beng. 75, 3 (1914) 282; McPherson, Pitcher

Plants of the Old World 1 (2009) 581-586, f. 316-317. TYPE: Wray & Robinson 5309

46 Gard. Bull. Singapore 64(1) 2012

(lecto SING, isolecto BO), Peninsular Malaysia, Pahang, Gunung Tahan, 990 m, 29

May 1905.

Nepenthes gracillima auct. non Ridl., Danser, Bull. Jard. Bot. Buitenz. II, 9 (1928)

296; Shivas, Pitcher Plants of Peninsular Malaysia and Singapore (1984) 31, quae

pro parte = N. ramispina Ridl., N. gracillima Ridl., N. alba Ridl. et N. macfarlanei

Hemsl. — Nepenthes gracillima auct. non Ridl., Kiew, J. Wildlife and National Parks

10 (1990) 36; Jebb & Cheek, Blumea 42 (1997) 43; Cheek & Jebb, Fl. Malesiana

15 (2001) 69, quae pro parte = N. gracillima Ridl., N. alba Ridl. et N. benstonei C.

Clarke.

Terrestrial or epiphytic climber to 6 m tall. Stems cylindrical, 3—6 mm diameter. Rosette

leaves thin-coriaceous, sessile, blade narrowly oblong-elliptic, up to 6 cm long x 1.5

cm wide, apex acute, base cuneate, margins sometimes decurrent for 3—4 mm; tendrils

up to 10 cm long, without a curl. Leaves of the short shoots coriaceous, sessile, blade

elliptic to sub-spathulate, wp to 18 cm long x 5 cm wide, apex acute, base cuneate,

clasping the stem for 3/4 of its circumference, margins not decurrent; tendrils up to

35 cm long, without a curl. Leaves of the climbing stems coriaceous, sessile, blade

oblong or narrowly elliptic-oblanceolate, apex generally acute, base cuneate, clasping

the stem for 1/2—1/3 its circumference, margins not decurrent; tendrils up to 30 cm

long, usually curled or with a kink in the middle. Longitudinal nerves of all leaf types

obscure, three on each side of the midrib; pennate nerves numerous, reticulate, often

indistinct; tendril insertion simple. Pitchers of seedling rosettes arising abruptly from

the tendril, ovate in cross section, infundibular in the lower 1/6, ovoid for the next

|—2/6, with a pronounced hip; upper parts cylindrical throughout; up to 8 cm tall x 1.5

cm wide; two wings, up to 3 mm wide, bearing multicellular fringe elements up to 8

mm long, run from the top to the bottom of the pitcher at the front; mouth ovate, acute

near the lid, oblique, concave; peristome loosely cylindrical at the front and sides,

flattened and slightly expanded towards the rear, 2-3 mm wide, ribs distinct, 0.3 mm

apart, outer edge entire, inner edge with distinct, but minute teeth up to 0.2 mm long;

lid broadly ovate, slightly peaked towards the apex, base simple or sometimes slightly

cordate, no appendages, lower surfaces sparsely to densely covered with simple,

white or red hairs, up to 3 mm long, particularly towards the margins, nectar glands

large, numerous, round to elliptic, crater-like, 0.5—0.7 mm diameter; spur flattened,

simple or sometimes with two short, thick bristles at the apex, or divided into two or

three branches, up to 5 mm long. Lower pitchers ovoid or broadly cylindrical with a

pronounced hip 1/3—1/2 way up from the bottom, up to 18 cm high x 7 cm wide, mouth

ovate up to 5 cm long x 4 cm wide, oblique; peristome flattened, expanded towards

the rear, up to 25 mm wide; teeth distinct, up to 5 mm long near the apex; lid ovate,

up to 5 cm long x 3.5 cm wide, barely or not cordate at the base; spur simple, up to 12

mm long, straight; other parts as for the rosette pitchers. /ntermediate pitchers arising

abruptly from the tendril, narrowly infundibular in the lower half, with a pronounced

hip; upper parts cylindrical or slightly infundibular; up to 22 cm tall x 4 cm wide;

mouth ovate, oblique, not extended into a pronounced neck at the rear; peristome

Nepenthes on Peninsular Malaysia s Gunung Tahan 47

loosely cylindrical, up to 8 mm wide, slightly wider towards the apex, teeth distinct

but short, up to 3 mm long; lid ovate, not cordate at the base, lower surface covered

with short, simple hairs, particularly towards the margins, spur simple, straight, up

to 12 mm long; other parts as for the rosette pitchers. Upper pitchers similar to the

intermediate ones but much smaller and narrower, up to 10 cm long x 1.5 cm wide.

Male inflorescence a raceme, peduncle up to 10 cm long, rachis up to 20 cm long:

partial peduncles 2 flowered at the base, 1—2 flowered above, 3-4 mm long, with a

filiform bract, up to 5 mm long; tepals elliptic, 3.5 x 2 mm; androphore 2.5 mm long:

anther head 1.5 x 1.5 mm. Female inflorescence a raceme, peduncle up to 10 cm

long, rachis up to 8 cm long; fruit valves 18 x 3 mm approx. Indumentum of simple,

very short hairs, up to 0.05 mm long, in the axils, around the exterior apex of the

pitchers, and inflorescence; simple red or white hairs, up to 1 mm long, on the lower

surface of the lid, particularly towards the margins. Colour of living pitchers dark

green, heavily marked with dark purple-brown speckles, occasionally suffused with

dark pink colouring throughout.

Observations on Gunung Tahan. Between N 4.6456°, E 102.1926°, 1276 m asl, below

Belumut Camp, and N 4.6287°, E 102.2183°, 1694 m asl, above Bonsai Camp, on the

western route to Gunung Tahan, March 30-31, 2011.

Comments. Nepenthes gracillima is one of asmall number of species that is consistently

trimorphic with regard to pitcher structure. Nearly all Nepenthes are dimorphic and

plants of many species occasionally produce one or two “intermediate” pitchers as

they switch from producing lower to upper pitchers (Clarke 1997, 2001), but it is very

unusual for intermediate pitchers to be the predominant form on climbing stems, as in

this species. Prior to Cheek & Jebb’s (2001) revised description of N. gracillima, all

previous treatments were based on either upper pitchers alone, or specimens of both

N. alba and N. gracillima. Good collections of lower and intermediate pitchers of N.

gracillima were made, but due to the presence of lid hairs, all of these were identified

as N. macfarlanei. For instance, Cheek & Jebb (2001, Fig. 8a, e & f) used Ridley 16174

(K) in their illustration of N. gracillima. The components of the drawing derived from

this specimen include a fragment of a climbing stem bearing an upper pitcher, and

show detail of the pitcher lid. However, the duplicate of this collection at SING is a

basal rosette bearing a lower pitcher of NV. gracillima, but Danser (1928) identified it

as N. gracillima x N. macfarlanei, whereas Jebb & Cheek (1997) identified it as N.

macfarlanei. As a consequence, taxonomists based their concepts of N. gracillima on

the least common pitcher type that this species produces, and which bears a strong

resemblance to the upper pitchers of NV. alba. It is therefore unsurprising that confusion

about these taxa arose, particularly among researchers who only examined herbarium

specimens, as these are comprised of stem fragments that bear only one type of pitcher,

making it impossible to associate the lower, intermediate and upper pitchers of any

given taxon with confidence (particularly when collections comprising several sheets

with the same number are split among different herbaria).

48 Gard. Bull. Singapore 64(1) 2012

Regardless of any shortcomings in the herbarium material, it is perplexing

that no researchers who visited Gunung Tahan prior to us traced the climbing stems of

N. gracillima to the ground. Had this been done, it is possible that much of the recent

confusion surrounding N. alba, N. gracillima and N. macfarlanei could have been

avoided. Clarke & Moran (2011) stress the importance of making complete collections

of Nepenthes for herbaria, and our findings in this study emphasise the need for detailed

and accurate field observations in interpreting closely related taxa.

Collections from Gunung Tahan examined: Chua 26664 (KEP), Haniff 7890, 8306 (SING), Holttum

20644 (BO, SING, fragment bearing larger pitchers), K/oss 12211, 12212 (BO), 12259 (SING),

12297 (BO), Lim 56363 (KEP), Ng 1448, 020961 (KEP), Noramly Muslim 4603 first sheet (KEP),

Ridley 16096 (SING), 16098 (SING), 16174 (K, SING), Wong & Wvatt-Smith 60 (KEP).

Nepenthes sanguinea Lindl.

Nepenthes sanguinea Lindl., Gard. Chron. (1849) 580; Danser, Bull. Jard. Bot.

Buitenz. II, 9 (1928) 366, f. 20; Shivas, Pitcher Plants of Peninsular Malaysia and

Singapore (1984) 43, f. 4-5, 27-28; Kiew, J. Wildlife and National Parks 10 (1990)

36; Cheek & Jebb, Flora Malesiana 15 (2001) 40; Clarke, Nepenthes of Sumatra &

Peninsular Malaysia (2001) 108; McPherson, Pitcher Plants of the Old World I (2009)

568. TYPE: Griffith 4411 (K) Malaysia, Johor, Gunung Ledang.

Nepenthes pumila Griff., Post. Papers 4 (1854) 349; Jebb & Cheek, Blumea 42 (1997)

79, quae pro parte = N. benstonei C. Clarke et N. sanguinea Lindl.

Descriptions. See Lindley (1849), Jebb & Cheek (2001), Clarke (2001).

Observations on Gunung Tahan. Near Belumut Camp on the western route to Gunung

Tahan, N 04.6401°; E 102.1964°, 1450 m asl, on Gunung Tahan on April 1, 2011.

Comments. This is probably the only Nepenthes species from Gunung Tahan that has

not been the subject of some taxonomic confusion. Kiew (1990) states that this species

is not particularly common on Gunung Tahan, but several collections have been made

and conform with Lindley’s description of this species. We observed only one plant of

N. sanguinea on the western route to Gunung Tahan (see above), but were in no doubt

that it belonged to this species. We suspect that it may not be that rare on Gunung

Tahan, but as neither the southern or western trails pass through suitable habitats for

N. sanguinea and most plants appear to be epiphytic, there are few specimens or visual

records.

Collections from Gunung Tahan examined: Wong & Wyatt-Smith 59 (KEP), Holttum 20643

(SING).

Nepenthes on Peninsular Malaysia's Gunung Tahan 49

Summary and conclusions

The montane Nepenthes of Peninsular Malaysia are among the most difficult to

interpret, due to their collective similarities, but also minor differences that appear to

reflect recent reproductive isolation of populations on different mountains. More than

any other group within the genus, accurate identifications depend on both detailed

field observations and careful examinations of herbarium material. The present study,

along with those of Kiew (1990), Jebb & Cheek (1997), Cheek & Jebb (2001) and

McPherson (2009), represents an important step towards a resolution of the confusion

surrounding the montane Nepenthes of Peninsular Malaysia. This will facilitate a

better understanding of the ecology, biogeography and conservation of these species,

most of which are found nowhere else.

References

Cheek, M. & Jebb, M. (2001) Nepenthaceae. F/. Males. 15. Netherlands: Leiden.

Clarke, C.M. (1997) Nepenthes of Borneo. Malaysia, Sabah, Kota Kinabalu: Natural History

Publications (Borneo).

Clarke, C.M. (1999) Nepenthes benstonei (Nepenthaceae) — a new pitcher plant from Peninsular

Malaysia. Sandakania 13: 79-87.

Clarke, C.M. (2001) Nepenthes of Sumatra & Peninsular Malaysia. Malaysia, Sabah, Kota

Kinabalu: Natural History Publications (Borneo).

Clarke, C. & Moran, J.A. (2011) Incorporating ecological context: a revised protocol for the

preservation of Nepenthes pitcher plant specimens (Nepenthaceae). Blumea 56: 225-228.

Danser, B.H. (1928) The Nepenthaceae of the Netherland’s Indies. Bull. Jard. Bot. Buitenzorg.

Serie III 9: 249-438.

Jebb, M. & Cheek, M. (1997) A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42(1):

1-106.

Kiew, R. (1990) Pitcher plants of Gunung Tahan. J. Wildlife Natl. Parks (Malaysia) 10: 34-37.

Macfarlane, J.M. (1908) Nepenthaceae. In: Engler, A. Das Pflanzenreich IV, Il, Heft 36, 1-91.

McPherson, S. (2009) Pitcher Plants of the Old World, vol. 1. United Kingdom, Poole: Redfern

Natural History Publications.

Ridley, H.N. (1908) On a collection of plants made by H.C. Robinson and L. Wray from

Gunong Tahan, Pahang. J. Linn. Soc., Bot. 38: 320.

Ridley, H.N. (1909) The flora of the Telom and Batang Padang valleys. J. Fed. Malay States

Mus. 4: 59.

Ridley, H.N. (1915) The botany of Gunung Tahan, Pahang. J. Fed. Malay States Mus. 6: 127—

202.

Ridley, H.N. (1924) The Flora of the Malay Peninsula 3: 21-25.

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Gardens’ Bulletin Singapore 64(1): 51-131. 2012 51

A taxonomic revision of Amischotolype (Commelinaceae)

in Asia

Helena Duistermaat

Singapore Botanic Gardens,

1 Cluny Road, Singapore 259569

Present address:

NCB Naturalis, section Nationaal Herbarium Nederland,

P.O. Box 9514, 2300 RA Leiden, The Netherlands

duistermaat(@nhn.leidenuniv.nl

ABSTRACT. A taxonomic revision of the Indomalayan part of the paleotropical genus

Amischotolype Hassk. (Commelinaceae) reveals 22 species in Asia, of which eight are described

as new (A. hbarbarossa Duist., A. divaricata Duist., A. dolichandra Duist., A. lobata Duist., A.

parvifructa Duist., A. pedicellata Duist., A. strigosa Duist., A. welzeniana Duist.), and three

are new combinations (A. hirsuta (Hallier f.) Duist., 4. /eiocarpa (Hallier f.) Duist., 4. rostrata

(Hassk.) Duist.). The status of the closely related genus Porandra Hong is discussed although

results of a molecular study are required to make a final decision on its generic status. For now

the genera are kept separate, but the species of Porandra are included in the key to the species

of Amischotolype.

Keywords. Amischotolype, Commelinaceae, Forrestia, morphology, Porandra, revision,

taxonomy

Introduction

Amischotolype is a genus of the Commelinaceae (Spiderwort family), a family of about

650 species in 41 genera with a cosmopolitan distribution (although with no native

species in Europe; Faden 1998). Faden and Hunt (1991), in their Commelinaceae

classification based upon morphology, anatomy, palynology and cytology, placed

Amischotolype in the subtribe Coleotrypinae Faden & D.R.Hunt with the African genus

Coleotrype and Asian genus Porandra. Important characters of the subtribe included

leaf epidermal cells with silica, axillary inflorescences, flowers with 6 fertile stamens

and seeds with a linear hilum. Faden & Hunt’s classification was later supported by

cladistic analysis of morphological data (Evans et al. 2000) and by rbcL sequences

combined with morphology (Evans et al. 2003). The chromosome numbers (2n = 18,

36 with a basic chromosome number x = 9) and the karyotype of several species of this

subtribe are very similar (Yuan & Yang 2006).

Generic delimitation

The first species were described from Java by Blume (1827) in Campelia, as C.

mollissima and C. marginata. Campelia was described by Richard (1808) based on a

a2 Gard. Bull. Singapore 64(1) 2012

species from tropical America. A third Old World species was discovered in New Guinea

and described by Richard (1834) as a new genus associated with the Colchicaceae

and Asparagaceae. He named it Forrestia (with a single species F. hispida), after the

British explorer who was the first to visit New Guinea, Thomas Forrest (1729-1802)

of the East India Company. Hasskarl (1852), unaware of A. Richard’s work, added

another species from Java to Campelia (though with an illegal name, see discussion

under A. glabrata). Several years later, when Hasskarl received material on loan from

Berlin, he realised that the Asian species of Campelia, with sessile inflorescences,

were very different from the American species which had pedunculate inflorescences.

Comparison of the Javanese species with Richard’s description of Forrestia (1834;

in Kunth 1843) which states “flores rubri, supra vaginam exserti...” led Hasskarl to

translate this as “inflorescence terminal”, which was manifestly different from his

species that had axillary inflorescences. Hasskarl (1863) transferred the Asian species

of Campelia to a new genus next to Forrestia: Amischotolype. The name refers to the

sessile glomerules of flowers, the character that immediately distinguishes it from the

American species of Campelia (which are now united with 7radescantia, see Hunt

1986). A year later, after having studied the type material of Forrestia, Hasskarl (1864)

transferred his three species of Amischotolype to Forrestia. He also added two new

species, one from India (F’. hookeri) and one from Java (F. rostrata), bringing the total

number of species in Asia to six. In the following years, more than 10 species were

described from Asia, the latest A. sphagnorrhiza Cowley (Cowley 1996). The first

African species was described by Clarke (1881, in Buforrestia C.B.Clarke), followed

by three more species (Schumann 1897, Clarke 1901, De Wildeman 1909).

From Hasskarl (1864) onwards, the generic name of Forrestia was commonly

(but not exclusively) used, until Pichon (1946: 235) pointed out that the Forrestia

of Richard (1834) was a later homonym of Forrestia of Rafinesque (1806), the

latter a genus in the Rhamnaceae, and argued that the correct genus name should be

Amischotolype Hassk. While Richard (1834) did mention the existence of a genus

Forrestia Schweack. in the Rhamnaceae (see De Jussieu 1820; the author name is a

mystery and perhaps refers to Rafinesque’s mother’s name: Schmaltz), he considered

the name available for use in the Commelinaceae because Forrestia Raf. was united

with Ceanothus. A proposal to conserve Forrestia A.Rich. in the Commelinaceae

(Babu & Dutta 1968) was rejected (see discussion by Rao 1971).

Hong (1974) described the genus Porandra from China as closely related

to Amischotolype, differing by its climbing habit with branched and basally ‘woody’

stems, and by having thecae opening by apical pores (hence the name). These

characters, however, are not unique for Porandra, as noted by Faden (1998: 112). This

present revision reveals that 4. rostrata (Hassk.) Duist. shares with Porandra both

the branched and scrambling stem and thecae opening by an apical pore. A straggling

habit also occurs in 4. divaricata Duist., A. dolichandra Duist., A. glabrata Hassk.

and A. hispida (A.Rich.) D.Y.Hong, while the opening of the thecae by an apical pore

also occurs in A. gracilis (Ridl.) 1.M.Turner (but there also by a longitudinal slit) and

A. griffithii (C.B.Clarke) I.M.Turner. Faden (1998) argues that the two genera should

be united based on the morphological evidence, and he is probably correct. However,

Amischotolype in Asia

pending results from on-going molecular studies, the genera are here kept separate, as

in Thitimetharoch et al. (2003). Nonetheless, it is interesting to note that the few field

observations of anther colour in Porandra state violet (but sometimes only so at the

margins: R.Geesink 6850), whereas in Amischotolype they seem to be white to yellow

(with rare exceptions in A. marginata (Blume) Hassk.). For 10 out of the 22 Asian

species, however, this information is lacking.

Need for revision

Amischotolype has a paleotropical distribution and the species typically grow in a

forest environment, even surviving for a long time after disturbance of the forest, on

rather dry to swampy soils. The most recent monographic revision of the genus was

that of Clarke (1881). Since then, 12 more species have been described. For Asia, local

flora treatments exist for the Himalaya (Hara 1966: | sp.), China (Hong & DeFilipps

2000: 2 spp.), Taiwan (Wang et al. 2000: 1 sp.), Japan and the Ryukyu Islands (Walker

1976: 1 sp.), Indo-China (Cherfils 1937: 7 spp.), Vietnam (Ho 1993: 3 spp., one species

with two forms and two varieties), Peninsular Malaysia (Ridley 1924: 7 spp.: these

all accepted in the checklist by Turner 1997), Singapore (Keng et al. 1998: 2 spp.),

Philippines (Merrill 1925: 1 sp.), Borneo (Hallier 1916: 4 spp., one species with four

varieties and one with two varieties). Java (Backer & Bakhuizen van den Brink 1968:

1 sp. with three forms), and New Guinea (Hallier 1913: 1 sp. with three varieties;

Lauterbach 1913: 1 sp. with two forms). Nomenclature and species delimitation in

these treatments differ considerably. Identification of species is further complicated

because most authors provide no key to their taxa. Several authors (e.g., Hasskarl

1864, Backer & Bakhuizen van den Brink 1968) considered the amount of indument

on the sheath and the lower surface of the leaf blade of high taxonomic importance, but

this later proved to be variable within many species.

Most problematic is the application of the name A. hispida(A.Rich.) D.Y.Hong,

now regarded as an East-Malesian species not reaching the Asian continent. Hooker

(1864), who had not seen the type material, matched several collections from Java,

Sumatera, Penang, Singapore, Sikkim and Assam with Richard’s description of F.

hispida and concluded that the distribution of the species ranged from NE India to

New Guinea. Only few collections were cited precisely by Hooker, but those included

specimens of at least A. marginata (Wallich s.n., Penang) and A. gracilis (Walker

155, Singapore). The epithet /ispida is now erroneously used for continental Asian

material (e.g., Hong & DeFilips 2000), most of which is correctly 4. glabrata. Owing

to this confusion, and because a number of species are quite variable, especially in the

amount of indument on the leaves, estimations of the number of species for the genus

differ considerably, ranging from six (Walker 1976) to 20 (Hong 1974). A revision of

the genus seemed therefore much needed. The African species are excluded from this

revision. There is no overlap in species between Africa and Asia.

Characters that have proved valuable for the delimitation of Asian species of

Amischotolype include the indument of the internodes, and upper surface and margin

of the leaf blade, the inflorescence position, the (relative) length and indument of the

sepals, the length of anthers, and size and indument of the capsules. The present revision

54 Gard. Bull. Singapore 64(1) 2012

of Amischotolype recognises 22 species for Asia, of which eight are new species (A.

barbarossa Duist., A. divaricata Duist., A. dolichandra Duist., A. lobata Duist., A.

parvifructa Duist., A. pedicellata Duist., A. strigosa Duist., A. welzeniana Duist.).

Three new combinations are made (A. hirsuta (Hallier f.) Duist., A. leiocarpa (Hallier

f.) Duist., A. rostrata (Hassk.) Duist.). Amischotolype glabrata is the most widespread

species, occurring from E Pakistan through China to S. Japan (Ruykyu Islands) and

south to Nusa Tenggara (Lombok), excluding the Philippines and Sulawesi.

Amischotolype Hassk.

Amischotolype Hassk., Flora 46: 392 (1863); Pichon, Notul. Syst. 12: 233 (1946); Hara,

Fl. East. Himalaya 1: 399 (1966); Morton, J. Linn. Soc., Bot. 60: 167-221 (1967);

Ho, Ill. Fl. Vietnam 3: 477 (1993); Keng, Chin & Tan, Conc. Fl. Singapore 2: 23

(1998); Hong & DeFillips in Wu & Raven (eds.), Fl. China 24: 23 (2000); Wang,

Chen & Peng in Huang (ed.), Fl. Tatwan, ed. 2, 5: 153 (2000). TYPE SPECIES: A.

glabrata Hassk. (designated by Pichon 1946).

Forrestia A.Rich. in Dumont d’Urville, Voy. Astrolabe 2: 1 (1834); Miquel, Fl. Ned.

Ind. 3: 547 (1855); Hasskarl, Flora 47: 625 (1864); Hasskarl, Commelin. Ind.:

83 (1870); Hooker f., Fl. Brit. India 6: 383 (1894); Ridley, Fl. Malay. Penins. 4:

359 (1924); Merrill, Enum. Philipp. Fl. Pl. 1: 197 (1925); Briickner in Engler &

Prantl, Nat. Pflanzenfam. ed. 2, 15a: 169 (1930): Cherfils in Lecomte, Fl. Gen.

Indo-Chine 6: 852 (1937); Backer & Backh. f., Fl. Java 3: 15 (1968); Walker, Fl.

Okinawa S. Ryukyu I.: 298 (1976); non Rafin. (1806). TYPE SPECIES: F. hispida

A.Rich. (= A. hispida (A.Rich.) D.Y.Hong)

Campelia auct., e.g., Blume, Enum. Pl. Javae 1: 7 (1827); Kunth, Enum. pl. 4: 109

(1843), p.p.; Hasskarl in Miquel, Pl. Jungh.: 154 (1852); Miquel, Fl. Ned. Ind. 3:

546 (1855); non L.C.M. Rich. (1808).

Plant perennial, herbaceous, rather succulent, exudate sticky and colourless. Stem

solid, nodes very short and not thickened. Leaves with involute vernation, alternate

in two ranks on erect part of the stem but at apex spirally arranged, herbaceous to

somewhat coriaceous. Sheath tubular, closed, tightly covering the internode (loose

dry material owing to shrinkage of the stem), the lower withering and deciduous as

the plant matures, generally green (in species description mentioned only if otherwise

coloured), longitudinally veined, more weekly transversely veined, apex truncate. Leaf

blade (rather) thickly herbaceous, oblong to linear-lanceolate (2.3—7.4 times as long

as wide), smooth or bullate, generally green (in species description mentioned only

if otherwise coloured), longitudinally veined often with veins alternately thickened

and more obvious on lower surface than on upper surface (except in A. /obata where

they are very obvious on both surfaces), oblique transverse veins present (though

not always distinct), base attenuate, apex acuminate to caudate, submarginal hairs (a

row of hairs on the upper or lower surface of the leaf blade along the entire margin

and here not included as leaf blade indumentum) present or absent. Inflorescence

Amischotolype in Asia 55

a more-or-less condensed thyrse, strictly axillary, breaking through the base of the

sheath if leaf is present; bracts broadly to narrowly triangular to rhomboid, 2—25 by

2.5—10 mm, thick herbaceous to coriaceous, glabrous to ciliate or hispid, pale green to

purplish, midvein keeled, semiamplexicaulous, more-or-less decurrent on branchlet,

apex obtuse to acuminate. Flowers almost actinomorphic, bisexual. Sepals 3, free,

succulent, imbricate, outermost the longest (size given in species description refers

to the longest sepal) and overlapping the inner two, innermost the shortest, persistent,

midvein more-or-less keeled, apex obtuse. Petals 3, free, subequal, membranous,

oblanceolate, obtuse, deliquescing soon after anthesis. Stamens 6, (sub-) equal,

inserted on the receptacle; filaments free, tortuose; anthers basifixed, subglobose to

oblong (deltoid in A. glabrata). Ovary 1, locules 3, equal; style 1, simple; stigma 1,

capitate, apical. Fruit a (usually) dehiscent, succulent, white, pink, red, lilac, (brown-)

purple or green capsule with 3 locules and (1—)2 seeds per locule, 3-lobed at apex or

not, valves finally free to fused for up to 4/5 of length. Seeds uniseriate, reniform,

surface coarsely rugose, hilum linear; aril fleshy.

Chromosomes. Basic chromosome number x = 9 (Morton 1967); mitotic counts: 2n =

18, 36. Published counts of 2n = 20 and 2n = 30 are considered doubtful (see Yuan &

Yang 2006, and references therein).

Distribution. Paleotropic, few (1 to 4) species in tropical Africa, 22 species in S, E and

SE Asia (including India, China, Taiwan, Japan and New Guinea, but not Sri Lanka,

Australia or Oceania).

Ecology. Primary forest and along forest streams, surviving a long time after

disturbance, on rather dry to swampy soils, also on limestone. Altitude: 0-1980 m asl.

Notes. 1. Hasskarl’s (1863) name Amischotolype (Greek, amischos = without stalk;

tolupe = cluster) refers to the sessile glomerules of flowers.

2. The species delimitation in the work of Cherfils (1937), presenting seven

species, is very different from the one presented here. I have seen 24 specimens (P)

that he identified; of these all his F. glabratus, F. hookeri, F. monosperma and F.

mollis belong to A. divaricata, whereas specimens he identified as F. marginatus are

in fact A. glabrata and A. divaricata. In his F. hispida | saw specimens of Pollia sp.,

A. hispida and A. glabrata. His species descriptions can in no way be related to mine

or any other. Moreover, it seems that measurements have been taken from material

belonging to other genera (e.g., sepals 5-35 mm long, 2-13 mm wide for F. glabratus

whereas maximum sepal length in Amischotolype is 18 mm, and maximum width

5.5 mm). Ho (1993) seems largely based on Cherfils (1937). Both these accounts are

largely ignored in the species protologues below. The new combinations made by Ho

(1993) are all invalid, as he did not cite the basionym (McNeill et al. 2006, Art. 33.4).

3. It greatly helped that I had the opportunity, while based in Singapore, to

study living material, both in the field and in cultivation, and make my own collections.

The descriptions are based on my observations on living material of A. glabrata, A.

56 Gard. Bull. Singapore 64(1) 2012

gracilis, A. hookeri, A. irritans, A. marginata, A. monosperma and A. parvifructa,

supplemented with many observations on herbarium material. Unless mentioned

otherwise, the stem is not scrambling and aerial (stilt-) roots have not been observed.

The area between the horizontally creeping rhizome and the erect stem is called the

knee. Hairs on different organs are colourless unless stated otherwise. Whether the

surface of the leaf blades is bullate or smooth cannot be observed from herbarium

material, and is here mentioned only for the species for which field observations are

available. An inflorescence with more than 40 flowers is considered many-flowered.

In some species sepals elongate after anthesis and change colour from whitish or green

to pink or violet. As the colour is lost in herbarium material, it is unknown unless

annotated by collectors. Petals and stamens are difficult to study because they are

lost soon after anthesis. Herbarium material of some species have capsules with three

0.5—1.5 mm long lobes at the apex of the valves. As I have not seen fresh material of

these species, I do not know if these lobes are artifacts, e.g. the result of the drying

of fruits with very deeply depressed apex. The colour of the ripe fruit can vary in a

single species from white to green, pink, red or purple. As far as the colour of the aril

is known for the species, it is orange (to dark red) and strikingly contrasting with the

colour of the capsule.

4. The dichotomous key to the 22 Asian species presented below uses

characters in leaves, inflorescence position, sepals, petals, stamens and fruits. The

characters that separate the two leads best are mentioned first, followed by characters

with more overlap (if present). Not infrequently all characters cannot be observed in

a single specimen. Character synopses are included in several Appendices in tabular

form to allow comparison of species.

5. As Ihave seen in the field, mixed populations of two or three species do occur.

Evidence from herbarium material suggests the same, e.g., mixed in a single accession

or collections with successive numbers from identical locations. The combinations

involve at least the following species: A. barbarossa, A. gracilis, A. hirsuta, A.

irritans, A. laxiflora, A. leiocarpa, A. marginata, A. mollissima, A. monosperma, and

A, parvifructa.

6. The genus is, for the time being, considered separate from Porandra (see

Introduction). I have not seen type material of the three species that have been described

in this genus. Based on 47 collections, 17 of which were cited by Thitimetharoch et al.

(2003) as belonging to the genus Poranda, I recognise two species, P. ramosa Hong

and P. scandens Hong. They are included in the key to the species below. Based on the

descriptions, I consider P. microphylla Wan to be a synonym of P. scandens.

Key to the species

la. Leaf blade hairy on both surfaces, hairs (0.5—)1—4 mm long and often yellow; if

hairs on one surface less than | mm long, then hairs on the other surface at least

1.5 mm long; submarginal hairs present on upper surface ...........c:ccccccceeeeeeeeeeees 2

b. Leaf blade hairy on one surface or glabrous, if hairy on both surfaces then hairs

Amischotolype in Asia 5) 7/

Da:

3a.

4a.

Sas

6a.

Ta.

8a.

shorter than 1 mm long and either at least one surface with hairs up to 0.5 mm

long (excluding submarginal hairs and hairs near or on midvein), or submarginal

PeAISEPIEC SCH MOUN LOW Clu SULA CE occ oo ccxec-saversceconenssooesttvaciesocsonsedesdsossvoacsvaeeostonsecs 6

Sheath (except at mouth) glabrous or with a few hairs at base .............ceeeeeees 3

Sheath with several lines of 2-4(—10) mm long hairs ..................cccsccceseceeseeeeeees =

Sheath 7-13 mm diameter. Sepals in flower and fruit 4.5-7.5 mm long, sparsely

to moderately (rarely densely) hairy with hairs 0.5 mm long. Petals glabrous.

Capsule exceeding sepals by (1.5—)3.5—6 mm ...............:ccscccesseeeeees 6. A. griffithii

Sheath 10-20 mm diameter. Sepals in flower 6.5—10.2 mm long, in fruit 10.5—13

mm long, moderately to rather densely (dark-)red spiny-hairy with hairs1.5—2.5

mm long. Petals on dorsal surface subapically with a 2 mm long spot of 1—1.5 mm

long red spiny hairs. Capsule shorter than sepals by 3-4.5 mm. ............. 10. A. irritans

Sepals 10.5—16(—18) mm long. Capsule shorter than sepals by 4—9 mm, hairs soft;

valves fused for 1/4 to 1/2 of length. [Philippines, New Guinea] ...... 8. A. hispida

Sepals (4.5—)6—10 mm long. Capsule 2 mm shorter to 4 mm longer than sepals,

hairs stiff; valves free (almost to the base). [Sumatera, Borneo] ...................005 5

Sepals glabrous or margins and keels ciliate with colourless hairs 0.5—1 mm

long. Capsule (equaling or) exceeding sepals by up to 4 mm, hairs yellow

ae ee ee eyes woe DEI Sak es Scho dds coe cea seas 7. A. hirsuta

Sepals rather densely hairy with red, spiny hairs 2 mm long. Capsule shorter than

eae MMII MM AITS REO ar = Seec tic eat Sao ceed oecatteeessedesesv.e sav 21. Ay Strigosa

Inflorescence on the creeping, leafless part of the stem (rhizome) on the ground or

at the (leafy) knee just above the ground; peduncle (0—)2—40 mm long ............. fj

Inflorescence on the erect, leafy part of the stem well above the ground, peduncle

o=Saumilone, iatthe knee thenpeduncle/absent:..0...0...0... ccc ccsescseeceesee stevens 10

Stem with many above-ground sphagnum-like orange branched roots. Sheath

glabrous. Leaf blade smooth, lacking submarginal hairs ...... 20. 4. sphagnorrhiza

Stem with above-ground roots absent or simple and whitish to brownish. Leaf

blade bullate. Either sheath with lines of 1-6 mm long hairs and/or leaf blade with

ReNU REET ARUN WET NESOFESE Blip ct o-oo co vats woscn se eos tu Sen slew on <lacaobatens soe casaeesenabecatesduetvoescces 8

Sheath with lines of orange to orange-brown or red hairs. Leaf blade green on

upper surface, (usually) red to purple on lower surface, (6.5—)9—16 cm wide.

Sepals with margins and keel sparsely to rather densely (0.5—)1—2 mm long

orange- to red-brown-hairy. Capsule with orange to red-brown hairs 1—2 mm long

Pear creed Seams Rc LA Eee Senecio Sada aa Aicec coe Saba cueeastvestoceoseveabl 16. A. monosperma

. Sheath usually with lines of (pale) yellow(-brown) hairs, rarely glabrous or with

red, appressed hairs 0.1—0.5 mm long. Leaf blade (2.7—)4—8(—10.6) cm wide.

58 Gard. Bull. Singapore 64(1) 2012

Sepals glabrous to rather densely hairy with 0.2—0.5(—1) mm long colourless- to

red hairs. Capsule glabrous or with colourless hairs 0.1—0.5(—1) mm long ......... 9

9a. Sepals (9—-)10—14 x 1.5—2.5 mm, white, green, pinkish or red-purple in flower and

red, (deep) purple (and green), magenta or violet in fruit. Capsule shorter than

sepals by :2=5 iminl:......:Saee Antes eee eae ee eee 11. A. laxiflora

b. Sepals (6—)7—10 x (2—)3-4(—S5.5) mm, cream to beige or yellow-brown in flower,

pink to (red-) purple in fruit. Capsule 1.5(—2) mm shorter to 1.5(—3) mm longer

that’s pallls:...zscsavdcseecisss <osteeeeees wate ee 14. A. marginata

10a. Sepals and capsule with orange to dark-red or brown hairs ......... 1. A. barbarossa

b. Sepals glabrous or with colourless, white or yellow hairs. Capsule glabrous or

with colourless, white, yellow or rarely brown haltS .............:::cscccsssceesnreceeeeeees 11

lila ‘(Capsule shorter than:-sepals by (1=)2—9 sania seeegeeeee eee eee eee 12

b. Capsule equalling or exceeding sepals by up to 9 MM ..............:ccsccssceteesseeseenres 17

12a. Capsule 3-lobed at apex when dried, lobes 0.5—1.5 mm long ................:cc0008 13

b: Capsule not lobed at apexiwihen! dried. -2 5s scce cc sceeceee ee ee eee 1)

13a. Anthers 2—2.2 mm long. Capsule glabrous; valves fused for 4/5 of length.

[Myanmar, Thailand) .:...4::..22ecteececeeeeees ee 22. A. welzeniana

b. Anthers 1—1.3 mm long. Capsule covered in 0.2—2 mm long hairs; valves free to

fused! for 1/3/of length: (Sumatera, Jaya; Borne oll eeeere-seanenese: cetera ne eee 14

14a. Leaf blade with veins distinct on upper surface, base very abruptly narrowed,

pseudopetiole 3.5—9.5 cm long and 0.5—l1 mm wide winged. Capsule hairs 1—2

mm long, ‘bristly. [Bomeo | wn.c-5: ace ee ee ee 13. A. lobata

. Leaf blade with veins indistinct on upper surface, base very gradually to abruptly

narrowed, pseudopetiole indistinct or up to 4 cm long with wings 1.5—2 mm wide.

Capsule hairs 0.2—1 mm long, soft. [Sumatera, Java] ................ 15. A. mollissima

15a. Flowers with pedicel 2-6 mm long. Leaf blade (19—)28-48 cm long. Capsule

apex slightly depressed)cc.-...:<sscesaee seem tenes ee ee 8. A. hispida

. Flowers without pedicel. Leaf blade 15—28 cm long. Capsule apex obtuse ....... 16

16a. Sepals 7.5—-9 mm long. Filaments glabrous. Anthers 3 mm long. Capsule valves

free, [India] a22.5...exibess Secs Sea ee ee 3. A. dolichandra

b. Sepals (9—)10—14.5 mm long. Filaments with 1-3 mm long hairs in upper half.

Anthers 0.6—1.0 mm long. Capsule valves fused for 1/3 to 1/2 of length. [Pakistan,

E through India and China to Japan, and S to Nusa Tenggara] ......... 4. A. glabrata

17a. Submarginal hairs absent or on lower surface (and sometimes margin) of leaf

blade... .......dccessseeadsaticntenlea tote Gee ieee ee ee ne 18

Amischotolype in Asia 59

18a.

19a.

20a.

Olay

22s

23a.

. Submarginal hairs present on upper surface (and sometimes margin) of leaf blade

Inflorescence 4—10-flowered. Anthers opening by an apical pore. Stem simple or

Mmanenemwbcatiolade AS —2 3ycti Lome se: ac: 20.0 04 foo oe es ade sda een eda ee ctiesee acaadens 19

. Inflorescence 10—30-flowered. Anthers opening by a longitudinal slit in upper

half or along the length of anther. Stem simple. Leaf blade 18—35 cm long ..... 20

Inflorescence 4—7(—10)-flowered. Anthers drip-shaped to deltoid, purple. [India,

China, Myanmar, Lao P.D.R., Vietnam, Thailand] .................... Porandra ramosa

. Inflorescence 7—10-flowered. Anthers subglobose to oblong, white. [Sumatera,

VISE, IMGT ST el oe oo ee ce ee oe 19. A. rostrata

Leaf blade on lower surface glabrous or hairy only between the veins. Sepals 4.5—

6.5 mm long. Capsule 6.5—8.5 mm long. [Borneo] ...................... 12. A. leiocarpa

. Leaf blade on lower surface hairy on the veins. Sepals 7-10 mm long. Capsule

pm long alimdiay Bameladeshy) .22c.c.icdsccesdsens svancssssnstscsetacscccenees 9. A. hookeri

Submarginal hairs 0.1—0.4(—1) mm long, if longer than 0.5 mm, then sheath with

a ring of 0.5—1 mm long hairs at node, anthers 2—2.5 mm long and capsule valves

ANUS KO Nl tat OA OME UL spare. shee aoe cescouctteedeseaderonchotaes dcdivesovsectecessseltedoubohicas D2

. Submarginal hairs (0.7—)1.2—2.5 mm long. Sheath glabrous or with hairs evenly

scattered or in lines. Anthers 1.0—1.8 mm long. Capsule valves free ................. 24

Sepals 4.5—6.5 mm long, glabrous or rarely sparsely 0.2—0.3 mm long hairy.

CA DSTINS EIB STROUD acs) si ee 12. A. leiocarpa

. Sepals 7-10.5 mm long, glabrous or 0.2—1.5 mm long hairy. Capsule 0.5—2.0 mm

GNA MITCH gy gers tamer eee MANN leas eT Ie dss 5, Ose da ion son cculoneee sie eaad be enddseceelec 23

Sheath with a few lines of hairs and/or a ring of hairs at the node, hairs 0.5—3 mm

long. Capsule 11.5—16 mm long, dark green to pink or reddish, bristle-hairy with

hairs 1.5—2 mm long. [Myanmar, Cambodia, Lao P.D.R., Thailand, Sumatera] ....

. Sheath with scattered hairs 0.1—0.2 mm long. Capsule 7-9 mm long, white to

pink, rather soft-hairy with hairs 0.5—1 mm long. [Peninsular Malaysia-Cameron

elit CoeeagnGH S| Pree emer) < SN, CRETE FF Jed se dteseeecasetialedss 17. A. parvifructa

. Sepals 4.5-7 mm long. Inflorescence 4—15-flowered. [SE Asia mainland,

SURAT Gas ONES ON pe ee et eee. me, EE A wend dal aceteos Ph abe besde seats bathe 5)

. Sepals (7—)8.5—11.5 mm long. Inflorescence 10—40-flowered. [Borneo] ........ 26

. Sheath glabrous or rarely with | or a few lines of 0.5—2 mm long hairs. Anthers

white to pale yellow. Capsule 0.7—1.5 mm long hairy, valves free. [Peninsular

Malaysia. singapore, Sumatera, BORMEO)|| -2....c2..c.csccceqenesn-cssnwacsascnne 5. A. gracilis

60 Gard. Bull. Singapore 64(1) 2012

b. Sheath with a single line of 1-3 mm long hairs. Anthers at least at margin violet

to purple. Capsule glabrous or 0.3—0.7 mm long hairy, valves fused in basal 1/2 to

2/3. [China, Laos, Vietnam, Thailand, Sumatera] .................... Porandra scandens

26a. Leaf blade abruptly narrowed into a 3.5—9.5 cm long pseudopetiole, veins very

distinct on upper surface. Capsule 3-lobed at apex when dried, lobes 1—1.5 mm

longsstyle remnant absemts....cceeere. eee eee 13. A. lobata

. Leaf blade gradually to rather abruptly narrowed with pseudopetiole indistinct or

up to 5 cm long, veins not distinct on upper surface. Capsule not lobed at apex

when dried, style remnant 0.7—1 mm long and persistent .......... 18. A. pedicellata

ion

1. Amischotolype barbarossa Duist. sp. nov. (Fig. 1 & 11A—B)

Amischotolype mollissimae et A. hispidae similis, internodiis pilis 0.5—1 mm longis,

vaginis capsulisque pilis aurantiacis ad rubris vel raro fulvis, laminis praecipue infra

pilis submarginalibus differt. TY PUS: Ridley 2950, Peninsular Malaysia, Perak, Larut

Hills (holo SING).

Amischotolype mollissima auct., sensu Turner, Gard. Bull. Sing. 47 (1997) 515; non

(Blume) Hasskarl., Flora 46 (1863) 392.

Forrestia mollis auct., sensu C.B.Clarke in A.Candolle & C.Candolle, Monogr. Phan.

3 (1881) 236, p.p. [A.C.Maingay 1712 p.p.; see note under A. griffithii|; Hooker f., Fl.

Brit. India 6 (1894) 383, p.p.; Ridley, Mat. Fl. Malay. Penins. 2 (1907) 124; J. Malayan

Branch Roy. Asiat. Soc. 1 (1923) 103; Fl. Mal. Pen. 4 (1924) 360; non Hasskarl, Flora

47 (1864) 628, nom. superfl. (= A. mollissima (Blume) Hassk.); non Ridley, J. Straits

Branch Roy. Asiat. Soc. 33 (1900) 170 (= A. gracilis).

Forrestia mollissima auct., sensu Briickner in Engler & Prantl, Nat. Pflanzenfam. ed.

2 (1930) 169; non (Blume) Koord.

Stem ascending from short creeping rhizome; erect part 50-200 cm long, simple;

internodes (moderately to) densely (0.5—)1 mm long hairy. Sheath 10-17 mm diam.,

with at least a few frontal lines of 1.5—4 mm long, yellow to orange or brown hairs, in

between glabrous to moderately 0.1—0.5 mm long hairy, mouth ciliate. Leaf blade 21—

45 x (S—)7—13 cm, 2.9-4(—S) times as long as wide, base gradually to rather abruptly

narrowed into a winged pseudopetiole 34 cm long and 1.5—3 mm wide; lower surface

moderately to densely 0.5—1 mm long yellow- to reddish brown-hairy, upper surface

glabrous or rarely sparsely to moderately 0.1—0.2 mm long hairy; submarginal hairs on

lower surface, rarely also on margin and upper surface, 1-2 mm long, dense, yellow to

yellow-brown. Inflorescence on erect stem and at knee, sessile, 2—3.5 cm diam., (very)

dense with branches obscure, 15—30-flowered. Pedicel 0.5 mm long. Sepals 7.5—12.5

Amischotolype in Asia 61

NWA ' [vi

oe ff AW Hy Wa NW Sf 4

6/4 ¥, SAR ANT Why \\ VW

WALSMIT SACHS 2012 CS ) Alin TANI Hh VY, NS SW

(pele pA A A A une ALAN AN!

Fig. 1. Amischotolype barbarossa Duist. A. Habit. B. Immature fruit with persistent sepals.

Drawing by A. Walsmit Sachs, from Kiah 31743 (L).

62 Gard. Bull. Singapore 64(1) 2012

x 2.5-3.5 mm, not or little elongating in fruit, (pale) pink, moderately to densely 0.5—

1.0 mm long orange- to dark red- or brown-hairy at least in upper half, tip hooded.

Petals 7-8 x 2 mm, much shorter than sepals, translucent (greenish) white, glabrous,

margin minutely fringed. Stamens with filaments c. 7 mm long, white, in upper part

with many 2—2.5 mm long hairs; anthers 1.5 x 0.5 mm, white, thecae opening by a

longitudinal slit. Capsule 7-10 =< 5.5—7 mm, ovoid, 2 mm shorter to 1.5 mm longer

than sepals, white to red or lilac, entirely or upper half (rarely only at apex) moderately

to rather densely covered in 1—1.5 mm long orange- to red-brown hairs; valves free;

apex slightly depressed, lobes absent or up to 0.3 mm long; style remnant 0.5—1.5 x

0.3—0.5 (base) mm, finally deciduous. Seeds 2 per locule, aril red.

Distribution. Thailand (Peninsular Thailand: Nakhon Si Thammarat), Peninsular

Malaysia (Kedah, Perak, Pahang, Negeri Sembilan, Malacca, Johor), Sumatera (Riau,

Jambi).

Ecology, In primary dipterocarp forest, in lighter shaded areas, on dry sandy soil or

moist to wet or swampy places. Altitude: 30—250(—800) m asl.

Notes. 1. This species was never recognised as different from 4. mollissima (Blume)

Hassk. Differences from A. mollissima and A. hispida, both unknown from the Asian

mainland, include the 0.5—1 mm long hairs on the internodes, the orange to red (or

rarely yellow) hairs on the sheaths, sepals, and capsules, and the submarginal hairs

predominantly on the lower surface of the leaf blade.

2. Ridley (1907) gave an acurate morphological description of the species, but

unfortunately did not realise the difference from F. mollis Hassk., a synonym for A.

mollissima. He cited only one specimen which is chosen here as the type specimen.

The epithet barbarossa means red beard and refers to the red hairs on the sheaths,

sepals and capsules.

Specimens examined: THAILAND. Peninsular Thailand: Nakhon Si Thammarat, Khao

Luang, May 1968, C.F. van Beusekom & C. Phengkhlai 920 (L).

PENINSULAR MALAYSIA. Kedah: Gunung Baling, Nov 1929, G.A. Best (SF) 21261

(SING); Baling, Ulu Legong, Kampung Keda, Water Catchment Area, Apr 2006, K. /min et

al. FRI 50644 (KEP). Perak: Maxwell’s Hill, Mar 1892, H.N. Ridley 2950 (SING); Temango,

Jul 1904, H.N. Ridley 13472 (SING); Belum, Sg Sara above OrangAsli village of Ciong, Sept

1993, 1.M. Turner & J.W.H. Yong 111 (SINU). Pahang: Temerluh, Krau GR, Bkt Rengit, Nov

1999, S. Damahuri et al. FRI 45316 (KEP); Temerluh, Kemasul FR, Feb 1936, Ja ’amat 40872

(KEP); near Kuala Teku, Jul 1936, Kiah SF 31743 (K, L, SING); Teku river, Gunong Tahan,

Jun 1922, Md.Haniff & Md.Nur SF 8092 (SING); Bukit Belar, Ulu Sungai Tembeling, Mar

1968, Md.Shah 1635 (K, KEP, L, SING); Jenka FR, May 1964, M.E.D. Poore 1420 (KLU);

Jerantut, Gng Aais FR, Sg Lurut, Jul 2004, ¥ ¥ Sam FRI 49056 (KEP); Bentung, Lintang FR,

Jun 1934, CF Symington 40515 (KEP); Jenka FR, Oct 1963, E.A. Turnau 908 (K, KLU).

Negeri Sembilan: Pasoh FR, Nov 1976, K. Jong s.n. (KLU); Pasoh F.R. 12 1979, R. Kiew

824 (SING); J.V. LaFrankie 2024 (KEP, L), Oct 1987, J.V. LaFrankie 2530 (KEP); 1989, JV.

LaFrankie 4284 (KEP); Jelebu, Pasoh Forest Reserve, Sept 1977, Mat Asri FRI 25737 (K, KEP,

Amischotolype in Asia 63

L); Petasik Jelebu, Triang Forest Reserve, Jul 1927, E.A. Strouts 13024 (KEP, SING). Malacca:

A.C. Maingay 1712, p.p. (L); base of Ophir, Jun 1982, H.N. Ridley s.n. (SING). Johor: north

of Gunung Belumut, May 1923, R.E. Holttum SF 10617 (SING); Gunong Lombak, Nov 1922,

R.E. Holttum SF 9394 (K, SING); Path to Bukit Tangga Tujoh via Kota Tinggi, Aug 1970,

Mad.Shah et al. 2085 (L, SING); Genuang, Mar 1915, H.N. Ridley s.n. (BM); Seduok, Aug

1908, H.N. Ridley 14374 (BM, SING); Sungei Juasseh, Labis, Jun 1970, Samsuri Ahmad 268

(L, SING); Mersing, Endau-Rompin, Ulu Endau, Sept 1985, R. Kiew s.n. (KEP).

SUMATERA. Riau: Tigapulu Mountains, Nov 1988, JS. Burley et al. 1318 (K, KEP, L,

SING); Anak Talang, Oct 1939, P. Buwalda 7129 (L). Jambi: Sri Mengopeh, Sept 1938, 7: W.

Rutten-Kooistra 32 (U); P.T.IFA concession, Pasir mayang, Feb 1982, MJ. Vreeken-Buijs 4 (L).

2. Amischotolype divaricata Duist. sp. nov. (Fig. 2)

Amischotolype hookeri similis, internodiis glabris, laminis supra pilis submarginalibus

plerumque brevioribus, infra in venibus non conlectis, filamentis (fere) glabris differt.

TYPUS: Larsen et al. 31476, August 1972, Thailand, NE, Chatyapum Province, near

Chulaphorn Dam, 16°30”N, 101°50”E (holo L; iso KLU).

Stem ascending or decumbent from rhizome, scrambling or winding; erect part 50—

300(—600) cm long, simple; internodes glabrous. Sheath 7-12 mm diam., at the node

with a few or ring of 0.5—1 mm long hairs and/or few lines of 1-3 mm long hairs,

mouth ciliate. Leaf blade 19-36 = 3.5—6.5 cm, 4.2—6.6 times as long as wide, lower

surface (light) green, tinged with purple or striped brown, base gradually narrowed

into a winged pseudopetiole 0.5—1.5 cm long and 2 mm wide; lower surface glabrous

to moderately 0.2—0.3 mm long hairy or rarely with 2 mm long hairs along midvein,

upper surface glabrous or rarely sparsely to moderately 0.2 mm long hairy; submarginal

hairs on upper surface, rarely also on margin, 0.2—0.5(—1) mm long, dense, yellow.

Inflorescence on erect stem, sessile, 24.5 cm diam., dense with branches obscure,

(10—)15—20(—30)-many-flowered. Pedicel absent. Sepals 7—-10.5 x 3-4.5 mm, not

elongating in fruit, light green with pink or violet tip (in flower) to greenish red or

purple (in fruit), glabrous or very sparsely 0.2—1.5 mm long ciliate keel and margins,

tip hooded. Petals c. 8 x 2.5 mm, distinctly shorter than sepals, white, glabrous, margin

minutely fringed. Stamens with filaments 8.5—10 mm long, white, glabrous or upper 3

mm with | mm long hairs; anthers (1.5—)2—2.5 x 0.6—1 mm, white, thecae opening by

a longitudinal slit (rarely in upper 2/3 only). Capsule 11.5—16 x 6—8 mm, ovoid, longer

than sepals by 5-9 mm, dark green to pink or reddish, in upper half to 2/3 sparsely

to rather densely 1.5—2 mm long bristle-hairy; valves fused for 1/4 to 1/2 of length;

apex acute to obtuse and slightly depressed, lobes absent; style remnant 0.5 < 0.3 mm,

persistent. Seeds 2 per locule, aril orange.

Distribution. Myanmar (Tenasserim), Lao P.D.R. (Xiangkhouang, Vientiane,

Khammouan, Champasak), Vietnam (Central Highlands: Kon Tum; South Central

Coast: Khanh Hoa; Southeast: Dong Nai), Cambodia (Pursat, Kampong Speu,

Kampot), Thailand (N: Chiang Mai, Phayao; E: Chaiyaphum; C: Nakhon Nayok; SE:

Chantaburi, Trat; Peninsular Thailand: Songkhla), Sumatera (N).

64 Gard. Bull. Singapore 64(1) 2012

gap

WALSMIT SACHS 2017

Fig. 2. Amischotolype divaricata Duist. A. Habit. B. Fruit with persistent sepals. Drawing by

A. Walsmit Sachs, from K. Larsen et al. 31476 (L).

Amischotolype in Asia 65

Ecology. Rather open primary, disturbed or secondary evergreen (bamboo) or seasonal

forest, on damp or moist to marshy places near streams, shaded to almost in full sun;

on white sand with large rocks, sandstone, granite or shale bedrock. Altitude: 50-1225

m asl.

Notes. The epithet refers to the scrambling habit of the plant. The species is similar

to and until now (mis)identified as 4. hookeri, but differs in the glabrous internodes,

the generally shorter submarginal hairs on the upper surface (and margin) of the leaf

blade, the hairs on the lower surface of the leaf blade not concentrated on the veins,

and the (almost) glabrous filaments. Hara (1966), Hong (1974, 1997) and Hong &

DeFilipps (2000) mention A. hookeri for the Chinese province of Yunnan. I have not

seen their specimens and their descriptions are wide enough to include both species.

Study of the specimens from Yunnan is needed to elucidate which of the two taxa are

present.

Specimens examined: MYANMAR. Tenasserim: Tavoy distr, Oct 1961, Keenan et al. 1604

(E);

LAO P.D.R. Xiangkhouang: Xieng-Khouang, 1926, M. Delacour s.n. (P). Vientiane: Phon

Kheng, Apr 1953, JE. Vidal 2301 (P); route du Phou Khao Khouay, Oct 1965, J.E. Vidal 4041

(P); Phou Khao Khouay, Oct 1971, JE. Vidal 5547 (P). Khammouan: Boneng-Thakhek, Apr

1950, J.E. Vidal 1274 (P); Nyalak, Oct 2005, MF Newman et al. LAO 530 (L); Tha Phabat

Distr., Path to Tad Leuk Waterfall, May 2006, MF Newman et al. LAO 1539 (L, P); Nakai

Oudom Souk, Apr 2005, B. Svengsuksa et al. BT 32 (L). Champasak: Boloven plateau, entre

Noug Bok Kao et Phong, Oct 1928, E. Poilane 15837 (P).

VIETNAM. Central Highlands: Kon Tum, nr Kontum, entre Ta Bai et Giang Lo, Sep 1930,

E. Poilane 18284 (P). South Central Coast: Khanh Hoa, sur le bas Song Cao et le Song Cay,

pres Nhatrang, Jul 1921, F Evrard 571 (P); Southeast: Dong Nai, Bien Hao, ?Gioray, Jul 1919,

E. Poilane 186 (P); Dong nai, nord de Blao Kil, Haut Donai, Oct 1931, E. Poilane 19869 (P):

Dong nai, Blao, du Haut-Donai, Apr 1933, E. Poilane 22419 (P). Cochinchine, Déon bu., Apr

1866, L. Pierre s.n. (P); Cochinchine, Camchoy, L. Pierre 1] (P); Cochinchine, 1862-1866,

Thorel s.n. (several in P); MeKong, Sep 1874, Harmand 1920 (P):; Dalat, Oct 1924, F Evrard

1467 (P).

CAMBODIA. Pursat: 1876, Harmand 3366 (P). Kampong Speu: Kompong Spen, Sep 1870,

L. Pierre s.n. (several in P). Kampot: Phnom Bokor, Dec 1965, M.A. Martin 815 (P). Mt.Kam

Chay, Poporkville waterfall, Mar 1970, B.C. Stone 9302 (KLU).

COCHINCHINE. 1862-66, Thorel 1139 (P).

THAILAND. N: Chiang Mai, Doi Sutep, east side, Pah Laht, Jul 1987, J.F Maxwell 87-729

(L); Phayao, Doi Luang NP, E-side, Jahm Bah Tawng Falls, May 1997, O. Petrmitr 15 (L). E:

Chaiyaphum, nr Chulaphorn Dam, Aug 1972, K. Larsen et al. 31476 (KLU, L). C: Nakhon

Nayok, Muang, Kow Yai NP, summit ridge E-side, Kow Kieo, Jul 2003, J.F. Maxwell 03-160

(L). SE: Chantaburi, western slopes of Kao Sabab, Feb 1935, G. Seidenfaden 2825 (SING):

Chantaburi, foot of Khao Soi Dao, Nov 1969, C.F. van Beusekom & T. T. Smitinand 2064 (E,

L); Trat, Klawng Yai, 7. Smitinand 269 (E); Trat, Hauy raeng, Dong tapit,7. Smitinand 1378

(E). Peninsular Thailand: Songkhla, Na Tawee, Jul 1928, G. Rahil 81] (L). Sapi foot, Apr

1922, Put 798 (L); Chantaboon?, 1936?, A. Vesterdal 7-P (SING).

SUMATERA. Sumatera Utara: ENE Sibajak, Oct 1928, J.A. Loerzing 14103 (L).

66 Gard. Bull. Singapore 64(1) 2012

3. Amischotolype dolichandra Duist. sp. nov. (Fig. 3)

Amischotolype glabrata similissima, sepalis 7.5—9 mm longis, filamentis glabris,

antheris 3 mm longis, capsulae valvibus liberis differt. TYPUS: Koelz 26448, 1950,

India, Manipur, Karong, 1050 m alt., in deep forest, scandent (holo L).

Stem rhizome unknown, scrambling; erect part c. 90 cm long, with c. 40 cm long

aerial (stilt) roots, simple; internodes glabrous. Sheath 8-11 mm diam., with 0.3—0.5

mm long hairs in upper third to half of the frontal part, mouth ciliate. Leaf blade

18-19 x 2.7-4.1 cm, 4.6—-6.7 times as long as wide, base gradually narrowed into an

indistinct winged pseudopetiole to 0.5 cm long and 2 mm wide; lower surface glabrous

or moderately 0.5 mm long hairy, upper surface glabrous; submarginal hairs on upper

surface, 0.8—1.5 mm long, dense, yellow. Inflorescence on erect stem, sessile, 1.6—1.7

cm diam., dense with branches obscure, 5—10-flowered. Pedicel absent. Sepals 7.5—9 x

3-3.5 mm, possibly slightly elongating in fruit, pink, glabrous or sparsely 0.3 mm long

ciliate margins, tip hooded. Petals c. 9.5 x 3 mm, slightly longer than sepals, colour

unknown, glabrous, margin minutely fringed. Stamens with filaments c. 9 mm long,

colour unknown, glabrous; anthers 3 x 0.7 mm, colour unknown, thecae opening by a

longitudinal slit. Capsule 6.5 x 5 mm, obovoid, shorter than sepals by c. 2 mm, pink,

moderately 1.5 mm long hairy; valves free; apex obtuse, lobes absent; style remnant

1.2 x 0.4 mm, persistent. Seeds 2 per locule, aril colour unknown.

Distribution. India (NE: Meghalaya, Mizoram, Manipur).

Ecology. Deep forest. Altitude: 1050-1200 m asl.

Notes. The epithet refers to the strikingly long anthers. The species most closely

resembles A. glabrata which has a much wider distribution, and has longer sepals,

stamens with hairy filaments and much shorter anthers, and capsules with basally

fused valves.

Specimens examined: INDIA. NE. Meghalaya: Khasi Hills, Cherapunjee, Jun 1952, Thakur

Rup Chand 5921 (L) & 5944 (L). Mizoram: Tengawl, Lushai Hills, Nov 1927, N.E. Parry 286

(K). Manipur: Karong, Oct 1950, W.N. Koelz 26448 (L).

4. Amischotolype glabrata Hassk. (Fig. 9A & 11C—D)

Amischotolype glabrata Hassk., Flora 46 (1863) 392; Coode et al., Checklist Fl. Pl.

Gymn. Brun. Dar. (1996) 352; Turner, Gard. Bull. Sing. 47 (1997) 515; Beaman &

Beaman, Plants of Mount Kinabalu, 3 (1998) 109. — Campelia glabrata auct., sensu

Hasskarl in Miquel, Pl. Jungh. (1852/53) 154; Miquel, Fl. Ned. Ind. 3 (1855) 546; non

Kunth, Enum. pl. 4 (1843) 109 (see note 2). — Forrestia glabrata (Hassk.) Hassk.,

Flora 47 (1864) 630; Commelin. Ind. (1870) 94; Clarke, Commelyn. Cyrtandr. Bengal.

(1874) 62, tab. XLII; Clarke in A.Candolle & C.Candolle, Monogr. Phan. 3 (1881)

Amischotolype in Asia 67

Fig. 3. Amischotolype dolichandra Duist. A. Habit. B. Fruit with persistent sepals. Drawing by

A. Walsmit Sachs, from W.N. Koelz 26448 (L).

68 Gard. Bull. Singapore 64(1) 2012

238; Hooker f., Fl. Brit. India 6 (1894) 384; Koorders, Excursionsfl. Java 1 (1911) 282;

Hallier f., Nova Guinea 8 (1913) 906; Ridley, Fl. Malay. Penins. 4 (1924) 360; Ridley,

J. Bot. 63 (1925) 124; Briickner in Engler & Prantl, Nat. Pflanzenfam. ed. 2 (1930)

169. — Forrestia mollissima (Blume) Koord. forma glabrata (Hassk.) Backer, Handb.

Fl. Java I, 3 (1924) 33, p.p.; Backer & Bakhuizen van den Brink, FI. Java 3 (1968)

15, p.p. — Forrestia mollissima (Blume) Koord. var. glabrata (Hassk.) Subba Rao

& Kumari, Bull. Bot. Surv. India 12 (1970) 209 (nom. inval.: Vienna Code Art. 33.4,

no full reference to basionym). LECTOTYPE (designated here): Junghuhn s.n., Java:

Ungaran prope Medini (L: L0041658). Other syntypes: Junghuhn s.n., Java: Merapi

(L), Gede et Tangkuban prau (L).

Campelia marginata var. B Blume, Enum. Pl. Javae (1827) 7; Kunth, Enum. PI. 4

(1843) 109; Miquel, Fl. Ned. Ind. 3 (1855) 547 (‘var foliis minoribus margine subtus

tantum ciliolatis’). TYPE: Zippelius s.n., Java (L).

Forrestia glabrata (Hassk.) Hassk. forma minor Hassk., Flora 47 (1864) 631;

Hasskarl, Commelin. Ind. (1870) 94. — Campelia n. sp., Moritz, Syst. Verz. (1845-

46) 93; Zollinger, Syst. Verz. (1854) 65. SYNTYPES: Nagler s.n., 1858, Java (photo

B: Barcode: B 10 0296343 / Imageld: 302936); Blume s.n., s.d., Java, Preanger ad

ped. mnt. Pangerangoh prop. Tjipanwar (L); Zollinger 305, Java, Tjapus (photo B:

Barcode: B 10 0296344 / Imageld: 302937); Zollinger s.n., 23 Oct 1858, Java, Idjen

(le):

Forrestia chinensis N.E.Br. in Forbes & Hemsley, J. Linn. Soc., Bot. 36 (1903) 158;

Walker, Fl. Okinawa & S Ryukyu Isls. (1976) 298. — Amischotolype chinensis [E.H.

Walker ex Hatusima, Fl. Ryukyus (1971) 771, nom. inval.: see note 4] ex K.-S. Hsu,

Fl. Taiwan 5 (1978) 158, t. 1305. SYNTYPES: Oldham 578/1 (P); Henry 149, 1618A,

5670 (all BM or K, n.v.); Ford 256 (BM or K, n.v.).

Tupistra cavaleriei H.Lév., Repert. Spec. Nov. Regni Veg. 10 (1912) 372; McKean,

Notes Roy. Bot. Gard. Edinburgh 45, 1 (1988) 153, in syn. A. hispida. SYNTYPES:

Jul. Cavalerie 2565, 3517, China, Guizhou (“Kweichow’), Lo-Fou (photo: E).

Forrestia sumatrensis Ridl., J. Malayan Branch Roy. Asiat. Soc. 1 (1923) 102. TYPE:

Ridley s.n., 8 Feb. 1921, Sumatera, Berastagi Woods (holo K). (See note 5).

Forrestia hispida auct., sensu C.B.Clarke in A.Candolle & C.Candolle, Monogr. Phan.

3 (1881) 236, p.p. — Amischotolvpe hispida auct., sensu Hong, Acta Phytotax. Sin.

12, 4 (1974) 461; McKean, Notes Roy. Bot. Gard. Edinburgh 45, | (1988) 153; Hong

in Wu, FI. Reipubl. Pop. Sin. 13, 3 (1997) 71; Hong & DeFilipps, Fl. China 24 (2000)

22; Wang, Chen & Peng, Fl. Taiwan, 2nd ed., 5 (2000) 153; Yang, Liu & Lin, Man. FI.

Taiwan 5 (2001) 56, pl. 103; Yuan and Yang, Bot. J. Linn. Soc. 152 (2006) 400; non

A. Rich. (See note 3).

Amischotolype in Asia 69

Stem ascending from short rhizome or basally decumbent and with long roots,

scrambling; erect part 50-200 cm long, sometimes with aerial (stilt) roots, simple;

internodes glabrous or moderately to densely 0.1—0.3 mm long hairy. Sheath (6—)8—12

mm diam., glabrous or at its front moderately 0.3—0.5(—1) mm long hairy or with one

or a few lines of 1-3 mm long hairs, mouth ciliate or rarely glabrescent. Leaf blade

15-28 x 3.2—6.5(—8.3) cm, 3+4.5(—6.3) times as long as wide, smooth, lower surface

sometimes red, base gradually to abruptly narrowed into a winged pseudopetiole up

to 2.5 cm long and 1 mm wide; lower surface glabrous or rarely sparsely to densely

0.1—0.5(—2) mm long hairy or 2 mm long hairy on midvein, upper surface glabrous

or rarely very sparsely to moderately 1 mm long hairy; submarginal hairs on lower

surface, rarely (also) on margin or upper surface, 0.2—0.5(—1) mm long, rather dense,

colourless to yellowish. Inflorescence on erect stem, sessile, 1.5—4 cm diam., dense

with branches obscure, (5—)10—20(—25)-flowered. Pedicel absent. Sepals (9—)10—14.5

x (1.5—)2-3(-4.5) mm (in fruit), (dark) green, pinkish purple or dark violet, sometimes

base white, probably not elongating in fruit, sparsely to moderately 0.2—1.5 mm long

colourless- or orange-ciliate keel or rarely glabrous, tip hooded. Petals c. 8 x 3 mm, as

long as sepals, white, glabrous, margin entire. Stamens with filaments c. 11 mm long,

white, upper half with 1-3 mm long hairs; anthers 0.6—1 « 0.5—1 mm, deltoid, white,

thecae opening by a longitudinal slit. Capsule 5-8 = (3.5—)4.5—6.5 mm, obovoid,

shorter than sepals by (1—)2.5—5(—6) mm, at first white and later turning to green and

finally red, upper half sparsely to moderately 1—2.5 mm long hairy or rarely glabrous;

valves fused for 1/3 tol/2 of length; apex obtuse, lobes absent; style remnant 1-3 x

0.3—0.5 mm, persistent. Seeds 2 per locule, aril orange.

Chromosomes. 2n = 36 (Yuan & Yang 2006).

Distribution. Pakistan (E: Sind), India (Sikkim, Meghalaya, West Bengal, Assam),

China (Guizhou, Yunnan, Guangxi Zhuang, Hainan), Taiwan, Japan (Ryukyu Isl.),

Vietnam (Tonkin, Lang Giu, Ninh Binh), Myanmar (Tenasserim), Thailand (N: Chiang

Mai, Chiang Rai, Lampang), Peninsular Malaysia (Kedah (Bukit Telim), Pahang

(Cameron Highlands), Selangor (Ulu Langat)), Sumatera (Toba, Talang, Karo plateau,

Sago, Singalang, Asahan), Borneo (Sabah: West Coast), Java (West: Preanger, Salak,

Bandoeng, Batavia, Bogor, Talaga-bodas, G.Patoeka, G.Tiloe; Central: Ungarang;

East: Mount Idjen, Besoeki, Pasoeroean), Nusa Tenggara: Lombok (Rindjani-vulcano).

Ecology. Primary or disturbed (mountain, oak-laurel) evergreen, mixed or deciduous

(monsoon) forest, often near stream, on dry to moist sandy or loamy soil, limestone,

granite or andesite breccia, shaded. Altitude: (75—)210—1850 m asl.

Notes. 1. This is the most widespread species of the genus. It is the only species

occurring in Pakistan, China, Taiwan, and Japan.

2. As Hasskarl (1863) noted, Campelia glabrata as used in Asian literature

is not based on Kunth (1843). The inflorescence in Kunth’s original material in B

70 Gard. Bull. Singapore 64(1) 2012

(Barcode: B 10 0184765 / Imageld: 245454), Kunth s.n. (Febr. 1833, Hortus Berol.),

has a long peduncle with two leaf-like bracts at the apex enclosing the flowers, whereas

the Asian material has very condensed inflorescences. The genus Campelia is restricted

to the American continent, and is now united with 7radescantia (Hunt 1986).

3. This species has often been referred to as A. hispida, a confusion that started

with Hooker (1864) who mentioned the species is distributed from New Guinea to

India. Since that time onward, the name hispida has been in use for continental Asian

material (e.g. Hong & DeFilips 2000), most of which should be identified as A. glabrata.

Both A. hispida and A. glabrata have capsules distinctly shorter than the sepals with

valves fused in the basal part. Differences include position and length of submarginal

hairs (with slight overlap), presence of pedicel, length of hairs on capsule (with some

overlap), length of anthers, shape of capsule apex, and length of style remnant. The

species co-occur only on Borneo. However, 4. hispida has been found only in E and

SE Kalimantan, whereas 4. glabrata has only been found in the mountainous areas of

Sabah.

4. Amischotolype chinensis E.H. Walker ex Hatusima is invalid because

reference to the basionym was incomplete and indirect (McNeill et al. 2006, Art. 33.4).

Hsu (1978) was the first to validate the name. Of the type material I have only seen

Oldham 3578/1, which has submarginal hairs on the upper surface, a state that is only

rarely seen in other specimens of A. glabrata. However, Brown’s original description

in Forbes & Hemsley (1903) agrees with the present delimitation of A. glabrata. The

illustration in Hsu (1978) clearly shows the persistent style remains; the fruit, however,

is depicted with four valves instead of three.

5. Ridley (1923) did not cite specimens when he described F’. sumatrensis,

he just mentioned that it was found in Berastagi woods (Sumatera). There is only one

collection of Ridley (K) that fits his description with matching locality, which should

therefore be the holotype. Ridley (1923) mentioned that the stamens are “apparently

quite glabrous”. Unfortunately, these could not be checked from the photo. The

filaments of other Sumatran material (Bartlett 66/2) are not glabrous, but compared

to material from other regions they are much less hairy and with shorter hairs. In the

present treatment this is regarded as variation within the species.

Specimens examined: PAKISTAN. E: Chittagong Hill Tracts, between Langlagklopara and

Kairwa, Jan 1965, M.S. Khan 952 (K).

INDIA. Sikkim: J.D. Hooker f. & T: Thompson s.n. (K, L); E. Himalaya, Rongbe, Jul 1913, G.H.

Cave s.n. (E). Meghalaya: Khasia, Shillong, Oct 1872, C.B. Clarke 19048 (BM); Shillong, Aug

1886, C.B. Clarke, 44390A (BM). West Bengal: Darjeeling, Rishap, Aug 1870, C.B. Clarke

12518 (K, BM); Darjeeling, Oomong, Oct 1870, C.B. Clarke 13645 (K); Darjeeling, Jun 1893,

Cowan s.n. (E). Assam: Garo Hills, Tura Mountain, 1929, NV.E. Parry 809 (K).

CHINA. Guizhou: Lo-Fou, Aug 1909, J. Cavalerie 2565 (E) & 3517 (P, E); 1936, S.W. Teng

90984 (L). Yunnan: A. Henry 10884 (E) & 12204 (E) & 13377 (E). Guangxi Zhuang: Na

Pao Kou, San Chiang Hsien, Sep 1933, 4.N. Steward & H.C. Cheo 110] (P). Hainan: Fung

Sheung Tsuen, Ngai distr, Aug 1932, Lau 427 (P, E); Chim Fung Mt., Feb 1935, Lau 5422 (E);

Yaichow, Aug 1933, Liang 62393 (E).

TAIWAN. Mt Arisan, Aug 1914, S. Kawagoe s.n. (L); Tamsui, 0, 1864, R. Oldham 578/1 (P);

Amischotolype in Asia 71

Taipei Hsien, Pingling Hsiang, Taiyuku, Nov 1992, Liu et al. 2] (E).

JAPAN. Ryukyu Isl.: Okinawa, Kunigami, Taminato Ugan, Jul 1951, E.H. Walker et al. 6130

(L); Okinawa, Motobu Peninsula, foot of Mt Awa, Dec 1953, E.H. Walker 7607 (L); Nantou

Co., Sun-Moon-lake, Aug 1972, C.-S. Kuoh 38824 (L).

VIETNAM. ?Dalat, ?Adran, Nov 1953, N. Schmid s.n. (P). Tonkin: Mont-Baui(?), Jun 1886,

B. Balansa, B. 4100 (P); Tien-yen, Ho Yung Shan and vicinity, 1940, WT. Tsang 30748 (E, P,

SING). Lang Giu: Than Mai, 1916, Eberhardt 3277 (P). Ninh Binh: Cuc Phuong NP, Nov

1998, D.D. Soejarto et al. 10380 (L); Cuc Phuong NP, site CP71, Jul 1999, N.M. Cuong 300

(L); Nho Quan District, Cuc Phuong NP, Dec 1999, D.D. Soejarto et al. 11318 (L).

MYANMAR. 1885, R.H. Beddome 8001 (BM). Tenasserim: Tavoy distr, Aug 1961, J. Keenan

et al. 715 (E); Aug 1961, J. Keenan et al. 1048 (E).

THAILAND. N: Chiang Mai: Doi Angka, E. slope nr rivulet, Oct 1927, H.B.G. Garrett 483

(L); Doi Chiang Dao, E foothills, Pa Blawng Cave area, Oct 1989, JF Maxwell 89-1287 (L).

Chiang Rai: Ban Pa Deuah, Bah Sahng subdistrict, Feb 1991, JF Maxwell et al. 98 (L).

Lampang: Jae Sawn NP, Wahng Die, Pah Ngahm (Nahn Kaht) cave, Jan 1996, .F Maxwell

96-76 (L).

PENINSULAR MALAYSIA. Kedah: Bukit Telim, Nov 1908, H.N. Ridley 13892 (BM, K,

SING). Pahang: Cameron Highlands, Boh plantation, Apr 1937, Md. Nur SF 32944 (SING).

Selangor: Ulu Langat, 1912, C.B. Kloss s.n. (K).

SUMATERA. P.W. Korthals s.n. (L). Sumatera Utara: Berastagi Woods, Feb 1921, H.N.

Ridley s.n. (K); Brastagi, Dec 1921, E.M. Burkill 30 (SING); N of Berastagi, Deleng Sing.,

Feb 1927, H.H. Bartlett 6612 (L); plateau above Berastagi, Jan 1929, J.A. Loerzing 15073 (L);

Asahan, East Coast, Aek Si Tamboerak, Oct 1936, Rahmat Si Boeea 10647 (L); vic. Taloen na

Oeli, nr headwaters Aek Mandosi, Nov 1936, Rahmat Si Boeea 10952 (L). Sumatera Barat:

G.Singalang, May 1918, H.4.B. Buennemeyer 2588 (L); Mt.Talang, Jul 1953, J. van Borssum

Waalkes 2796 (L, SING). N slope of Mt Sago, Pajakumbuh, Air Kesimbultan, Mar 1955,

W.J.C. Meijer 3654 (L).

BORNEO. Sabah: West Coast: Kinabalu, Menteki River, /.H. Beaman et al. 10785 (K): Kota

Belud, Kinabalu, Golf Course Site, Oct 1983, J.H. Beaman et al. 7205 (L); Kota Belud, Mount

Kinabalu, Penibukan, Dahobang falls, Jan 1933, J. & M.S. Clemens 30681 (L, BM); Kota

Belud, Mt Kinabalu, Penataran basin, Jul 1933, 7. & M.S. Clemens 34200 (L, BM): Kota Belud,

Kinabalu, Penataran River, Jul 1933, 7. & M.S. Clemens 34272 (BM); Ranau, Kundasan, Nov

1957, Kidinan Cox 951 (L); Ranau, S slope Kinabalu, Penosok plateau, Sungai Mesilau, Jul

1963, H.P. Fuchs et al. 21111 (L); Ranau, Mt.Kinabalu, Mesilau camp, Jan 1964, WL. Chew

& E.J.H. Corner 4001 (SING, L); Ranau, Kinabalu, Pinosok plateau, 1964, MZ E.D. Poore

3863 (L); Kota Belud, Kinabalu NP, along road between HQ and Tenumpok, Jan 1969, S.

Kokawa & M. Hotta 3095 (L); Ranau, Mesilau trail few trails from road, Sep 1972, /.L. Chow

SAN 76433 (L, SING); Ranau, Kinabalu, West Mesilau River, Nov 1983, /.H. Beaman et al.

7452 (L, K); Ranau, Kinabalu, West Mesilau River, Feb 1984, J.H. Beaman et al. 8637 (L);

Penampang, Penampang: Crocker Range, Mar 1984, J.H. Beaman et al. 8849 (L); Tambunan,

km 63 KK-TambunanRd, Jul 1984, Amin et al. SAN 60351 (SAN, L); Ranau, Kundasang,

Melisau, Mar 1985, A. Zuraidah bt 0027/7195 (SAN); Ranau, S.Bambangan, Sep 1988, Amin

et al. SAN 123564 (SAN); Tambunan, Kawasan simpanan Rafflesia, Jul 1989, Norhayati

Sandapali 03 (SAN); Ranau, Kampong Bundu Tuhan, Sep 1992,S. Doinis 24 (KEP); Kota

Belud, Mt.Kinabalu, halfway along Liwagu Trail, Feb 1995, K.M. Cameron s.n. (K); Kota

Belud, Kinabalu Park, Mountain Garden, Jul 2005, H. Duistermaat 392 (L, SING).

JAVA. Kuhl & v.Hasselt s.n. (L). Jakarta: Pasir Limoes (Boerangrang), Jul 1920, R.C.

Bakhuizen van den Brink sr. 4367 (L). Jawa Barat: A. Zippelius s.n. (L); Salak, A. Zippelius

72 Gard. Bull. Singapore 64(1) 2012

s.n.(2269a) (L); Gedeh, FW. Junghuhn s.n. (L); Pangerangoh, nr Tjipanjawar, C.L. Blume s.n.

(L); Tjibodas, J.G. Boerlage s.n. (L); tussen Tjibodas en Tjikeureum, S. Bloembergen 86 (L);

G.Patoeka, P W. Korthals s.n. (L); Pangerango, supra Tjibodas, Apr 1894, V. Schiffner 1692 (L);

Tjibodas, Oct 1896, S.H. Koorders 26031B (K); Tjigenteng, Jan 1897, S.H. Koorders 26329B

(L); Tjibodas, Oct 1898, S.H. Koorders 31912B (L); Tjibodas, 1905, R.H.C.C. Scheffer s.n.

(L); WZW van Buitenzorg, Nirmala, Dec 1913, C.S. Backer 10673 (L); Tjiboden, 1915, Sapiin

450 (SING); Tjibodas, 1915, Sapiin 2037B (L, U); G.Beser, May 1917, WE: Winckel, WF: 28

(L); Tjadas, Malang-Tjidadap, Tjibeber, Jun 1917, R.C. Bakhuizen van den Brink sr. 1935 (L);

Talaga-bodas, PW. Korthals s.n. (L); G.Djaja, Mar 1920, H.J. Lam 166 (L); G. Perbakti, Sep

1922, R.C. Bakhuizen van den Brink Jr. 1729 (U); G.Tiloe, Rioeng goenoeng, Jan 1927, J.H.

Coert 575 (L); Tjibodas, Dec 1932, WU.C. Kooper s.n. (U); Gedeh, Tjibodas, Apr 1932, A.

Kloos 38 (L); voetpad Tjibodas-Tjibeurum, Jan 1939, C. Holstvoogd, C. 210 (L); Tjisaroea,

Feb 1950, S.J. van Ooststroom 12843 (L). Jawa Tengah: Ungarang, prope Medini, FW.

Junghuhn s.n. (L); Djati-Kalangan, Waitz (845’) (L). Jawa Timur: Banyuwangi: Mount Idjen,

Oct 1858, H. Zollinger s.n. (L); Besoeki, Banjoelinoe-Kalibendo, Jul 1916, S.H. Koorders &

A. Koorders-Schumacher 40490B (L); Pasuruan, N...kadjadja, May 1923, J. Dorgelo 513 (L);

Ranse Pani near Sendoero, Feb 1934, J.H. Coert 1120 (L).

NUSA TENGGARA. Lombok: Rindjani-vulcano, SO-seite, Sappittal, Pussu, Jun 1909, J.

Elbert 1799 (L); Rindjani-Vulkangebirge, Sapittal, S-side Pussuk, Jun 1909, J. Elbert 1839

(L, SING).

5. Amischotolype gracilis (Ridl.) 1.M.Turner (Fig. 9B, 11E—-F & 12A—D)

Amischotolype gracilis (Ridl.) 1.M. Turner, Gard. Bull. Sing. 45 (1993) 53, 47 (1997)

515. Keng et al., Conc. Fl. Sing. Volume II: Monocotyledons (1998) 23. — Forrestia

gracilis Ridl., J. Straits Branch Roy. Asiat. Soc. 41 (1904) 37; Mat. Fl. Malay. Penins.

2 (1907) 123; Fl. Malay. Penins. 4 (1924) 359; Briickner in Engler & Prantl, Nat.

Pflanzenfam. ed. 2 (1930) 169; Keng, Gard. Bull. Sing. 40 (1987) 124. LECTOTYPE

(designated here): Ridley 238], Pulau Tijau (SING). Other syntypes: Cantley s Coll.

(Curtis s.n., Kwala Lumpor; A/vins Y.H. 2020, Gunong Bumbon) (SING); Ridley

2382, 6320, 10512 (all SING); Ridley 4130 (n.v.), Walker 155 (BM, n.v.).

Forrestia hispida auct., sensu Hooker f., Bot. Mag. 90 (1864) t. 5425; non A.Rich.

Forrestia mollis auct., sensu C.B.Clarke in A.Candolle & C.Candolle, Monogr. Phan.

3 (1881) 236, p.p. (Walker 155); Hooker f., Fl. Brit. Ind. 6 (1894) 383, p.p.; Ridley,

J. Straits Branch Roy. Asiat. Soc. 33 (1900) 170; non Hasskarl, Flora 47 (1864) 628,

nom. superfl. (= A. mollissima (Blume) Hassk.); non Ridley, Mat. Fl. Malay. Penins.

2 (1907) 124; J. Malayan Branch Roy. Asiat. Soc. 1 (1923) 103; Fl. Malay. Penins. 4

(1924) 360 (= A. barbarossa Duist. sp. nov.).

Stem ascending from short creeping rhizome; erect part 90—250(—300) cm long, often

with short or long stilt roots, simple; internodes glabrous. Sheath (4—)6—11(—13) mm

diam., glabrous or rarely with | or a few lines of 0.5—2 mm long hairs, mouth ciliate or

rarely glabrous. Leaf blade (16—)20-—35(-45) = (3.5—)4-7(-8.7) cm, (3.3—)4—6(-7.3)

Amischotolype in Asia 73

times as long as wide, smooth, lower surface green to dark purple, upper surface green

or variegated with silver-white (see note 3), base gradually to abruptly narrowed into

a winged pseudopetiole up to 5.5 cm long and 1—3 mm wide; lower surface glabrous

or sparsely to densely 0.1—1 mm long hairy, upper surface glabrous or rarely sparsely

0.1—0.3(—0.5) mm long hairy; submarginal hairs on upper surface, (0.7—)1.5—2.5 mm

long, (rather) dense, (yellow-)orange to red-brown. Inflorescence on erect stem,

sessile, 1.5—3.5 cm diam., (rather lax to very) dense with branches obscure or up to

3(—6) mm long, 5—15-flowered. Pedicel 0—1 mm long. Sepals 4.5—7 x 2.5-4.5 mm, not

elongating in fruit, white to green in flower, green to purplish in fruit, margins and keel

or entirely sparsely to moderately (0.1—)0.2—0.5 mm long hairy or rarely glabrous, tip

hooded. Petals 4—7 x 2-3 mm, not to slightly longer than sepals, white, glabrous or

with few 0.2 mm long hairs at midvein near apex, margin entire or minutely fringed

at apex. Stamens with filaments 3—7 mm long, white to pale yellow, in upper 0.5—3

mm with many 1.5—2 mm long, white hairs; anthers 1.0—1.8 x 0.5—0.7 mm, white to

pale yellow, thecae opening by an apical pore or a longitudinal slit 0.3-1.8 mm long.

Capsule 7—11(—14) x 4.5—8 mm, obovoid, white to purplish red, purple or rarely green,

longer than sepals by (2.5—)4—6(—7) mm, upper 1/3 partially to entirely rather sparse

to moderately 0.7—1.5 mm long white- to yellow-hairy (hairs longest at apex); valves

free; apex slightly depressed, lobes absent: style remnant 0.8 < 0.2—0.4 mm, finally

deciduous. Seeds 2 per locule (rarely 1 and double-sized, or | abortive), aril orange,

rarely containing 2 seeds.

Distribution. Peninsular Malaysia (Kedah, Kelantan, Terengganu, Perak, Pahang,

Selangor, Negeri Sembilan, Malacca, Johor), Singapore, Sumatera (Aceh, Anambas

Natuna Islands, Jambi, Palembang), Borneo (Sarawak: Kuching; see note 2).

Ecology. In rather dry to wet or swampy, primary or recently logged-over, lowland,

mixed, dipterocarp or montane rain forests, rarely in kerangas or peatswamp forest,

in dry streambeds and on slopes; partly shaded; on granite, clay, and alluvial soil.

Altitude: 0—1350(—1500) m asl.

Notes. 1. As Ridley (1904) noted, this is the most common species of the genus in

Peninsular Malaysia and Singapore. Earlier authors included it in Forrestia mollis (but

see also A. barbarossa), an illegitimate name as Hasskarl (1864) based it on Campelia

mollissima Blume. It is most obviously different from 4. gracilis in having capsules

much shorter than the sepals and with apical lobes.

2. The species is known from Borneo from only two collections, both from

Sarawak. Another specimen is close to this species but deviates from the above

description in having 2 mm long hairs on the upper surface of the leaf blade (S [S.7.Lai]

54495: SAR n.v., KEP, L).

3. Material from disturbed forest along Jalan Gombak, Selangor, Peninsular

Malaysia, has silver-white variegated leaf blades (Duistermaat 353; Fig. 1E). The

variegation slowly disappeared when the plant was grown in the greenhouse.

74 Gard. Bull. Singapore 64(1) 2012

Specimens examined: PENINSULAR MALAYSIA. Kedah: Sungai Batang, Mar 1938, Sow

34656 (KEP); Baling, Lata Mengkung, Dec 1990, Khairuddin Hj. Itam 8 (KEP); Baling, Ulu

Legong, Kampung Keda, Water Catchment Area, Apr 2006, K. Imin et al. FRI 50645 (KEP,

L); Baling, G. Inas FR, Bkt Iboi, Apr 2008, 1 Kamarul Hisham et al. FRI 59170 (KEP).

Kelantan: Chaning Woods, Feb 1917, H.N. Ridley s.n. (K); Ulu Sungei Ketil, Mar 1972,

Mad.Shah 2588 (KEP, SING); Bukit Berangkat Reserve, via Kpg. Perak Bertam, Feb 1973,

Ma.Shah & Md.Ali 2862 (KEP, SING); Sg. Lebir, Eastern bank opp. Jentah, Apr 1976, B.C.

Stone 12549 (KEP, KLU); Jeram Pasu, May 1982, B.C. Stone et al. 15211 (KLU); Merapoh,

near base camp, Jun 1995, Ong 654 (KLU); Merapoh, bird trail forest near base camp, Jun

1995, Ong 572 (KLU). Terengganu: Rantau Panjang F.R., Jun 1921, HL. Hume 7589

(SING); Kemaman, Bukit Kojang, Nov 1935, E.J.H. Corner s.n. (SING, K); Kemaman, Ulu

Bendong, Nov 1935, E.J.H. Corner s.n. (SING); Ulu Brang, Jul 1937, L.M.J. Moysey & Kiah

SF 33726 (SING); Dungun, Jambu Bongkok F.R., Aug 1964, Kassim M. s.n. (KLU); Dungun,

May 1968, Soepadmo KLU 9151 (KLU); Bukit Yong, Jerteh, Mar 1975, Md. Shah et al. 3542

(KEP, KLU, SING); Dungun, Bukit Bauk Forest Reserve, Feb 1998, Tham S.M. 56 (KEP);

Besut, Bukit Bintang, Hilltop around Telekom station, Sep 2006, Yao TL. et al. FRI 53122

(KEP, L). Perak: Telok Sera, Mar 1896, H.N. Ridley s.n. (SING); Bukit Kapayang, 1904, H.N.

Ridley s.n. (SING); Ulu Temango, Jul 1904, H.N. Ridley 14376 (BM, SING); Taiping Hill,

Jan 1923, MR. Henderson 10413 (SING); Taiping Hill, Feb 1923, MR. Henderson SF 10451

(SING); Kuala Kangsar, Bubu Forest Reserve, Apr 1933, C.F) Symington 29943 (KEP); Batu

Kurau, Apr 1968, Sidek bin Kiah S 331 (SING); Maxwell Hill, Nov 1980, Keng Wee et al. 58

(SINU); Kuala Kangsar, Kledang Saiong Forest, 1987, S. Anthony 769 (KEP). Pahang: Pulau

Tiyau, Pahang River, 1891, H.N. Ridley 2381 (SING); Tahan River, 1891, H.N. Ridley 2382

(SING); Titi Bangor, Temerloh, Mar 1923, MR. Henderson SF 10631 (SING); Fraser’s Hill,

Aug 1923, Md.Nur SF 11039 (SING); Pulau Tioman, Gunong Rokam, May 1927, Md. Nur SF

18773 (SING); base Gunong Senyum, Aug 1929, MR. Henderson SF 22335 (SING); Raub,

Bilut FR, Apr 1932, Osman 28480 (KEP); Bentung, Lentang FR, Jun 1935, Symington 40519

(KEP); Raub, Bukit Kajang, Jun 1935, C.F! Symington 40535 (KEP); Fraser’s Hill, Ring Road,

Apr 1955, J.W. Purseglove 4173 (K, L, SING); Jengka For. Res., Nov 1963, E.A. Turnau 909

(KLU); Pekan, Bukit Chini FR, Oct 1968, PS. Bray FRI 11652 (KEP); Ulu Sungai Sepia,

near Kuala Aur, Jul 1970, Md.Shah & Md.Noor 1905 (KEP, L, SING); Aug 1970, Rao et al.

s.n. (SING); Ginting Highlands Road, May 1972, B.C. Stone 10761 (KLU); Taman Negara,

Kuala Trenggan, path to salt lick, Aug 1972, B.C. Stone 10839 (KLU); Fraser’s Hill, path to

the Lodge, Aug 1972, Md.Shah 2731 (SING); Kuala Tahan, Taman Negara, 1973, B.C. Stone

11545 (KLU); Fraser’s Hill, Nov 1973, Keng et al. 34 (SINU); nr basecamp at Ulu Sungei

Kinchin, Rompin-Endau, Aug 1996, 1.M. Turner 96-103 (SING); Temerluh, Krau Game

Reserve, Oct 1999, Chua L.S.L. FRI 40873 (KEP); Nenasi FR, Nov 2005, Shamsul Khamis

et al. N-42 (KEP); Temerloh, Bukit Rengit, Krau wr, Rentis Tualang, Mar 2006, K. Jmin et al.

FRI 50603 (KEP); Rompin, P.Tioman, Kpg.Paya, Mar 2007, K. Jmin et al. FRI 50949 (KEP,

L). Selangor: KL, Univ. Malaya Bot. Garden Greenhouse, Khatijah 103 (KLU); Bukit Kuda,

Jun 1889, H.N. Ridley s.n. (SING); Kwala Lumpor, Feb 189x, C. Curtis s.n. (SING); Batu Tiga,

Feb 1904, H.N. Ridley 11878 (SING); Klang Gates, Aug 1908, H.N. Ridley s.n. (SING); Ulu

Langat, Feb 1912, C.B. Kloss s.n. (BM); Kwala Lumpur, Welds Hill, Mar 1915, H.N. Ridley

s.n. (BM, K, SING); Selangor, Ulu Gombak, Sep 1921, H.L. Hume 8625 (SING); Selangor,

Ulu Gombak, Oct 1921, H.L. Hume 9152 (SING); Ginting Simpah, Oct 1921, H.L. Hume 9328

(SING); Petaling, Sungai Buloh, Feb 1928, E.J Strugnell 13939 (KEP); Hulu Langat, Sungai

Lalang Forest Reserve, Mar 1930, C.F’ Symington 22671 (KEP); Ginting Simpah, Oct 1937,

Md.Nur SF 34263 (K, SING); Gombak, Ulu Gombak, Apr 1958, Abdullah b.1. & A.H. Willard

Amischotolype in Asia 75

304 (KEP, S INU); Ulu Gombak, Jun 1962, B.L. Burtt & P.J.B. Woods 1632 (E); Genting

Simpah rd. 22nd mile, Ulu Gombak FR, May 1965, B.C. Stone 5751 (KLU); Ulu Gombak

16th mile, UM Field Studies Centre, Nov 1965, Soh Kim-Gai 11] (KLU); Templer Park forest

reserve, Jun 1971, Teo L.E. 406 (L, SING); Ulu Lanlat, Sungei Serai water shed, Mar 1976,

Lee D.W. 23338 (KLU); Petaling, Air Hitam Forest Reserve, Sep 1978, S. Anthony 208 (KEP):

Klang Gates Ridge, Jan 1982, R. Kiew 1180 (SING); Teluk Selangor, Dec 1990, Khairuddin

Hj. Itam 7 (KEP); JIn Gombak km48/47(Bentong), FRIM-examination plot, Jun 2005, H.

Duistermaat 348 (L, SING); N. of Kuala Lumpur, Jin Gombak km 36/37(KL), Jun 2005, H.

Duistermaat 355 (L, SING); Gombak, Ulu Gombak FR, Miles 15, Mar 2006, K. Imin et al.

FRI 50567 (KEP). Negeri Sembilan: Gunong Bumbon, Jun 1885, YH. Alvins 2020 (SING);

Gunong Tampin, Nov 1922, R.E. Holttum SF 9536 (K, SING); Gunong Angsi, N.W.side, Nov

1923, Md.Nur SF 11587 (SING); Gunong Telepak Burok, Seremban, Mar 1977, J.F. Maxwell

77-155 (L, SINU); Pasoh FR, Aug 1987, J.V. LaFrankie 2457 (KEP); Pasoh FR, Sep 1987,

JV. LaFrankie 2483 (KEP); Pasoh FR, 1989, J.V. LaFrankie 4253 (KEP); Gunung Angsi, Jun

2005, H. Duistermaat 343 (SING) & 344 (L, SING); Seremban, Berembun FR, Air Pantai, Sg

Batang Benar, Apr 2008, M.A. Mohd Hairul et al. FRI 60040 (KEP); Jelebu, Berembun FR,

G. Telapak Buruk, Apr 2008, K. Jmin et al. FRI 63020 (KEP); Jelebu, Berumbun FR, Jeram

Toi, forest trail to ridge, Apr 2008, Yao T.-L. et al. FRI 65283 (KEP). Malacca: Sungai Buloh,

Oct 1899, H.N. Ridley 10512 (SING). Johor: Tanjong Bunga, 1894, H.N. Ridley 6320 (BM,

SING); between K.Tinggi and Rombon, K.T., Sep 1929, Z. Teruya 1027 (SING); S.Rhu Reba,

Jason Bay, Jun 1934, E.J.H. Corner s.n. (SING, K); Lengor F.R. Mersing, Jul 1957, Whitmore

406 (SING); Kuala Sedili, New Road, Jun 1959, Kadim Md. Noor 163 (SING); Lombong Batu,

Mersing, Sep 1966, Jumali & E.H. Heaslett s.n. (SINU); Kluang, path to Gunung Blumut, Sep

1970, Md.Shah & Sanusi 2123 (SING); Gunung Panti top westside, Oct 1972, Samsuri Ahmad

Shukor 784 (SING); Mt.Ophir, Apr 1972, R.B. Phillips 1637 (KLU); Endau State Park, Pantai

Burung, Apr 1998, Damahuri S. & Mustapa D. s.n. (KEP); Gunung Panti, top, May 2005, H.

Duistermaat 336 (L, SING); Segamat, Ulu Sg.Segamat, trail up the ridges, Aug 2007, Chew

M.Y. & Kueh H.L. FRI 55597 (L); Kota Tinggi, Panti FR, G.Panti, Jul 2008, Yao T.-L. et al. FRI

65379 (KEP).

SINGAPORE. MacRitchie Res, Sinclair 4931 (E); Tampines, 189x, J.S. Goodenough s.n.

(SING); Reservoir Woods, 1893, H.N. Ridley s.n. (SING); Pulau Ubin, 1893, H.N. Ridley 4810

(SING); Chan Chu Kang, 1894, H.N. Ridley s.n. (SING); Reservoir woods, 1906, H.N. Ridley

s.n. (SING); Bukit Timah Forest Reserve, Sep 1959, Md.Shah 747 (SING); Bukit Timah Nature

Reserve, Sep 1965, Hess/lett 3468 (SINU); Bukit Timah Nature Reserve, May 1967, Jumali s.n.

(SINU); SW of MacRitchie Reservoir, May 1993, W.S. Chee et al. 2158 (KEP); Bukit Timah,

Fern Valley, Jun 1998, L.M.J. Chen 245 (SING) & 246 (SING); Bukit Timah Nature Reserve,

2000, C.F. Teo D.H.L. 25 (SINU); Nee Soon, Jun 2003, Samsuri et al. 27 (SING); Bukit Timah

Nature Reserve, Tiup Tiup path, Mar 2004, Samsuri et al.41 (SING); SBG, rainforest off Liana

Rd close to Cluny Rd, Apr 2005, H. Duistermaat 328 (L, SING); Bukit Timah, Fern Valley,

Apr 2005, H. Duistermaat 331 (L, SING) & 333 (SING); Bukit Timah NR, Rangas Path, May

2005, H. Duistermaat 340 (SING); MacRitchie Reservoir, patch behind Gardenia Rd, May

2005, Gwee A.T. et al. 2005-142 (SING).

SUMATERA. Aceh: Jul 1939, A.H. Batten Pooll s.n. (SING); W.van Kroeng Loeas, O van

Troemon, Aug 1941, Asdat 4/ (L); Bukit Lawang-Bohorok-Langkat, Feb 1973, R. Soedarsono

262 (L, SING) & 307 (L); Gng Leuser NP, G.Mamas, Feb 1975, W.J.J.O. de Wilde & B.E.E.

de Wilde-Duyfjes 14789 (L); Gng Leuser NP, Alas river valley, Jul 1979, W.J.J.O. de Wilde &

B.E.E. de Wilde-Duyfjes 18480(vel 1) (L); Gng Leuser NR, Sikundur FR, Aug 1979, W..J.O.

de Wilde & B.E.E. de Wilde-Duyfjes 19293 (L); Kloet NR, along Krung (=river) Lembang, Jul

76 Gard. Bull. Singapore 64(1) 2012

1985, WJ.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 19740 (L); Kloet NR, along Krung (=river)

Lembang, Jul 1985, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 19904 (L). Sumatera Barat:

Afd.4 Kota S.Koeriman, May 1918, H.4.B. Buennemeyer 3312 (SING, L); Afd. Agam Brani,

May 1918, H.4.B. Buennemeyer 3355 (L). Kepulauan Riau: Anambas / Natuna Islands,

Terempak, Siantan, Apr 1928, .R. Henderson SF 20254 (SING); Siantan Isl., E of Terompa,

Apr 1928, C.G.G.J. van Steenis 918 (L, SING, U). Jambi: S Kejasung Tabin, Jambi Sunake,

Mar 1980, ¥. Laumonier 804 (L); P.T. IFA Concession, Pasir Mayang, Feb 1982, M.J. Vreeken-

Buijs. 45 (L). Sumatera Selatan: Palembang, Rawas, Jun 1917, J.M. Dumas 1598 (L).

BORNEO. Sarawak: Kuching: Lundu, Jalan Lundu, Othman et al. S 61977 (E); Kuching,

Lundu, Sentunggan, Dec 1995, Lai S.T et al. S 54495 (cf; KEP, L).

6. Amischotolype griffithii (C.B.Clarke) I.M.Turner (Fig. 9C)

Amischotolype griffithii (C.B.Clarke) I.M.Turner, Novon 6 (1996) 221; Coode et al.,

Checklist Fl. Pl. Gymn. Brun. Dar. (1996) 352 (see note 2); Turner, Gard. Bull. Sing.

47 (1997) 515. — Forrestia griffithii C.B.Clarke in A.DC. & C.DC., Monogr. Phan.

3 (1881) 236; Hooker f., Fl. Brit. India 6 (1894) 383; Ridley, Mat. Fl. Malay. Penins.

2 (1907) 123; Fl. Malay. Penins. 4 (1924) 360; Briickner in Engler & Prantl, Nat.

Pflanzenfam. ed. 2 (1930) 169; non Cowley & Furness, Kew Bull. 52 (1997) 469;

non Beaman & Beaman, The Plants of Mount Kinabalu 3 (1998) 109 (= see note

2). LECTOTYPE (designated by Turner (1996)): Maingay 1712, p.p. (K), Peninsular

Malaysia, Malacca.

Stem ascending from rhizome; erect part c. 100 cm long, simple; internodes glabrous.

Sheath 7-13 mm diam., glabrous, mouth (sparsely) ciliate. Leaf blade (18—)25-35 x

4.5—7.5 cm, 3.3—5.6 times as long as wide, base gradually to rather abruptly narrowed

into a winged pseudopetiole 1—3 cm long and c. 2 mm wide; lower surface sparsely

to densely (0.5—)1—2 mm long hairy, upper surface sparsely to moderately 1-2 mm

long yellow-hairy; submarginal hairs on upper surface, (0.8—)1—3 mm long, (rather)

dense, yellow to reddish gold or rarely red. Inflorescence on erect stem, sessile,

2—3.5 cm diam., (rather) dense with branches obscure, 10—20-flowered. Pedicel 1—1.5

mm long. Sepals 4.5—6.5(—7.5) x 2.5 mm, not elongating in fruit, (white or) red to

purplish, sparsely to moderately (rarely densely) 0.5 mm long ciliate margins and keel

or hairy all over, tip hooded. Petals 44.5 = 1.5 mm, slightly shorter than sepals, white,

glabrous, margin entire. Stamens with filaments c. 8 mm long, colour unknown, upper

half with 1.5 mm long hairs; anthers 1.1 x 0.7 mm, yellow, thecae opening by an apical

pore. Capsule (7—)8—12 = 5—8.5 mm, pyriform, longer than sepals by (1.5—)3.5—6 mm,

white to purple, upper half only to entirely sparsely to moderately (0.7—)1(—2) mm

long hairy; valves free; apex depressed, lobes absent; style remnant 0.5—0.7 x 0.2—0.3

mm, deciduous. Seeds 2 per locule, aril colour unknown.

Distribution. Peninsular Malaysia (Penang, Kelantan, Perak, Pahang, Negeri Sembilan,

Malacca, Johor), Sumatera (Riau).

Amischotolype in Asia WI

Ecology. Moist to wet or swampy primary or regenerating (dipterocarp) forest, by

streams or riversides; shaded or rarely unshaded. Altitude: 60—1310 m asl.

Notes. 1. Maingay 1712 is a mixture (see Clarke, 1881), with one specimen (L) which

is A. barbarossa, and another (K; only seen from a photo) which is the present species.

Two specimens at SING [Holmberg S56 and Ridley s.n. (Perhantian Tinggi)] identified

by Ridley as F. griffithii are in accordance with the above description.

2. According to Ridley (1907) this species is endemic to Peninsular Malaysia.

Today it is also known from Sumatera where it has been found only twice. It has been

mentioned for Borneo (Coode et al. 1996, Beaman & Beaman 1998), but specimens

available for study are all 4. hirsuta Hallier f., a closely similar species with hairy

internodes and hairy sheaths (J. & M.S.Clemens 26039, 26476; Cowley 19), and A.

lobata (Shea & Aban SAN 76938), a species with distinctly lobed capsules.

Specimens examined: PENINSULAR MALAYSIA. Penang: 189x, H.N. Ridley s.n. (SING).

Kelantan: Sungei Merkeh, Bukit Baka, Machang, Md.Shah & Ahmad Shukor 3186 (KEP,

KLU, SING); Sungei Sat FR, Bukit Baka, base, May 1982, B.C. Stone in Chin S.C. 3200

(KLU); Kuala Krai, Gua Bogo, track to G.Stong and camp Ba, Feb 2007, Yao TL. et al. FRI

55864 (KEP, L). Perak: Kinta, Ulu Sungai Kinta, Jun 1938, Sow 47255 (KEP). Pahang: Tras,

May 1903, A.D. Machado s.n. (SING); Fraser’s Hill, upon the Selangor border, Sep 1922,

HM. Burkill & R.E. Holttum 8869 (SING); Gunung Lesong FR, May 1939, Fontian Forester

31645 (KEP); Fraser’s Hill, near Selangor Residency, Aug 1959, Md.Shah & Md.Noor 731

(SING); Fraser’s Hill, Aug 1969, B.C. Stone 8753 (KLU); Temerluh, Krau GR, Kuala Lompat,

Sep 1981, R.W. Kiew 1049 (KEP). Negeri Sembilan: Perhantian Tinggi, 18xx, H.N. Ridley s.n.

(SING); Gunong Tampin, Sep 1913, H.C. Robinson s.n. (K); Tebong FR, Dec 1959, M.E_D.

Poore 134 (KLU); Port Dickson, Sungai Menyala Forest, Oct 1978, R.W. Kiew 675 (KEP):

S.Menyala F.R., Oct 1978, R.W. Kiew 688 (SING). Malacca: Ager Punnus (Ayer Panas), W.

Griffith 5485, p.p. (K); A.C. Maingay 1712, p.p. (K); 1891, D.F:A. Hervey s.n. A (SING) & s.n.

B (SING); Selandor, Jan 1885, Y.H. Alvins 473 (SING); Sep 1885, ¥.H. Alvins 2102 (SING);

Bukit Singei, Mar 1886, YH. Alvins s.n. (SING); base of Mount Ophir, Aug 1888, R.W. Hullett

s.n. (844) (K); Jasin, Apr 1890, Anon. 499 (SING); Bukit Sidanan (Sadanan), Aug 1891, P./.

Holmberg 856 (BM, K, SING). Johor: Gunung Pulai, 1892, Mar. s.n. (SING); Gunung Pulai,

Dec 1904, H.N. Ridley s.n. (SING); Bukit Semanau, Aug 1915, H.M. Burkill SF 1337 (SING);

192x, YH. Alvins 1587 (SING); Endau Rompin, nr Jasin River just S of junction Endau River,

Mar 1992, JS. Klackenberg & R. Lundin 587 (L).

SUMATERA. Riau: Muara Padjanki, Apr 1939, P. Buwalda 6485 (L); Tigapulu Mts, Bukit

Karampal area, Nov 1988, J.S. Burley et al. 1237 (L, SING).

7. Amischotolype hirsuta (Hallier f.) Duist. comb. nov. (Fig. 9D & 13A)

Forrestia hirsuta Hallier f., Beih. Bot. Centralbl. 34 (1916) 47; Merrill, J. Straits

Branch Roy. Asiat. Soc. (1921) 113; Briickner in Engler & Prantl, Nat. Pflanzenfam.

ed. 2 (1930) 169. TYPE: Winkler 2617, 28 Jun 1908, SE Borneo, Hayup (holo B, photo

(Barcode: B 10 0296342 / Image Id: 302935); iso L).

78 Gard. Bull. Singapore 64(1) 2012

Amischotolype griffithii auct., sensu Coode et al., Checklist Fl. Pl. Gymn. Brun. Dar.

(1996) 352: Beaman & Beaman, Plants Mount Kinabalu 3 (1998) 109; non C.B.Clarke

(= A. griffithii).

Amischotolype mollissima auct., sensu Cowley & Furness, Kew Bull. 52 (1997) 469;

Beaman & Beaman, Plants Mount Kinabalu 3 (1998) 110; non Blume (= A. mollissima).

Forrestia mollis auct., sensu C.B.Clarke in A.Candolle & C.Candolle, Monogr. Phan.

3 (1881) 236, p.p. [Beccari 827]. non Hasskarl, Flora 47 (1864) 628, nom.superfl. (=

A. mollissima (Blume) Hassk.); non Ridley, Mat. Fl. Malay. Penins. 2 (1907) 124; J.

Malayan Branch Roy. Asiat. Soc. 1 (1923) 103; Fl. Malay. Penins. 4 (1924) 360 (= 4.

barbarossa Duist. sp. nov.).

Stem ascending from short rhizome; erect part 50-200 cm long, sometimes scrambling,

simple; internodes sparsely to densely (0.3—)1—2.5 mm long hairy or rarely glabrous.

Sheath (8—)10—15(—22) mm diam., with at least several frontal lines of hairs and a

single line of hairs at its back, hairs 24(—7) mm long, yellow, mouth ciliate and

sometimes glabrescent. Leaf blade 20—35(-42) = (4—)5—7(-11) cm, 3.5—6.0(—7.4)

times as long as wide, lower surface red tinged or not, base (very) gradually to rather

abruptly narrowed into a winged pseudopetiole up to 3 cm long and c. 3 mm wide;

lower surface sparsely to densely (0.5—)1-4 mm long white- to yellow-hairy, upper

surface sparsely (mostly near midvein) to rather densely (1.5—)2—4 mm long yellow-

hairy; submarginal hairs on upper surface, 1.5—2.5 mm long, (rather) dense, yellow

or rarely orange. Inflorescence on erect stem or rarely at knee, sessile, 1.8-4.5 cm

diam., dense with branches obscure, 15—40-flowered. Pedicel 1.5—3 mm long. Sepals

(4.5—)6—10 « 24(—5) mm, not elongating in fruit, green (in flower) to lilac or purple

(in fruit), glabrous to margins and keels 0.5—1 mm long ciliate, tip hooded. Petals c.

8 x 2.5 mm, as long as sepals, pale pink, glabrous, margin minutely fringed. Stamens

filament c. 8 mm long, colour unknown, upper | mm with (0.5—)2 mm long hairs;

anthers 1—1.3 = 0.8—1 mm, colour unknown, thecae opening by a longitudinal slit.

Capsule 6—9 « 3.5—6 mm, ovoid to pyriform, equalling or exceeding sepals by up to 4

mm, (pale) green or pinkish to reddish- or deep purple, narrowed part of apex to upper

1/3 1-2 mm long, yellow bristle-hairy; valves (almost) free; apex depressed, lobes

absent; style remnant c. 0.5 x 0.4 mm, finally deciduous or rarely persistent. Seeds 2

per locule, aril orange.

Distribution. Sumatera (Padang), Borneo (Brunei; Sabah: West Coast, Interior,

Sandakan, Tawau; Sarawak: Kuching, Samarahan, Sri Aman, Sibu, Bintulu, Miri;

Kalimantan: Dusun, Central, West, East, SE).

Ecology. Primary, disturbed, logged-over or secondary, lowland, mixed dipterocarp,

swamp, alluvial, beach or montane forest, on flat land, river bank, hill side or ridge,

on damp, rich dark or yellow sandy soil, shale or sandstone. Altitude: 6—1400 m asl.

Amischotolype in Asia 79

Notes. |. Hallier (1916) described Forrestia hirsuta as being very close to Ridley’s F.

irritans but with hairy sheaths and with smaller and almost glabrous inflorescences.

However, it shows more resemblance to A. griffithii with which it has been confused

(see there, note 2), and which is almost confined to Peninsular Malaysia.

2. This species is almost restricted to Borneo, with only two specimens known

from Sumatera (Beccari 827, Padang, near Ajer mantjur; Korthals s.n., s.loc.).

Specimens examined: SUMATERA. P.W. Korthals s.n.A (L). Sumatera Barat: Padang, Abita,

nr Ajer mantjur, Aug 1878, O. Beccari 827 (K, L).

BORNEO. Brunei: Belait: Labi, Rampayoh, J. Cowley 157 (K); Belait, Labi, Rampayoh, Jul

1993, Cowley et al. 19 (L); Belait: Labi, Mendaram, Wasai Wong Kadir, Jun 1995, Ariffin Kalat

BRUN 16792 (K). Sabah: West Coast: Kota Belud, Mt.Kinabalu, Dallas, Sep 1931, J. & MS.

Clemens 26476-26039 (BM, L): Mount Kinabalu, Penibukan, near camp, Mar 1933, J. & M.S.

Clemens 35012 (BM); Ranau, about 8 miles E of Kampung Merungin, Nov 1975, Leopold &

Saikeh SAN 82649 (SAN): Ranau, Kampong Nalumad, Aug 1996, A. Daim 566 (KEP): Interior:

Tambunan, Mnt Trusmadi, Mar 1969. H.P Nooteboom 1444 (L); Tambunan distr: Crocker

Range, Oct 1983, J.H. Beaman et al. 7272 (K, L); Sipitang, Lumaku FR, L39, Mondolong SFI

Concession, Aug 1988, Muhammad Saedi et al. 64 (SAN); Tambunan, Trusmadi, Loloposon

(Gua), Jul 2005, J.J. Vermeulen in H. Duistermaat 398 (L, SING). Sandakan: Dagat Camp,

Apr 1963, Wing J.Ah SAN 35497 (K, SAN); mile 32 Labok Rd, agric.stat, Mar 1970, H_P.

Nooteboom 1605 (L); Ulu Dusun, oil palm Res. Sta., Mar 1977, B.C. Stone 12875 (KLU);

Kinabatangan, Kampong Ulu Menanam, Sep 1978, S. Dewol & T. Kodoh SAN 89259 (KEP):

Beluran, Ulu Sg.Muanad, Nov 1984, Jrang Ak.Angi SAN 67433 (L, SAN); Beluran, Jan 1985,

Amin et al. SAN 69486 (KEP, L, SING); Kinabatangan, Tongod, Pinangah FR, Feb 1985, S.

Dewol & Mansus SAN 67483 (SAN); Kinabatangan, Ulu Sungai Melikop, Jun 1985, S. Dewol

& T. Langkap SAN 109288 (KEP, L); Lahad Datu, Ulu Segama, Danum Valley.at entrance to

west trail, Mar 1987, G. Argent et al. GA 681987 (E, SAN); Sepilok FR, Jan 1989, P. Kessler

SAN 118180 (SAN); Lahad Datu, Base Gunung Nicola, Feb 1992, A. Berhaman SAN 134457

(SAN); Kinabatangan, Lake Linumunsut, Oct 2001, Sidkan A. MB 1063 (SAN); Lahad Datu,

Danum Valley, May 2004, 7.M.A. Utteridge et al. 486 (KEP); R.I.C. Sepilok, Sep 2004, S.

Dewol et al. SAN 142184 (SAN); Tawau: Elphinstone Province, 1922-23, 4.D.E. Elmer 20507

(BM, L, SING, U): Apas. May 1948, Kadir A 2084 (KEP, SING): Kalabakan, Ulu Sungai

Kalabakan, May 1984, Fedilis & Matin SAN 103611 (L, SAN). Sarawak: Kuching: Meatang,

Aug 1884, R.W. Hullett s.n. (SING); Setapok, Kuching, Sep 1955, J. W. Purseglove & Md.Shah

P 4394 (L, SING); Lundu, Sematan Beach, Sep 1987, Abg. Mohtar et al. S 54298 (K, L); Bau,

Bukit Kho Ze San, Bau subdistrict, Dec 1994, Rantai Jawa et al. S 70126 (cf; KEP); Bau,

Gg.Lanyang, Apr 2002, S. Julia et al. SBC 2908 (SING); Samarahan: Gng Bun, Ulu Simunjan,

Sep 1975, P. Chai S 36723 (L): Serian, Bukit Rawang, Tebakang area, Apr 1983, Dvg. Awa &

I. Paie § 45586A (L); Sri Aman: Abok, Apr 1990, G. Rena et al. S 538955 (K, KEP, L): Sibu:

Kanowit, Sana, Sungei Tau, May 1956, J.W. Purseglove 3120 (L, SING): Bintulu: Oct 1985, 7.

Paie S 50161 (K, KEP, L); Miri: Marudi, Gng Mulu NP, Jul 1977, B. Lee S 38062 (L): Marudi,

G. Mulu NP, Apr 1978, R. Kiew RK 283 (KEP): Marudi, 4th Div, Route to Bt Lawi, Bario,

Aug 1985, Dyg. Awa & L. Bernard § 50535 (K, L); Marudi, Sungei Tutoh, Bt Pelamau, Apr

1997, R.M_A.P. Haegens et al. 408 (K, L). Kalimantan C: Duson-area, P W. Korthals s.n. (L);

KualaKuayan 112 28E 2 00S, Mar 1984, C. Hansen 1323 (L). Kalimantan W: Liang Gagang,

H. Hallier B2956 (L); Pontianak, Sep 1948, Moh. Enoh 355 (L); Gn Palung NP, Oct 1996, 7-G.

Laman et al. 51 (K, L). Kalimantan E: Nunukan, N. of Tarakan, Nov 1953, W. Meijer 1978

(L); Kebun raya univ.Mulawarman Samari, Aug 1974, H. Wiriadinata 298 (L); Jelini, along

80 Gard. Bull. Singapore 64(1) 2012

Sungai Belayan, NW of Tabang, Jan 1979, G. Murata et al. B 1139 (L); Tabang, West Kutai,

Dec 1980, M. Kato & H. Wiriadinata B 6098 (L); Long Bawan-Panado, Jul 1981, R. Geesink

9061 (L); P.T.ITCI, Kenangan-G.Meratus km 69, Mar 1995, P. Kessler et al. 944 (K, L); PT

KEM ENVIR. Lingau plateau, Jun 1996, Arbainsyah AA 1870 (L); Berau Forest, Swakelola,

Labanan, Oct 1996, P. Kessler et al. Berau 145 (K, L); PT Kelian, Mar 1997, P. Kessler 2091

(L); Wartono Kadri, Mar 2005, C. Bernard et al. 7362 (L). Kalimantan SE: Hayup, Jun 1908,

H. Winkler 2617 (L).

8. Amischotolype hispida (A.Rich.) D.Y.Hong (Fig. 9E & 13B)

Amischotolype hispida D.Y.Hong, Acta Pytotax. Sin. 12, 4 (1974) 461, pro nomen. —

Forrestia hispida A.Rich. in Dumont d’Urville, Voy. Astrolabe 2 (1834) 2; Miquel,

Fl. Ned. Ind. 3 (1855) 547; Hasskarl, Flora 47 (1864) 629; Commelin. Ind. (1870) 86;

C.B.Clarke in A.Candolle & C.Candolle, Monogr. Phan. 3 (1881) 236, p.p. (See note

2); Lauterbach, Bot. Jahrb. 50 (1913) 64; Merrill, J. Straits Branch Roy. Asiat. Soc.

(1921) 113, p.p. (see note 3); Briickner in Engler & Prantl, Nat. Pflanzenfam. ed. 2

(1930) 169. — Forrestia hispida A.Rich. var. typica Hallier f., Nova Guinea 8 (1913)

906; Beih. Bot. Centralbl. 34 (1916) 46, p.p. (See note 3). — Forrestia mollissima

(Blume) Koord. forma hispida (A.Rich.) Backer, Handb. FI. Java I, 3 (1924) 33.

TYPE: Lesson s.n., August 1827, crescit in sylvis Novae-Guineae (holo P, in herb.

Franquaville, 7.v.; klepto L in hb Hasskarl).

Forrestia nigricans A.Rich. ex Steud., Nomencl. Bot. ed. 2 (1840) 644. TYPE: not

indicated (see note 5).

Forrestia philippinensis Merr., Publ. Bur. Sci. Gov. Lab. 35 (1906) 5; Philipp. J. Sci.

2, 4 (1907) 266; Briickner in Engler & Prantl, Nat. Pflanzenfam. ed. 2 (1930) 169. —

Forrestia hispida A.Rich. var. glabrior Hallier f., Nova Guinea 8 (1913) 906; Beth.

Bot. Centralbl. 34 (1916) 46. — Forrestia hispida A. Rich. var. philippinensis (Merr.)

Merr., Enum. Philipp. Fl. Pl. 1 (1925) 197. TYPE: Merrill 4051 (holo PNH, n.v.; iso

K, photo).

Forrestia hispida A.Rich. forma glabrescens Lauterb., Bot. Jahrb. 50 (1913) 64;

Hallier f., Nova Guinea 8 (1913) 906 (forma dub.). SY NTYPES: Moszkowski 205, 11

July 1910, Tana, Flussufer, c. 90 m alt. (B, n.v.; L?, n.v.); Moszkowski 302, October

1910, Naumoni (B, n.v.; L?, 7.v.).

Forrestia hispida A.Rich. var. calva Hallier f., Nova Guinea 8 (1913) 906; Beth. Bot.

Centralbl. 34 (1916) 46. SYNTYPES: Atasrip 32 (L), Gjellerup 4 (L).

Stem ascending from long rhizome; erect part 45-250 cm long, often trailing or

scrambling, lower part occasionally with many stilt roots, simple; internodes glabrous

to densely 0.1—0.3 mm long hairy. Sheath (9—)11—20(—24) mm diam., with several

frontal lines of 2-4 mm long, colourless to straw-coloured hairs, rarely glabrous or

Amischotolype in Asia 81

0.1—0.2 mm long hairy, mouth ciliate or rarely glabrous. Leaf blade (19—)28-48 ~=

(4.1-)5.2-10.3 cm, (3.2—)4—6.5(—7.3) times as long as wide, lower surface purplish

tinged or not, base very gradually to rather abruptly narrowed into an indistinct winged

pseudopetiole to 8 cm long and 2 mm wide; lower surface glabrous to densely 0.1—0.5(—

1) mm long white-hairy, upper surface glabrous to densely 0.2—0.7 mm long white-

hairy and/or with 1.5—3.5 mm long yellow hairs; submarginal hairs on upper surface,

(0.5—)1—2 mm long, dense, yellow. Inflorescence on erect stem, sessile, 3—6 cm diam.,

dense with branches obscure, (10—)20—many-flowered. Pedicel 2—6 mm long. Sepals

10.5—16(—18) x 2-5 mm, unknown elongating in fruit, pink to pale purple in flower,

violet or reddish blue to brilliant (dark) red-purple in fruit, glabrous to moderately

0.2—0.5(—1) mm long ciliate margins and keel or rarely sides moderately 0.5 mm long

appressed red-hairy, tip hooded. Petals 9-11.5 x 2 mm, shorter than sepals, white or

with purple base or flushed with purple, glabrous, margin at apex minutely fringed.

Stamens with filaments 10-12 mm long, colour unknown, upper part up to 1 mm

below apex with 1-2 mm long hairs; anthers 1.2-2 x 0.5—-1 mm, white to yellow,

thecae opening by a longitudinal slit. Capsule 5-8 = 3.5—6.5 mm, ovoid, shorter than

sepals by 4-9 mm, (pale) purple or red, glabrous or apex to upper half moderately

0.1—0.5(—1) mm long soft-hairy; valves fused for 1/4 to 1/2 of length; apex slightly

depressed, lobes absent; style remnant 0.5—0.7 = 0.20.4 mm, finally deciduous. Seeds

2 per locule, aril orange.

Distribution. Philippines (Luzon, Polillo, Catanduanes, Mindoro, Negros, Samar,

Leyte, Mindanao), Borneo (Kalimantan: W, E, SE), Sulawesi (North, Central), Maluku

(Sula, Ceram, Buru, Ambon, Kai), New Guinea (Irian Jaya: Cendrawasih, Yapen

Island, NE, Central, Bomberai, S; Papua: Sepik, Madang, Morobe, Central, Gulf).

Ecology. Primary or secondary (mixed dipterocarp or 7erminalia) rain forest, on

ridges, (steep) slopes, in periodically inundated or riverine areas, gullies or (sagopalm-)

swamps, also in old Araucaria and cacao plantations; partially shaded; on clay, alluvial

or volcanic soil or limestone. Altitude: 0—1000(—1500) m asl.

Notes. 1. This species was the first described in the genus Forrestia. Hong (1974)

legally transferred it to Amischotolype, but the taxonomic use of the name has been

different (see note 2).

2. Amischotolype hispida has an East-Malesian distribution, reaching as far

north and west as the Phillipines and Borneo. It is absent from continental Asia and

it is the only species of the genus present in New Guinea. It has been mentioned for

Taiwan (Clarke 1881: Formosa: Wang et al. 2000, Yang et al. 2001) and China (Hong

1974, 1997: McKean 1988). but characters in the descriptions and all material studied

for these areas (Cavalerie, Jul. 2565, 3517; Henry 10884, 12204, 13377; Kawagoe

s.n., 8-1914; Kuoh, C.-S. 3882A; Lau 427, 5422; Liang 62393; Liu et al. 21; Teng, S.W.

90984) are referable to A. glabrata.

3. Hallier (1916) and Merrill (1921) included material of A. pedicellata in this

species (Hallier B2600). For differences see note under that species.

82 Gard. Bull. Singapore 64(1) 2012

4. This species is fairly variable with respect to indument of vegetative parts,

sepals and capsule. Hallier (1913) described three varieties, based on the amount of

indument on the sheath, occurring on New Guinea, Borneo and the Philippines. I see

no use in maintaining these entities. Likewise, in Sulawesi and Maluku the internodes

are always glabrous, but otherwise not different and therefore material from these

places are included in the present species.

5. Steudel (1840) claims that [A.]Rich. described F. nigricans. However, I

have not been able to find a publication with this name. It is most likely an ‘in sched.’

name. As it was mentioned for ‘N. Guinea’ it must refer to 4. hispida, the only species

of this genus on New Guinea.

6. I have seen only a photo of the type specimen, and none of the other

specimens that Merrill (1906) cited of F. philippinensis, which he described with

filaments glabrous. A specimen in SING (Baker 3357) identified by Merrill as F.

philippinensis is possibly A. hispida. Unfortunately, this specimen 1s a fruiting one

and has lost all of its stamens. However, the original description of Merrill’s F.

philippinensis 1s otherwise very similar to A. hispida and it is therefore treated as a

synonym of the latter.

Specimens examined: PHILIPPINES. Luzon: Atimonan, Tayabas, Mar 1905, E.D. Merrill 4007

(E); Tayabas, Lucban, May 1906, 4.D.E. Elmer 7410 (E, L); Laguna Province, Mt.Maquiling,

Nov 1912, V. Servinas BS 16877 (L); Mt Mabiling, May 1914, C.F) Baker 3357 (SING); Sorsogon

province, Irosin (Mt Bulusan), Aug 1916, 4.D.E. Elmer 14101 (L, U); Laguna Province, Los

Banos, Mt Maquiling, 1917, A.D.E. Elmer 17973 (L, U); Laguna, Mt. Maquiling, Dec 1935,

C.F. Symington 40928 (KEP); Sorsogon Prov., Mt.Bulusan, 1947, M.D. Sulit PNH 2791 (ct;

SING); Albay Province, Mayon Volcano, Jun 1953, D.R. Mendoza PNH 18305 (L); Sorsogon

Province, Mt Juban, Jun 1956, G.E. Edano PNH 37124 (L); Irosin, Sorsogon Province, Mt

Bangko, May 1957, G.E. Edano & H. Gutierrez PNH 37820 (L); Albay, Mt Malinao, Brgy.

Tagoytoy, So.Sadurong, Oct 1995, E.J. Reynoso et al. PPI 21180 (L). Polillo: Aug 1909, C.B.

Robinson BS 6935 (L); Oct 1909, R.C. McGregor BS 10387 (L). Catanduanes: 1928, M.

Ramos & G. Edano BS 75694 (cf; SING); San Miguel, Brgy Sulung, Mt Saan, Nov 1991,

Barbon et al. PPI 2362 (L); So.Boradan, Brgy.Summit, Nov 1996, E.J/. Reynoso & Majaducon

PPI 24872 (L). Mindoro: Baco River, Mar 1905, E.D. Merrill 4051 (K); Provident tr.Fms. San

Teodoro, Feb 1985, C.E. Ridsdale 802 (L). Negros: May 1908, A.D.E. Elmer s.n. (E). Samar:

Mtuguinao, Jan 1952, FS. Gachalian PNH 15500 (L); Rawis, Sohoton, National Park, May

1971, University of San Carlos 220 (L). Leyte: Dagami, Jul 1957, G.E. Frohne PNH 35671

(L). Mindanao: Davao, Todaya, Mt.Apo, Jul 1909, 4.D.E. Elmer 11156 (cf; E, L); Agusan

Province, Cabadbaran (Mt Urdaneta), Sep 1912, A.D.E. Elmer 13914 (E, L, U); Surigao

Province, Mt Kabatuan, Mar 1949, D.R. Mendoza & P. Convocar PNH 10320 (L); Surigao del

Sur, Aras-asan (Logging area), May 1975, University of San Carlos 793 (L); Dinagat, Agusan

del Sur, Sitio Campo Sies, Agusan, Oct 1991, F.J.M. Gaerlan et al. PPI 4825 (KEP, L).

BORNEO. Kalimantan W: Bukit Blitang, J.E. Teysmann 8244 (cf; L). Kalimantan E:

Boengaloen, Nov 1912, L. Rutten 736 (U); W.Koetai, no 9, nr M Antjaloeng, Jul 1925, FH.

Endert 2134 (L); E.Kutei, Sg Susuk region, Jun 1951, A. Kostermans 5553 (L). Kalimantan

SE: Duson, Muara Larran, PW. Korthals s.n. (L); Hayup (Hayocp), May 1908, H. Winkler

2120 (BM, L).

SULAWESI. North: Rec. Menado, sub div. Paloe, Aug 1937, Eyma 1721 (SING, U); Rec.

Amischotolype in Asia §3

Menado, subd. Loewoek, betw Boenta-Gonohop I, Sep 1938, Evma 3736 (L, U); Toraut Dam.

Mar 1985, E.F- de Vogel & J.J. Vermeulen 6625 (KEP, L); tributary of Sg Ilanga, Feb 1990, JS.

Burley et al. 3519 (SING). Central: E.peninsula, Pinapoean, Dec 1920, T.G. Kaudern 481 (L);

Mt Nokilalaki area, Apr 1975, W. Meijer 9627 (L); rd to Lake Lindu, 60 km SSE of Palu, May

1979, M. van Balgooy 3566 (L); E.of Tongoa, Feb 1981, /. 7. Johansson et al. 37 (L); Batui R.,

Oct 1989, M.J.E. Coode 5941 (L); Lore Lindu NP, Feb 2008, D. Cicurra 596 (cf; L).

MALUKU. Sula: Taliaboe, Kp.Likitobi, Atjeh (Exp.v.Hulstijn) 354 (L); kampong Mangoli, kali

Waj Mana, Sep 1939, S. Bloembergen 4672 (L, SING); Taliaboe, Samoeja, Z van kampong,

Oct 1939, S. Bloembergen 4780 (L). Ceram: Proho, Nov 1917, Kornassi (Exp. Rutten) 518

(L); weg Kairatoe-Honitetoe, Sep 1918, L. Rutten 1617 (L). Buru: Waeha, betw Waelanga-

Lake Kunturun, Nov 1984, M. van Balgooy 4625 (L); Wae Langa, km14, Nov 1984, H.P.

Nooteboom 5101 (L); Wae Duna river, N of Bara, Nov 1984, H.P. Nooteboom 5213 (L); c.10

km S of Bara, Dec 1984, M. van Balgooy 5063 (L). Ambon: 1792, Lahaie 933 (P); Beresih,

Nov 1895, H.G. Botter s.n. (L); 1913, C.B. Robinson 1831 (L). Kai: Groot Kei, Goenoeng

Daal, Apr 1922, H.G. Jensen 114 (L).

NEW GUINEA. PA. Lesson ex J.K. Hasskarl s.n. (L). trian Jaya: Ayambori bei Dore, J.E.

Teysmann 6774 (L); Lorentz river nr Alkmaar, Aug 1907, B. G. Versteeg 1587 (L); ‘Papuarand’

am Tal des Noord-Flusses, Oct 1909, L.S.A.M. von Rémer 397 (L): Moeraira, east coast

Geelvink bay, Jun 1912, R.Fr. Janowski 1 (L); Beaufort River, Nov 1912, A. Pulle 379 (L);

near Pionier biv. Forest, Jun 1920, HJ. Lam 450 (L) & Jul 1920, 659 (L): near Prauwen biv.,

Aug 1920, H.J. Lam 952 (L); Meervlakte, Batavia biv., Nov 1926, WM. Docters van Leeuwen

11179 (L); Bernhard bivouac, Aug 1938, E. Meijer Drees 563 (L); Roode river, Aug 1938, E.

Meijer Drees 663 (L, SING); Idenburg R, Bernhard Camp, Apr 1939, L.J. Brass 13874 (L);

Yapen Island, Jappen-Biak, Sg.Soemboi near Seroei, Aug 1939, Aet & Idjan (exp. L.J.v.Dijk)

505 (L); Yapen Island, Jappen-Biak, Seroei, Sep 1939, Aet & Idjan (exp. L.J.v.Dijk) 824 (L);

Sg Si-era (djera) nr Oeta, Jun 1941, Aet (exp. Lundguist) 261 (L); Vogelkop, Warsui, nr Ransiki

(S of Manokwari), Jul 1948, 4. Kostermans 2734 (L); Sorong, nr Klamono, Aug 1948, D.R.

Pleyte 592 (L); N Bird Head Peninsula, steenkool-Temboeni, Apr 1954, P. van Royven 4036

(L); Hollandia, Nemo, Mar 1956, C.F’ Kalkman BW 3471 (L); Manokwari, Rendani, Oct

1959, VW. Moll BW 9549 (L); Manokwari, Feb 1961, C. Versteegh BW 10450 (L); Hollandia,

Cycloop mountains, Baimoen Creek-Klifon, Aug 1961, P. van Royen & H. Sleumer 6535 (L);

Vogelkop Peninsula, Manokwari, Tafelberg, Oct 1961, P. van Royen & H. Sleumer 6687 (L);

Vogelkop Peninsula, Aifat R valley, nr Sururem, Oct 1961, P. van Royen & H. Sleumer 7029

(L); Wandammen Peninsula, Wondiwoi Mts., Feb 1962, C.F. Koster BW 13643 (L); Bomberai

Peninsula, Sjuga-Wagura area, Armina, May 1962, VW. Moll BW 13034 (L); Kabupaten

Manokwari, Arfak Plains Apr 1994, M.J.S. Sands et al. 6258 (L); Kabupaten Manokwari,

Kecamatan Kebar, May 1994, M.J.S. Sands et al. 6575 (L); Kabupaten Manokwari, ArfakMupi

Dessa, Apr 1995, Mahyar et al. 321 (L); Bird’s Head Peninsula, surr Ayawasi, Nov 1995, C.E.

Ridsdale 2178 (L); Mimika Reg., PT-Freeport Ind Cone Area, toKalikopi , Dec 1998, M/S.

Sands 7217 (L); Mimika Reg., PT-Freeport Ind Cone Area, KualaKence , Mar 1999, FR. Willis

et al. 129 (L); Mimika Reg., PT-Freeport Ind Cone Area, NewEast lev, Apr 1999, PJ. Rudall

107 (L). Papua: Tami-Fluss zwischen den Eti- und Arson-Fliissen, Mar 1910, K. Gjellerup 4

(L); Dieni, Ononge Road, 1933, L.J. Brass 3870 (L); Gogol R., Ninam logging area, Verdcourt

et al. 5133 (L); nr Horata village, Aug 1953, R.D. Hoogland 3571 (L); Naiesonge (Rawa),

nr Gogol R nr Mawan village, Jun 1955, R.D. Hoogland 4929 (L); Sepik, Aitape, Pieni R nr

Walwali vill., Jun 1961, PJ. Darbyshire & R.D. Hoogland 7989 (L); Morobe, Kalapit, Umi

River, Jan 1963, A.N. Miller & P. van Royen NGF 15638 (L); Tymne-Wago track, Mar 1963,

T.G. Hartley 11447 (L); Ossima village, Mar 1964, D. Savers NGF 13209 (L); W bank of

84 Gard. Bull. Singapore 64(1) 2012

Vailala R, c. 3 miles S of jnct w Lohiki R, Feb 1966, R. Schodde & A.L. Craven 4429 (L); nr

Ambunti, May 1966, R.D. Hoogland & L.A. Craven 10149 (L); Tumam, c. 4 km E of Dreikikir,

Jul 1966, PC. Hsyligers 1395 (L); Garagos, Nov 1966, H. Streimann NGF 27962 (L, SING);

Wampit, Bupu village, Jul 1967, A.N. Millar NGF 22925 (L); Ramu, tiver nr Aiome, Mar 1968,

M.J.E. Coode NGF 32759 (L); Vanimo subd., Ossima, Jan 1969, H. Streimann & A. Kairo NGF

39199 (L); Amanab subd, Bedwani mts, nr Kilifas on rv Yenabu, Mar 1970, M.S. Sands 1047

(L); Vanimo, Vanimo-hinterland, Dec 1971, H. Streimann LAE 53820 (L); Kerema, Vailala

Riv, nr Lohike vill, Jun 1980, E.E. Henty & P. Katik LAE 72498 (L); Vanimo, Blackwater creek

logg. area, Sep 1982, K. Kerenga LAE 56478 (L); Mawan, 1km E of Baku, Apr 1983, 4.N.

Vinas AV 226 (L); Angoram, Jul 1985, L.J. Harkink 15 (L); Amanab, above Aimane vill., Oct

1986, O.G. Gideon LAE 57565 (L); Bismarck Range, Ikil R tributary, Oct 1995, W. Takeuchi

10989 (L); Mt Jaya, Mimika Regency, Ecol.Plot 5, 1, 1998, R.J. Johns et al. 8748 (L, SING);

Josephstaal FMA area, nr Kumamdeber, Jul 1999, W. Takeuchi et al. 13578 (L).

9. Amischotolype hookeri (Hassk.) H.Hara (Fig. 9F)

Amischotolvpe hookeri (Hassk.) H.Hara, Flora Eastern Him. I (1966) 399; Hong, Acta

Pytotax. Sin. 12,4(1974) 461; Fl. Reipubl. Pop. Sin. 13,3 (1997) 71; Hong & DeFilipps,

Fl. China 24 (2000) 23 (See note). — Forrestia hookeri Hassk., Flora 47 (1864) 629;

Commelin. Ind. (1870) 89; Clarke, Commelyn. Cyrtandr. Bengal. (1874) 61, tab.XLI;

Clarke in A.Candolle & C.Candolle, Monogr. Phan. 3 (1881) 237 (see note); Hooker

f., Fl. Brit. India 6 (1894) 384; Briickner in Engler & Prantl, Nat. Pflanzenfam. ed. 2

(1930) 169; Rao et al., Proc. Indian Sci. Congr. (1960) 366. SYNTYPES: Hooker &

Thomson s.n., S.d., India contin. regionem tropicam montium Khasiae (inter 14000’)

(W, K, M, G-DC, P, all n.v.; B photo (Barcode: B 10 0296341 / Imageld: 302934), L);

Hooker & Thomson s.n., s.d., reg. trop. Sikkim (G-DC, K, all 7.v.).

Stem rhizome unknown; erect part 120-210 cm long, simple; internodes moderately

0.5 mm long hairy. Sheath 9-14 mm diam., 0.5(—1) mm long hairy and/or with one to

several lines of 1.5—2 mm long, pale-yellowish hairs, mouth ciliate. Leaf blade 20-35

x 7—10.4 cm, 2.9-4.4 times as long as wide, smooth, base gradually to rather abruptly

narrowed into indistinct pseudopetiole; lower surface on veins densely 0.1—0.5 mm

long hairy, upper surface glabrous; submarginal hairs on lower surface and on margin,

(0.5—)1.5—2 mm long, dense, whitish to yellow. Inflorescence on erect stem, sessile,

2—3.5 cm diam., dense with branches obscure, 10—25-flowered. Pedicel 0—0.7 mm

long. Sepals 7-10 = 3—3.5 mm, not elongating in fruit, green to deep purple, glabrous

or rarely sparsely 0.5 mm long ciliate margins and keel, tip hooded. Petals c. 9 x 2.5—

3.5 mm, c. as long as sepals, white to pale pink, glabrous, margin unknown. Stamens

filament c. 9 mm long, colour unknown, in upper | mm with 2 mm long hairs; anthers

(2—)2.2-3 = 0.7-0.9 mm, colour unknown, thecae opening by a longitudinal slit.

Capsule 9-14 = 5.5—7 mm, ovoid, longer than sepals by (1—)3.5—S5 mm, whitish to

red or purple, sparsely 0.5—1(—1.5) mm long hairy; valves free to fused in basal half;

apex acute, lobes absent; style remnant 0.3—0.5 = 0.4 mm long, persistent. Seeds 2 per

locule, aril colour unknown.

Amischotolype in Asia 85

Chromosomes. 2n = 36 (Kammathy 81236: L, E; unpublished but in agreement with

Rao et al. 1960).

Distribution. India (Assam, West Bengal, East Bengal, Meghalaya, Sikkim),

Bangladesh (Moulavi Bazar).

Ecology. Along stream in forest. Altitude: 1—-600(—1500) m asl.

Notes. The species is only known from NE India and Bangladesh. Several authors

included Yunnan in the distribution of this species (Hara 1966, Hong 1974, 1997

and Hong & DeFilipps 2000). However, I have not seen their specimens and their

descriptions are wide enough to include also A. divaricata. Study of the specimens

from Yunnan is needed to elucidate which of the two taxa are present.

Clarke (1881) mentioned the species for the island of Pinang (Campelia

marginata, Wallich Cat. 8977 p.p., non Blume). In the Wallich collection at K-W

(studied on microfiche), three sheets of Cat 8977 exist: one is 8977b, from Singapore

(= A. gracilis); two are 8977a, from Pinang, one of them without doubt 4. marginata;

the identity of the second sheet cannot be ascertained from the microfiche. A fourth

photo of Wallich Cat 8977 in K-W has been found on the Kew-website. The sheet

has three branches, so is not entirely similar to the ones on microfiche (perhaps

remounted?), and not indicated a or b: one with a creeping stem with inflorescences

agreeing with 4. marginata, one sterile, and one with inflorescences on the erect part

of the stem. The latter two cannot be identified with certainty. If, however, one of the

collections of Pinang is indeed 4. hookeri, man must have brought it there, probably

unintentionally with planting material for, for instance, the spice gardens (species of

the genus are often found growing together with wild gingers) that were etablished on

the island in 1794. Likewise, in Singapore it has only been found growing in the rain

forest in the Botanic Gardens where it was undoubtedly introduced, either by accident

or as an ornamental. It has not been observed in flower, and has not been collected for

the herbarium. Clarke 37582 from Brunei is most likely mislabelled.

Specimens examined: INDIA. cult., Bot. Gardens of Calcutta and Serampore, 1834-1841, /.O.

Voigt s.n. (P). Assam: W. Griffith 1345 (BM); Jenkins s.n. (L, P); Jenkins 9383 (BM); Naga

hills, Masters s.n. (L, P); Naga Hills, Pereumi, Aug 1935, N.L. Hor 6454 (K); Khasi Hills,

Nongpoh, Jun 1949, WN. Koelz 23015 (L); K.&J. Hills Distt., nr Exp.Plot 2, Nongpoh, Feb

1962, R.V. Kammathy 81236 (BM, E, K, L). West Bengal: Darjeeling, Selim, Sep 1869, C.B.

Clarke 9082 (K, SING); Darjeeling, Rishup, Oct 1870, C.B. Clarke 13629A (K); Darjeeling,

Garidoora, May 1884, C.B. Clarke 35539B,F (BM, K); Tista, Jun 1913, G.H. Cave s.n. (E):;

Darjeeling, Jun 1923, JM. Cowan s.n. (K). East Bengal: (1864), W. Griffith 5487 (L, P).

Meghalaya: montium Khasiae, J.D. Hooker & T. Thompson s.n. (BM, E, L, P, U); Khasia,

Bhorlasa, Aug 1886, C.B. Clarke 44469E (BM). Sikkim: J.D. Hooker s.n. (BM, K, P); J.D.

Hooker & T. Thompson s.n. (BM, E, L, P, U); Jun 1870, C.B. Clarke 11805 (SING) & 11806B

(BM); Mongpo, Oct 1884, C.B. Clarke 36264B (K); Dulkajhar, Oct 1884, C.B. Clarke 37048

(BM).

BANGLADESH. Moulavi Bazar: Adampur forest, Rajkandi range, Jun 2000, 4.M. Huq &

86 Gard. Bull. Singapore 64(1) 2012

M.K. Mia 10748 (L).

SINGAPORE. grown in Hort.Kew, 1904, Anon. 461-95 (K).

BORNEO. Brunei: Mar 1885, C.B. Clarke 37582 (BM).

10. Amischotolype irritans (Ridl.) 1.M.Turner (Fig. 9G & 13C)

Amischotolype irritans (Ridl.) 1.M.Turner, Novon 6 (1996) 221 & Gard. Bull. Sing.

47 (1997) 515. — Forrestia irritans Ridl., J. Straits Branch Roy. Asiat. Soc. 41 (1904)

38; Mat. Fl. Malay. Penins. 2 (1907) 124; Fl. Malay. Penins. 4 (1924) 361; Briickner

in Engler & Prantl, Nat. Pflanzenfam. ed. 2 (1930) 169; non Beaman & Beaman, The

Plants of Mount Kinabalu 3 (1998) 109 (see note 1). LECTOTYPE (designated by

Turner (1996)): Ridley s.n., Peninsular Malaysia, Selangor, on the Tras Route at the

15th mile (SING).

Stem ascending from a creeping or scrambling rhizome up to 200 cm long; erect part

50-120 cm long, simple; internodes rather sparsely to densely 0.2—0.5 mm long red-

hairy, hairs easily caducous, rarely glabrous. Sheath 10-20 mm diam., glabrous or

rarely with a few 0.5 mm long hairs basally at the back, mouth glabrous or (towards

pseudopetiole) ciliate. Leaf blade 23-44 x (4—)5—9 cm, 3.8—6.2 times as long as wide,

smooth, base rather gradually narrowed into c. 5.5 cm long and 2—3 mm wide winged

pseudopetiole; both surfaces moderately 1-2 mm long yellow- to orange-hairy;

submarginal hairs on upper surface, 1—1.5 mm long, rather dense, yellow. Inflorescence

on erect stem, sessile, 2.5—S cm diam., very dense with branches obscure, 20—many-

flowered. Pedicel c. 1.5 mm long. Sepals 6.5—10.2 (in flower), elongating in fruit to

10-13 « 1.5—2 mm, yellow-white or pale green (before and in flower) to pink or reddish

purple (in fruit), moderately to rather densely 1.5—2.5 mm long spiny red-hairy, tip not

hooded. Petals 9-10 x 1.8—2 mm, slightly longer than sepals, white, dorsal surface

subapically with 2 mm long spot of 1—1.5 mm long, red-spiny hairs, margin at apex

minutely fringed. Stamens with filaments 11-12 mm long, white, glabrous or in an

upper part up to 2-3 mm below apex with few 2—3 mm long hairs; anthers 1.5—2 x

0.5—0.7 mm, white or yellowish, thecae opening by a longitudinal slit. Capsule 5.5—6

x 3.5-4 mm, narrowly ovoid, shorter than sepals by 3—4.5 mm, bright purple, apex

1—2 mm long, red spiny-hairy; valves free; apex slightly depressed, lobes absent; style

remnant 0.5 x 0.2 mm, finally deciduous. Seeds 2 or rarely | per locule, aril colour

unknown.

Distribution. Thailand (Peninsular Thailand: Pattani), Peninsular Malaysia (Kedah:

Langkawi, Kelantan, Terengganu, Perak, Pahang, Selangor, Negeri Sembilan),

Sumatera (N: Leuser, Asahan).

Ecology. Primary to rather disturbed, riverine or lowland dipterocarp forest, forest

margin or forest regrowth, dry to wet (alluvial) soil. Altitude: 15—750 m asl.

Amischotolype in Asia 87

Notes. 1. Ridley (1904, 1907) described it as a rather rare endemic of Peninsular

Malaysia from mountain forests (2000 feet alt.). It has since been found at lower

altitudes, in Sumatera (three times) and in Thailand (once). The species has been

cited for Borneo (Beaman & Beaman 1998: Clemens 26141) but this is based on a

misidentification of A. monosperma.

2. Ridley (1904, 1907) mentioned he had never seen fruits and he did not

mention them in 1924. However, based on the number of collections with fruits, they

are normally formed, and were for the first time collected in 1921 (SING: Hume 9382,

Peninsular Malaysia: Selangor). The colour of the aril has never been annotated and,

unfortunately, I have only seen them in flower in the field.

Specimens examined: THAILAND. Peninsular Thailand: Pattani, Aug 1923, 4.FG. Kerr

7516 (L, P).

PENINSULAR MALAYSIA. Kedah: Langkawi, Dec 1990, Khairuddin Hj. Itam 10 (KEP).

Kelantan: path to Bukit Berangkat, via Kampong Perak, Bertam, Feb 1973, Md.Shah &

Md. Ali 2867 (KEP, SING); Gua Musang, Relai Forest Reserve, Sep 1992, Hamid 13 (KEP).

Terengganu: Kemaman, Bukit Kajang, Nov 1935, Corner SF 30238 (K, SING); Hulu

Terengganu, Tasik Kenyir, Sg. Cicir, Aug 2007, K. Imin et al. FRI 58335 (KEP). Perak: 188x,

HN. Ridley s.n. (SING); Bujong Malacca, 1898, H.N. Ridley 9784 (SING): Pondok Tanjong

Forest Reserve, Feb 1939, G.H. Spare SF 36220 (K, KEP, SING). Pahang: 7 mile Raub Road,

Jul 1924, G.A. Best SF 14132 (SING); Sungai Tahan, near Kuala Teku, Sep 1928, R.E. Holttum

SF 20810 (SING); Gunong Serudom, Kuantan, Oct 1975, Md.Shah et al. 3762 (KEP, KLU,

SING); Jerantut, Taman Negara, Gunung Tahan, Sunga Kelam, Aug 1996, R Kiew 3991 (L);

Temerluh, Krau GR, Sg. Perlok, Nov 2000, S. Damahuri & K. Avyau FRI 43603 (L). Selangor:

on the Tras route at the 15th mile, 1897, H.N. Ridley s.n. (SING); Semempik, Jul 1921, HL.

Hume 8093 (SING); Ginting Simpak, Oct 1921, H.L. Hume 9382 (SING); 16 mile Ulu Gombak,

Oct 1937, Md. Nur SF 34211 (SING): Ulu Gombak, Jun 1966, J.C. Carrick 1485 (K, L, SING):

Lalang FR, Semenyih, Kajang, Oct 1967, J. Dransfield s.n. (KLU); Pahang border, Gentiang

Sempah trail, Sep 1973, B.C. Stone 11661 (KLU); Gombak, Ulu Gombak Forest Reserve, Sep

1974, Tho Yow Pong FRI 21544 (KEP, L); Ulu Langat, road to JKR Quarry, Jul 1984, R Kiew

1335 (SING); N. of Kuala Lumpur, JIn Gombak km36 (KL), Jun 2005, H. Duistermaat 351

(L, SING). Negeri Sembilan: Sungei Ujong, Kupayang, Nov 1885, YH. Alvins 2202 (SING):

Gunung Angsi, Jun 2005, H. Duistermaat 346 (L, SING).

SUMATERA. Aceh: Leuser, Kluet NR, along Krung Lembang, Jul 1985, WJ.J.O. de Wilde

& B.E.E. de Wilde-Duyfjes 19846 (L). Sumatera Utara: Asahan, East Coast, Vale of Tangga,

May 1927, H.H. Bartlett 7707 (L); East Coast, vicinity of Tomoean Dolok, Aug 1936, Rahmat

Si Boeea 10005 (L).

11. Amischotolype laxiflora (Merr.) Faden (Fig. l|OA & 14A—B)

Amischotolype laxiflora (Merr.) Faden in Coode et al., Checklist Fl. Plants & Gymn.

Brunei Dar. (1996) 352, 434; Cowley & Furness, Kew Bull. 52 (1997) 469. —

Forrestia laxiflora Merr., Philipp. J. Sci. 29 (1926) 354. TYPE: Castro & Melegrito

1618, August 1923, Malaysia, Sabah, Banguey Island, in forests, altitude about 30 m

(holo UC, n.v.: iso K, BO).

88 Gard. Bull. Singapore 64(1) 2012

Forrestia marginata auct., sensu Hallier f., Beih. Bot. Centralbl. 34 (1916) 47, p.p.

(see note 1); non Blume.

Stem ascending from trailing, 100-150 cm long rhizome; erect part 20-75 cm long,

simple; internodes glabrous or sparsely to moderately 0.1—0.3 mm long hairy. Sheath

6—14 mm diam., with several moderate to dense lines of (2—)3—5 mm long, yellow(-

brown) hairs, mouth ciliate. Leaf blade 15—32 = (2.7—)4—8(—9.7) cm, 2.6-4(—S5.2) times

as long as wide, bullate, lower surface often purple, base gradually to rather abruptly

narrowed into an indistinct winged pseudopetiole to 4 cm long and 4 mm wide; lower

surface glabrous or midvein in lower half with 1-2 mm long yellowish hairs or rarely

0.1 mm long colourless- to brownish-hairy, upper surface glabrous or rarely 0.1 mm

long hairy; submarginal hairs on upper surface or rarely absent, 0.1—0.5(—0.8) mm

long, sparse to rather dense, colourless or yellow. Inflorescence on rhizome or at knee,

peduncle (0—)3—15 mm long, 2—5 cm diam., rather dense to very lax with branches

obscure or the longest 0.8—3 cm long, 15—many-flowered. Pedicel 1.5—2.5 mm long.

Sepals (9-)10—14 x 1.5—2.5 mm, slightly elongating in fruit, white, green, pinkish or

red-purple (in flower) and purple (and green), red, magenta or violet (in fruit), glabrous

or sparsely 0.5 mm long ciliate margins and keel, tip hooded. Petals c. 6 x 2 mm,

shorter than sepals, white to yellow, glabrous, margin entire. Stamens with filaments

c. 5 mm long, white, upper 2 mm with many 2 mm long hairs; anthers | x 0.9 mm,

cream-coloured or yellow, thecae opening by a longitudinal slit. Capsule 5.5—8.5 x

3.5—5.5 mm, obovoid, shorter than sepals by 2—5 mm, red or purple, glabrous or apex

to upper half sparsely 0.2—0.5 mm long hairy; valves (almost) free; apex depressed,

lobes absent; style remnant 0.3 = 0.3 mm, deciduous. Seeds 2 per locule, aril orange.

Distribution. Borneo (Sabah: West Coast, Interior, Kudat, Sandakan, Tawau, Sarawak:

Kuching, Sri Aman, Kapit, Bintulu, Miri; Kalimantan: W, C, E, S), Sulawesi (S).

Ecology. Primary or disturbed mixed dipterocarp, riparian or swamp forest, along

streams, in wet or swampy places, on loam soil, often on limestone. Altitude: 0-800

m asl.

Notes. 1. The species was originally described from Banguey Island (N Sabah). It has

been collected outside Borneo only once, in Sulawesi (van Balgooy 3910). It is often

identified as the widespread 4. marginata, which also occurs on Borneo, and from

which A. /axiflora differs as mentioned in the key (lead 9). Kessler et al. 938 appears

to be a mixture of the two species, showing that the species can even grow in mixed

populations: the specimen in L 1s A. /axiflora, in US it is A. marginata (n.v.; pers.

comm. R.B. Faden).

2. The roots are said to have medicinal value (Forman 452, E. Kalimantan,

Gunung Sahari).

Specimens examined: BORNEO. Sabah: West Coast: Ranau, Sungai Lakus, Kg.Segindai, Aug

1983, S. Dewol et al. SAN 100162 (L, SAN); Interior: Pensiangan Kayu FR, Jul 1992, K.

Amischotolype in Asia 89

Fidilis SAN 136002 (KEP, L); Pensiangan, Pun batu, along the path to limestone, Oct 1996, A.

Berhaman et al. 139 (SAN); Kudat: Banguey Island, 1923, P. Castro & F. Melegrito 1618 (BM,

BO, K); Sandakan: Bettotan, near Sandakan, Jul 1927, Boden-Kloss SF 18736 (K, SING); Kabili

FR, May 1935, H.G. Keith 38907 (KEP); Kabili forest reserve, May 1935, H.G. Keith 4531 (L,

SING); Gomantong, Feb 1960, W. Meijer SAN 20549 (SAN); Sepilok FR, Aug 1968, K. Ogata

10790 (L); Sepilok FR, Dec 1975, PF: Stevens et al. 328 (KEP, L); Sepilok FR, Oct 1979,

R. Kiew 786 (KEP); Sepilok FR, Oct 1981, Kwiton Jong SAN 93914 (SAN); Kinabatangan,

Sungai Menanggul, Feb 1985, Amin & Matin SAN 69950 (SAN); Lahad Datu, Danum Valley,

West Trail, Feb 1986, S. Andrews 725 (E, SAN); Lahad Datu, Ulu Segama, Danum Valley,at

entrance to west trail, Mar 1987, G. Argent et al. GA 671987 (E, L, SAN); Lahad Datu, Danum

valley, Sep 1987, Campbell 23/9/4 (E); Lahad Datu, Lahad Datu, Tabin Wildlife Sanctuary,

Aug 1991, L. Madani SAN 132613 (KEP, L); Lahad Datu, Lahad Datu, Tabin WR, trail to

Lipad mudvolcano, May 1998, F: Borchsenius 513 (SAN); Tawau: floodplain of Tawau River,

Tawau Hills Park, Jun 1984, J.H. Beaman 10210 (KK); Tawau Hills NP, path to waterfall, May

2005, J.J. Vermeulen in H. Duistermaat 357 (L); Sarawak: Kuching: lower slopes gng Lundu,

B.L. Burtt & P.J.B. Woods 2723 (E); Sri Aman: Bukit Danau, Aug 1938, Daud & Tachun SF

35750 (KEP, SING); Kapit: Belaga, hills behind airfield, Apr 1963, P.S. Ashton S 18262 (L);

Sungai Apa, Mar 1975, P-C.K. Chai S 36195 (KEP, L); Belaga, Ulu Belaga, Sungai Semawat,

Oct 1981, C. Hansen 716 (KEP, L, SING); Ulu Sg. Balleh, Sungei Sedupak, Yii P.-C. et al. S

52067 (KEP, L); Belaga, Ulu Batang Balui,Ulu Sungai Kebhor, Mar 1989, Yii PC. S 62338

(L); Song, Ulu Katibas, trib.Sg.Beloh, Sg.Joi, Jun 1993, Yii PC. et al. S 64900 (L); Bintulu:

Tatau, Bukit Sarang group, Bukit Lebik, Aug 2005, J.J. Vermeulen in H. Duistermaat 401 (L,

SING); Miri: Subis, Niah., Oct 1954, Ahmad 5 (SING) & 5 (B) (SING); Marudi, Gng Api,

gorge on S side, Feb 1966, J.A.R. Anderson S 24025 (L); Marudi, C.Mulu NP, Sg.Mendalam,

Oct 1977, J.A.R. Anderson S 39393 (L); Marudi, Dulit Range, Sg Sirui, Oct 1983, Dyg. Awa

& Vii P.-C. S 46611 (L); valley in front of Niah Caves, B.L. Burtt & P.J.B. Woods B 2008 (EB);

Kalimantan W: Liang Gagang, “Oberlauf des Manddeistromes”, Mar 1894, H. Hallier B3001

(L); Sungei Pary, 1896/97, Jaheri (Exped. Nieuwenhuis) 1136 (BO); Gn Palung NR, 100 km

S of Pontianak, Jun 1986, M. van Balgooy & A.K. van Setten 5618 (L). Kalimantan C: Bukit

Raya, Tumbang Tapi, Jan 1983, H. Wiriadinata 3357 (L). Kalimantan E: W. Koetai, n.5 nr

Lahoen, Jun 1925, FH. Endert 1768 (L, SING); W.Koetai, Aug 1925, FH. Endert 2790 (L):

E.Kutei, Sg. Susuk region, Jul 1951, 4. Kostermans 5744 (L); Gunung Sahari, Belajan river,

Aug 1956, L.L. Forman 452 (L); Balikpapan, Wanariset, Sep 1982, Ramlanto 8 (L); Wanariset

area, Samboja-Semoi km 11, Feb 1991, Ambri & Arifin W 647 (L); PT. ITCI, Kenangan-G.

Meratus km 69, Mar 1995, P. Kessler et al. 938 (L). Kalimantan S: Kabupaten Tabalong, Jul

2000, K. Sidivasa & Z. Arifin 2137 (L).

SULAWESL S: Soroako-Wasuponda Road, km 19, Jul 1979, M. van Balgooy 3910 (K, KLU, L).

12. Amischotolype leiocarpa (Hallier f.) Duist. comb. nov. (Fig. 10B)

Forrestia rostrata Hassk. var. leiocarpa Hallier f., Beih. Bot. Centralbl. 34 (1916)

49: Merrill, J. Straits Branch Roy. Asiat. Soc. (1921) 113. TYPE: Hallier f’ B3002, 29

March 1894, W. Borneo, Liang Gagang (holo BO, n.v.; iso L).

Stem rhizome unknown; erect part 90-200 cm long, stilt roots sometimes present,

simple; internodes glabrous or moderately to densely 0.2—0.3 mm long hairy. Sheath

5—9 mm diam., glabrous or moderately 0.2—-0.5 mm long hairy, mouth (sparsely)

90 Gard. Bull. Singapore 64(1) 2012

ciliate. Leaf blade 18—28 x (3.3—)4—7(—10) cm, (2.4—)3—6(—7.6) times as long as wide,

base rather to very abruptly narrowed into 1.5—5 cm long and 1 mm wide winged

pseudopetiole; lower surface glabrous or rarely moderately 0.2 mm long hairy, upper

surface glabrous; submarginal hairs absent or on lower surface, rarely on upper

surface, 0.1—0.3 mm long, sparse. Inflorescence on erect stem, peduncle 0—5 mm long,

1.6-3.5 cm diam., (rather) lax or rarely dense with longest branches 0.3—1.3 cm long,

10—30-flowered. Pedicel 1—1.5 mm long. Sepals 4.5—6.5 x 2—3 mm, not elongating

in fruit, green or reddish, glabrous or very sparsely 0.2—0.3 mm long ciliate keel and

margins, tip hooded. Petals unkown. Stamens with filaments unknown; anthers with

thecae opening by a longitudinal slit in upper half only, otherwise unknown. Capsule

6.5—8.5 « 3.8—6.5 mm, short-pyriform, longer than sepals by 1.5—4.5 mm, (light) green

to reddish or purplish, glabrous; valves (almost) free; apex (slightly) depressed, lobes

absent; style remnant 0.5—1.5 = 0.2—0.3 mm, persistent. Seeds 2 per locule, aril orange.

Distribution. Borneo (Sabah: West Coast, Interior, Sandakan; Sarawak: Kapit, Miri;

Kalimantan: W).

Ecology. Primary or logged over mixed dipterocarp (hill or riparian) forest, on hillside

or along stream, moist, rocky or on granite. Altitude: (230—)500—1200(—1500) m asl.

Notes. 1. Described as a variety of A. rostrata by Hallier (1916), said to differ from the

species only in the entirely glabrous capsules. However, this study revealed differences

in the number of flowers per inflorescence, the length and shape of the capsule, and the

valves being free or fused, warranting its recognition at species level. It is endemic to

Borneo.

2. Most collections are in fruit. The single collection with anthers was returned

from loan before all details were described (Kokawa & Hotta 4846, SAN; in L without

anthers).

Specimens examined: BORNEO. Sabah: West Coast: Dallas, Sep 1931, J. & M.S. Clemens

26396 (BM); Mount Kinabalu, Dallas, Oct 1931, J. & M.S. Clemens s.n. (BM); Mount

Kinabalu, Penibukan, Jan 1933, J. & M.S. Clemens s.n. (BM); Kota Belud, Mt Kinabalu,

Mahandui river, Mar 1933, C.E. Carr SF 26297 (SING); Mt Kinabalu, Penibukan, Mar 1933, J.

& M.S. Clemens 32085 (BM, L); Ranau, Kinabalu NP, along Sg.Mamut, nr Poring, Feb 1969,

S. Kokawa & M. Hotta 4846 (L, SAN); Kinabalu, Penibukan, Bahandoi (Sg Tahubang), Mar

1970, H.P. Nooteboom & Aban 1503 (L, SAN); Ranau, Langanan Fall, Poring, Feb 1989, E.P.

Tay et al. 247 (KEP, SING); Kota Belud, Kinabalu P, Kg.Sayap, end rd to Wariu Waterfall, Jan

1991, Jamili Nais SNP 4674 (SAN); Kota Belud distr, Kpg Sayap, Kemontis, May 1996, Yalin

Surunda 101 (KEP); Kota Belud, Kinabalu P, Sayap, Lumangis trail, Jun 2000, 4.D. Poulsen

et al. 1620 (SAN); Ranau, Ulu Tungud FR, Jul 2005, L.G. Saw et al. SAN 146080 (KEP);

Interior: Tambunan, Mt.Trusmadi, Mar 1969, H.P. Nooteboom 1475 (L, SAN); Tenom, Kaang,

Mar 1987, Asik Mantor SAN 120105 (SAN); Keningau, Hs.Trusmadi, May 1988, J. Bousi

et al. SAN 123971 (SAN); Trus Madi FR, May 1988, B. Joseph et al. SAN 124017 (E, KEP,

L); Sandakan: Lahad Datu, Ulu Segama, Dec 1982, K. Fidilis SAN 95555 (KEP, L, SING).

Sarawak: Kapit: Belaga, Batang Balui, Ulu sg Penuan, NWsl BtTasu, Mar 1987, Yii PC. S

Amischotolype in Asia 9]

53627 (L); Miri: Marudi, Mulu National Park, May 1985, 4.P. Abang Mohtar & I. Othman

S 49695 (KEP): Marudi, Sg Silat Bassin, Sg Palutan, Mar 2003, N. Normaya et al. S 91072

(KEP). Kalimantan W: Liang Gagang, Mar 1894, H. Hallier B3002 (L); Serawai, Feb 1995,

A.C. Church et al. 1971 (L).

13. Amischotolype lobata Duist. sp. nov. (Fig. 4)

Cum Amischotolype mollissima capsulae apicis lobis longis congruens, pseudopetiolo

angustissime alatis, laminis supra venis distinctis, capsulis calyce superantibus pilis

setosis differt. TYPUS: Argent et al. SAN 108301, March 1985, Malaysia, Sabah,

Lahad Datu, Ulu Sg.Segama (holo SAN, iso E).

Stem ascending from rhizome; erect part 50-150 cm long, simple; internodes glabrous

or rather sparsely to moderately 0.3-0.7 mm long hairy. Sheath 6-13 mm diam.,

moderately 0.3—0.5 mm long hairy at least at the front in upper half and/or with a

few lines of 24 mm long, yellow-brown hairs, mouth (sparsely) ciliate. Leaf blade

(17-)25-33 x (3.7—)5.6—7.5 cm, 3.7—5.7 times as long as wide, lower surface purplish

or not, base very abruptly narrowed into 3.5—9.5 cm long and 0.5—1 mm wide winged

pseudopetiole; lower surface sparsely to densely 0.2—0.5(—1) mm long hairy or rarely

glabrous, upper surface with veins very distinct, glabrous or sparsely to moderately 0.2

mm long hairy or rather sparsely 1.5—3 mm long hairy near midvein; submarginal hairs

on upper surface, 1.2—-2 mm long, dense, yellow. Inflorescence on erect stem, sessile,

2-3 cm diam., dense with branches obscure, 10—40-flowered. Pedicel 1—1.5 mm long.

Sepals 9-11.5 x 34.5 mm, not elongating in fruit, white and purplish tinged or pale

green (in flower) to magenta or purple (in fruit), glabrous or sparsely to moderately

0.5—0.7 mm long ciliate margins and keel, tip hooded. Petals c. 8 x 2 mm, shorter

than sepals, colour unknown, glabrous, margin unknown. Stamens with filaments c.

6 mm long, colour unknown, upper 2 mm with 0.5—1 mm long hairs; anthers 1.2

0.6 mm, colour unknown, thecae opening by a longitudinal slit. Capsule 7.5—10 =

5.5—6 mm, obovoid, 1 mm shorter to 0.5—1 mm longer than sepals, green, upper 1/3

to half moderately 1-2 mm long, yellowish bristle-hairy; valves (almost) free; apex

depressed, lobes 1—-1.5 mm long lobes; style remnant absent. Seeds 2 per locule, aril

colour unknown.

Distribution. Borneo (Sabah: West Coast, Interior, Kudat, Sandakan, Tawau;

Kalimantan: E).

Ecology. Primary, severely logged-over or secondary dipterocarp hill or riverine forest,

on sandstone. Altitude: 90-800 m asl.

Notes. The species is endemic to the northeastern part of Borneo, mostly Sabah,

and only one collection is from East Kalimantan (Geesink 9284). Even vegetatively,

this species is readily recognised by the distinct and only very narrowly winged

pseudopetioles and the distinct veins on the upper surface of the leaf blade. In fruit, the

92 Gard. Bull. Singapore 64(1) 2012

Wi WALSMIT SACHS 2042

Fig. 4. Amischotolype lobata Duist. A. Habit. B. Immature fruit with persistent sepals. Drawing

by A. Walsmit Sachs, from R. Geesink 9284 (L).

Amischotolype in Asia 93

long lobes at the apex of the capsule are striking (hence the name): this is a character

it shares only with A. mollissima. Differences from A. mollissima include the shape of

the leaf blades, the bristly hairs on the capsules, and the capsules generally exceeding

the sepals.

Specimens examined: BORNEO. Sabah: West Coast: Ranau, NW of Kampong Pinawantai,

May 1973, G. Shea & Aban SAN 76938 (SAN): Ranau, c. 8 miles from Kampung Merungin,

Novy 1975, Leopold & Saikeh SAN 82580 (SAN); Interior: Tenom, Crocker Range, Melalap, Sg

Losong, Nov 1968, K. Ogata 11466 (L); Sipitang, G-Lumaku, Mar 1969, H.P. Nooteboom 1119

(L); Kudat: Kota Marudu, Kampong Monggis, May 1996, . Rumutom 287 (KEP); Sandakan:

Labuk Sugut, W-side of Bt Doji and Telupid-UluKaram, Oct 1968, S. Kokawa & M. Hotta 485

(L); Kinabatangan, Sungai Kuamut, Jan 1976, PF. Stevens et al. 439 (KEP, L): Lahad Datu,

N. of camp 3 Ulu Sg. Danum, Sep 1976, B.C. Stone et al. SAN 85248 (KLU): Mt.Tawai FR

Karamuak, Jul 1978. S. Dewol & M. Alexius SAN 88695 (SAN); Lahad Datu, Ulu Sg.Segama,

Mar 1985, G. Argent et al. SAN 108301 (E, SAN); Lahad Datu, Danum Valley W1355, Oct

1985, MJ. Still SAN 112110 (SAN); Lahad Datu, Danum Valley, Palum Tambun Nature Trai,

Jul 1990, Campbell EG 109 (E):; Kinabatangan, Sungai Imbak, Jun 2000, A.D. Poulsen &

K. Kjeldsen 1609 (AAU, KEP): Tawau: Tawau hill park, Apr 1992, 4. Berhaman et al. SAN

134514 (SAN). Kalimantan E: Papadi-Pamilau, Aug 1981, R. Geesink 9284 (L).

14. Amischotolype marginata (Blume) Hassk. (Fig. 1OC, 14¢C—D, 1SA-F & 16A-B)

Amischotolype marginata (Blume) Hassk., Flora 46 (1863) 392; Turner, Gard. Bull.

Sing. 45 (1993) 53; Coode et al., Checklist Fl. Pl. Gymn. Brun. Dar. (1996) 352:

Turner, Gard. Bull. Sing. 47 (1997) 515; Beaman & Beaman, The Plants of Mount

Kinabalu 3 (1998) 110; Keng et al., Conc. Fl. Sing. Volume II: Monocotyledons

(1998) 23. — Campelia marginata Blume, Enum. PI. Javae (1827) 7; Kunth, Enum.

pl. 4 (1843) 109: Moritz, Syst. Verz. (1845-46) 93; Zollinger, Syst. Verz. (1854) 65;

Miquel, Fl. Ned. Ind. 3 (1855) 547. — Forrestia marginata (Blume) Hassk., Flora 47

(1864) 630, p.p. (see note 5); Commelin. Ind. (1870) 90, p.p.; Clarke in A.Candolle

& C.Candolle, Monogr. Phan. 3 (1881) 237: Hooker f., Fl. Brit. India 6 (1894) 383;

Ridley, J. Straits Branch Roy. Asiat. Soc. 33 (1900) 170; Mat. Fl. Malay. Penins. 2

(1907) 123 (see note 3); Koorders, Exkursionsfl. Java 1 (1911) 282: Hallier f., Beth.

Bot. Centralbl. 34 (1916) 47, p.p.; Merrill, J. Straits Branch Roy. Asiat. Soc. (1921)

113; Ridley, Fl. Malay. Penins. 4 (1924) 360; Briickner in Engler & Prantl, Nat.

Pflanzenfam. ed. 2 (1930) 169: Cherfils in Lecomte, Fl. Gen. Indo-Chine 6 (1937)

856, t.85, f. 1-3 (‘marginatus’; see note 4); Keng, Gard. Bull. Sing. 40 (1987) 124 (see

note 3). Forrestia mollissima (Blume) Koord. forma marginata (Blume) Backer,

Handb. FI. Java I, 3 (1924) 33: Backer & Bakhuizen van den Brink, FI. Java 3 (1968)

15. — Amischotolype mollissima (Blume) Hassk. forma marginata P.H.H6, Cayco

Vietnam 3, 1 (1993) 478, illus. (“Amischolotype’), nom. inval., no basionym (see note

4). TYPE: Blume s.n., ‘In umbrosis montium Gede, Salak, etc.’ (holo L).

Forrestia hispida auct., sensu Hook.f., Bot. Mag. 90 (1864) t.5425; non A.Rich.

94 Gard. Bull. Singapore 64(1) 2012

Stem finally ascending from creeping and branched, up to 300 cm long rhizome; erect

part 30-100 cm long, simple; internodes glabrous to rather densely 0.2—0.5 mm long

(red-)hairy. Sheath (6—)10—20(—24) mm diam., with a purple tinge or not, with few

to many lines of 1-6 mm long, yellowish hairs, rarely glabrous or densely 0.1—0.5

mm long red-hairy, mouth ciliate. Leaf blade (15—)20—35(-46) = (3.4—)4.5—7(-10.6)

cm, (3.1—)4-6(—7) times as long as wide, bullate, lower surface purple or not, base

gradually to rather abruptly narrowed into an indistinct or 2-3 cm long and 2-3 mm

wide winged pseudopetiole; lower surface glabrous or sparsely to very densely 0.1—

0.5(—1) mm long hairy or rarely midvein in lower half with 2-3 mm long hairs, upper

surface glabrous or rarely sparsely to moderately 0.1—0.5 mm long hairy; submarginal

hairs on upper surface, 0.1—0.5(—1) mm long, rather sparse to moderately dense,

colourless or yellowish. Inflorescence on rhizome and knee, peduncle 2—15 mm long,

(1—)2.5—5(—6.5) cm diam., rather dense to very lax with branches obscure or longest

0.3—3 cm long, 20—many-flowered. Pedicel 0—0.5 mm long. Sepals (6—)7—10 x (2—

)3—4(—5.5) mm, not or slightly elongating in fruit, cream to beige or yellow-brown in

flower, pink to (red-)purple in fruit, glabrous to rather densely 0.2—0.5(—1) mm long

colourless- or red-hairy mostly on keel, tip hardly to distinctly hooded. Petals 5.5—8.0

x 2.5-4.2 mm, slightly longer than sepals, white, glabrous, margin entire or minutely

fringed. Stamens with filaments c. 8 mm long, white, in upper 2—3 mm with many 3-4

mm long hairs; anthers 1-1.6 « 0.8—1.4 mm, yellow, rarely pink, red, blue or purple,

thecae opening by a longitudinal slit. Capsule 6—8.5 « 3.5—4.5 mm, narrowly obovoid,

1.5(—2) mm shorter to 1.5(—3) mm longer than sepals, pink to red-purple, glabrous or

apex to upper half sparsely 0.1—0.5(—1) mm long hairy; valves free; apex depressed,

lobes absent; style remnant 0.5 x 0.2 mm, finally deciduous. Seeds 2 per locule, rarely

1 abortive, aril orange.

Distribution. Myanmar (Tenasserim), Thailand (SW: Kanchanaburi, Petchaburi;

Peninsular Thailand: Ranong, Nakhon Si Thammarat, Trang, Satun, Pattani,

Narathiwat), Peninsular Malaysia (Kedah: Langkawi and mainland, Penang: Pulau

Pinang, Kelantan, Terengganu, Perak, Pahang, Selangor, Negeri Sembilan, Malacca,

Johor), Singapore, Sumatera (Aceh, Sumatera Utara, Sumatera Barat: Kepulauan

Mentawai, Riau, Kepulauan Riau, Jambi, Palembang: Bangka Island, Lampung), Java

(West: Bogor, Bandung, Priangan), Philippines (Palawan: Puerto Princesa, Brooke’s

Point), Borneo (Brunei; Sabah: West Coast, Interior, Sandakan, Tawau; Sarawak:

Kuching, Samarahan, Sri Aman, Betong, Bintulu, Miri, Limbang, Kapit; Kalimantan:

W, C, SE, E), Sulawesi (Central).

Ecology. Lowland dipterocarp, peat swamp or montane primary, disturbed or old

secondary, evergreen or (semi-)deciduous forest, in deep to partly shaded or open

vegetation in valley, along streams, in forest clearings or margins or on roadsides, on

rather dry to wet, rocky or clayey (volcanic) soils with humus absent or present, on

sandstone, shale, granite or limestone (terra rossa). Altitude: 0—-1500 m asl.

Notes. 1. This species is variable with respect to sheath indumentum and width, and

leaf blade indumentum and dimensions. Whether this is ecologically induced variation

Amischotolype in Asia 95

could not be established from herbarium material alone.

2. Cleistogamy is suspected in at least one collection (Duistermaat 350,

Peninsular Malaysia, Selangor, Gombak), in which the anthers shed their pollen in

tightly closed flowers.

3. In Singapore, there is only one record from the wild (Ridley 6433, 1892,

Bukit Timah, Fern Valley). Despite serious attempts we were unable to refind the

population. Ridley (1907, 1924) and Keng (1987) cited Wallich 8977b for Singapore,

but as seen from microfiche (K) this is A. gracilis; although Wallich 8977a is A.

marginata, this specimen is from Penang. At present, the species is growing in the rain

forest of the Singapore Botanic Gardens, but the origin of that population is at best

uncertain.

4. Although the species has been reported for Lao P.D.R. and Vietnam (Cherfils

1937; Ho, 1993), I doubt its presence there. Material identified by Cherfils as F.

marginatus are A. glabrata (Balansa 4100, Eberhardt 3277) and A. divaricata (Thorel

s.n.). | have not seen any material of A. marginata from Lao P.D.R. nor Vietnam. The

description by Cherfils (1937) includes long rhizomes, and inflorescence in the part

of the stem where the leaf blade has been shed, i-e., the rhizome. These characters

agree with A. marginata. However, the illustrations and other parts of the description

conflict with this species. Both cited sources illustrate a plant with the inflorescence

on the node that also bears a leaf with the leaf blade still attached, 1.e., flowering on

the erect stem. Also, sepals are reported as 4.5—9 mm long and thecae as opening in

the apical region, whereas in fact sepals are at least 6 mm long and the thecae open

by a longitudinal slit only. It seems that a description of A. marginata was copied,

supplemented with characters from material of at least one other species. (See also

note 2 under Amischotolype.)

5. A. laxiflora and A. monosperma are quite similar in habit with inflorescences

on the rhizome, but differ in width of sepals, and colour of indumentum.

Specimens examined: MYANMAR. Tenasserim: Tavoy distr, Aug 1961, J. Keenan et al. 972

(E); Tawer distr, Yebyu Township, E of Kanbauk, Oct 1998, J.F. Maxwell 98-1092 (L).

THAILAND. SW: Kanchanaburi, Between Khao Yai and Khao Ngi Yai. E of Sangkhla, Apr

1968, C.F. van Beusekom & C. Phengkhlai 411 (E, L, P); Petchaburi, Kaeng Krachan NP, Khao

Phanoen Thung, km no 30, May 2008, P. Phonsena et al. 6006 (L). Peninsular Thailand:

Pattani, Bachaw, Jul 1923, A.F.G. Kerr 7181 (P); Trang, Chawrj, Apr 1928, A.EG. Kerr 15185

(L); Ranong, Khao Saideng. near Ranong, May 1968, C.F. van Beusekom & C. Phengkhlai

542 (E, P); Nakhon Si Thammarat, Khao Luang area, May 1968, C.F. van Beusekom & C.

Phengkhlai 846 (E, P); Waterfall at Yala, Oct 1970, Ch. Charoenphol et al. 4132 (KLU):

Khao Khlong Yang at Khao Phra Mi, Jul 1972, K. Larsen et al. 30655 (KLU, L): Satun, Kuan

Kah Long, Tong Sgui subdistr. nr Nam Rah vill, Aug 1984, J. Maxwell 84-100 (SINU):

Nakorksitaprarat, Lansagah, Gahrome Falls, Khao Luang Nat.Park. Apr 1985. J. Maxwell 85-

400 (SINU); Trang, Kao Chong Nat.Park, Jul 1987, Samsuri Ahmad 35 (SINU): Narathiwat,

Khao Chana, Kaluwotai, Narathiwat, Sep 1995, C. Nivomdham 1047 (P).

PENINSULAR MALAYSIA. Kedah: Kedah Peak, Jun 1893, H.N. Ridley s.n. (SING):

Langkawi, Gunung Raya Forest Reserve, Mar 1939, Md.Nor 31378 (KEP): Gunong Bongon

Forest Reserve, Jun 1941, G.H. Spare 3786 (SING): Langkawi, G.Raya, ascent from SE near

96 Gard. Bull. Singapore 64(1) 2012

the base, Aug 1972, B.C. Stone 10932 (KLU); Langkawi, N base of G.Raya, due S of Kg.Sg.

Itau, Nov 1979, B.C. Stone 14314 (KLU); Langkawi, Dec 1990, Khairuddin Hj. Itam 3 (KEP)

& 5 (KEP); Kuala Muda, Gng Jerai FR, Titi Hayun, compt 20,, Jul 2006, K. Jmin et al. FRI

50747 (L, KEP, SING). Penang: Pulau Pinang, NV. Wallich 8977a (BM, E, K); Pulau Betong,

Aug 1884, C. Curtis 1948 (A) (SING) & 1948 (B) (SING); Pulau Betong, 189x, Guard (Ridley)

2 (BM); Governement Hill, May 1898, C. Curtis s.n. (A) (SING) & s.n. (B) (SING). Kelantan:

Ulu Sg. Lebir, 1.5 mi S of Sg. Ternya, Sep 1967, B.C. Stone 7317 (KEP, KLU, L). Terengganu:

Kemaman, Bukit Kajang, Nov 1935, E.J.H. Corner s.n. (SING); Kemaman, Sungai Nipah

FR, Cptmt 56, Sungai Ayam, May 2006, 4.R. Rafidah et al. FRI 51647 (KEP, L, SING); Hulu

Terengganu, Ulu Telemong FR, Jul 2006, S.N. Phoon et al. FRI 51960 (KEP); Hulu Terengganu,

Petuang FR, Jul 2006, S.N. Phoon et al. FRI 51977 (KEP); Hulu Terengganu, Tasik Kenyir, Sg.

Cendana, Aug 2007, K. Imin et al. FRI 58538 (L, KEP). Perak: Taiping Hill, Dec 1892, H.N.

Ridley 11422 (K, SING); Maxwell’s Hill, 1892, H.N. Ridley s.n. (SING); Taiping Hills, Feb

1904, H.N. Ridley s.n. (SING); Temango, Jul 1904, H.N. Ridley s.n. (SING); Lenggong, Aug

1904, H.N. Ridley 14375 (BM, SING); Gunong Bujang Malaka, Aug 1959, Kadim bin Tassim

457 (SING); Bukit Kinta FR, Apr 1987, R. Kiew 2588 (KEP); Hulu Perak, Temenggor FR,

Sungai Halong, Aug 1993, M. Abdul Latiff et al. 3943 (KEP, L); Belum, near Sg. Halong, Sep

1993, 1. M. Turner & J.W.H. Yong 31 (SINU); Hulu Perak, Belum FR, Sungai Kejar, Sg. Kejar,

Jun 2007, S. Svahida Emiza & Kueh H.L. FRI 55186 (KEP, L). Pahang: Pulau Tioman, Aug

1889, HN. Ridley s.n. (SING); Telom, Nov 1901, H.N. Ridley 13813 (BM, K, SING); Telom,

Nov 1908, H.N. Ridley 13812 (BM, K, SING); Pulau Tioman, Juara Bay, Tawa Valley, Jun 1915,

HM. Burkill s.n. (SING); 7th mile Raub Road, Jul 1924, G.A. Best SF 14128 (SING); Pulau

Tioman, G.Kajang, Ayer Surin, May 1927, M.R. Henderson SF 18443 (SING); Tembeling,

Jul 1929,M.R. Henderson SF 21782 (SING); Sungei Lemoi, Sep 1931, Jaamat 28172 (KEP,

SING); Lipis, Sungai Serambun, Oct 1931, Osman 27998 (KEP); Tekek to Joara, May 1974,

B.C. Stone 11984 (KLU); Genting Highlands Road, Jun 1974, B.C. Stone & T. Hattink 12013

(KLU); Fraser’s Hill, Nov 1976, H. Keng et al. s.n. (SINU); Fraser’s Hill, Bishop’s Trail, Apr

1992, R. Kiew & S. Anthonysamy 3525 (SING); Pulau Tioman, G. Kajang, Saw L.G. et al. FRI

40117 (KEP); Pulau Tioman, 1km W of Kg Juara, track to Kg Tekek, Jul 2005, H. Duistermaat

394 (L, SING); Maran, Jengka FR, Hutan Lipur Jebak Puyuh, Gua kap, Oct 2008, 1.4. Mohd

Hairul et al. FRI 60076 (KEP). Selangor: Telok Reserve, Klang, Sep 1918, H.M. Burkill SF

3135 (SING); Talik reserve, Klang, Mar 1921, H.M. Burkill SF 7034 (SING); Hulu Langat,

Sungai Lalang Forest Reserve, Mar 1930, C.F) Symington 24057 (KEP); Kuala Lumpur, Univ.

Malaya, fern plot, Khantijah 102 (KLU); Telok Khantijah 107 (KLU); Telok FR (nr Klang?),

Nov 1971, K. Jong 9025 (KLU); N of Kuala Lumpur, JIn Gombak km36(KL), Jun 2005,

Hf. Duistermaat 350 (L, SING); Kepong, FRIM, forest trail behind herbarium, Jun 2005, H.

Duistermaat 356 (L, SING); Gombak, Kanching FR, along waterfall main trail, Sep 2006, H.

Duistermaat et al. FRI 51935 (KEP, L); Gombak, Kanching FR, lower part of quartz ridge,

Mar 2007, Chew M.Y. et al. FRI 53665 (L). Negeri Sembilan: Gunong Angsi, near bungalows,

Nov 1923, Md.Nur SF 11645 (SING); Pasoh FR, Oct 1987, J. V. LaFrankie 2529 (KEP); Pasoh

FR. nr Simpang Pertang, Jul 1989, 4. Gentry & J.V. LaFrankie 66983 (L); Gunung Angsi, Jun

2005, H. Duistermaat 342 (L, SING); Jelebu, Berembun FR, Jeram Toi , forest trail, Apr 2008,

Yao TL. et al. FRI 57936 (KEP, L); Jelebu, G.Telapak Buruk, Peak nr Telekom/Celcom tower,

Apr 2008, S.N. Phoon et al. FRI 60645 (L). Malacca: Maingay 1713 (L); Bukit Kedongong,

May 1890, R. Derry 602 (SING); Bt. Ledondong, May 1890, H.N. Ridley 602 (BM); Jasin, Bkt

Senggeh FR, Cpt.3, Jul 2008, YM. Chan FRI 64724 (KEP). Johor: Tanjong Kopang, 1892,

HN. Ridley s.n. (SING); Gunung Panti, Dec 1892, H.N. Ridley s.n. (SING); Kukub., 1909,

HN. Ridley 14168 (SING); Sg.Endau near Kg.Peta, Aug 1973, Khantijah 104 (KLU).

Amischotolype in Asia 97

SINGAPORE. Bukit Timah, 1892, H.N. Ridley s.n. (K); Bukit Timah, nr Fern Valley, 1892,

HN. Ridley 6433 (SING); Bukit Timah, Jun 1948, J. Sinclair 4840 (E); SBG, rainforest, nr

‘rattan-entry’ at palm valley, Apr 2005, H. Duistermaat 329 (L, SING); SBG, Liane Rd nr path

to greenhouses, Aug 2005, H. Duistermaat 395 (L, SING).

SUMATERA. Aceh: Ketambe, valley of Lau Alas, May 1972, W.J.J.O. de Wilde & B.E.E.

de Wilde-Duyfjes 12419 (L); Gng Leuser NP, Alas river valley, Jul 1979, W..J.O. de Wilde

& B.E.E. de Wilde-Duyfjes 18480 (L). Sumatera Utara: Sibolangit, May 1917, J.4. Lérzing

5125 (L); Sibolangit NR, Dec 1927, J.A. Lorzing 12746 (L); Bukit Lawang-Bohorok-Langkat,

Feb 1973, R. Soedarsono 309 (K, L). Sumatera Barat: Afd. Agam Brani, Jun 1918, H.A.B.

Buennemeyer 3344 (L); G.Sago, Jul 1918, H.A.B. Buennemeyer 3649 (L); Padang, nr Ajer

mantjur, Aug 1878, O. Beccari 828 (L); Kepulauan Mentawai, Island of Siberut, Aug 1924,

C. Boden Kloss SF 14497 (SING); Sipora Isl., Sioban omgeving, Oct 1924, [boet 417 (SING).

Riau: Indragiri, Sg.Gaung, Sep 1940, E. Polak 546 (L). Kepulauan Riau, Anambas/Natoena,

Boengoeran eiland, Apr 1928, C.G.G.J/. van Steenis 1397 (L). Jambi: Pesisip river, nr Dusun

Pasirmayang, Dec 1980, N.A.P. Franken & M. Roos 363 (L). Bangka-Belitung: Bangka Island,

Soengai Selan, Nov 1917, H.A.B. Buennemeyer 2106 (L). Lampung, Gunung Sugi, Jan 1908,

J. Elbert s.n. (L); Kalianidoea, Dec 1921, CM. Sloet 435 (L); Mt. Tanggamus, Apr 1968, ©.

Jacobs 8117 (KEP, L, SING).

JAVA. C.L. Blume 2047 (L). Jawa Barat: Bogor, C.L. Blume s.n. (L); Preanger, nr Gadongang

(Bandong), H. Zollinger 540, p.p. (E, L); Priangan, Tjampaka near Tjidadap, Dec 1937, P.

Buwalda 3436 (L); road Pelabuhan Ratu-Kiara-Lengkong,, Sep 1985, E.F: de Vogel 7649 (L).

PHILIPPINES. Palawan: Brooke’s Point, Addison Peak, S slope, Jan 1991, D.D. Soejarto &

O. Fernando 7356 (L); Munic. Brooke’s Point, S range of Mantlingajan Rang, Jun 1994, D.D.

Soejarto et al. 8330 (L); Mt. Cleopatra Ranges, Co. Calabayog, Puerto Princesa, Jun 1996, E.

Reynoso & R. Majaducon PPI 24356 (L); lrawan forest, Puerto Princesa, Aug 1996, E. Romero

et al. PPI 38462 (L).

BORNEO. Brunei: Belait, Labi, Kampong Tenajor, Sep 1988, A. Mohd. Haslani 17 (KEP, L,

SING); Tutong District, track from Panchong to Benutang, Oct 1989, L.L. Forman & J.B.J.

Blewett 983 (L, SING); Belait District, Labi, path to Rampayoh waterfall, Oct 1989, L.L.

Forman & J.B.J. Blewett 1046 (L, SING); Belait, Labi, Rampayoh, Jul 1993, J. Cowley 101 (L)

& 1] (L); Temburong, Amo, Apan, Sungai Baki, Jul 1993, /. Cowley 87 (K). Sabah: West Coast:

Mount Kinabalu, Penibukan, Jan 1933, J. & M.S. Clemens 30889 (BM); Kota Belud, above

Sg.Dahobang S-side nr Kinabalu Park, Sep 1965, 4. Kanis 51478 (L, SAN); Ranau, Sungai

Nabutan, Mar 1982, Ahan Gibot SAN 94552 (L, SAN); Kota Belud, NW side Mt Kinabalu,

May 1984, J.H. Beaman et al. 9754 (L); just outside Kinabalu Park along rd to Marai-Parai,

Sep 1993, J. Nais et al. SP 5321 (KEP); Ranau, Kampung Bundu Tuhan. Lugas, Kg.Himba,

Sep 1993, Doinis Soibeh 423 (SAN); Ranau, Kampong Takutan, Apr 1995, 7: Lomudin 221

(KEP); Ranau District, Kampung Takutan, Skm dari kpg, Jun 1995, 7. Lomudin 378 (KEP);

Kota Belud, Kinabalu P, Sayap, Lumangis trail, Jun 2000, A.D. Poulsen et al. 1619 (SAN);

Interior: Tenom, Oct 1961, 4. Buntar SAN 27364 (L); Keningau, Keningau, Ulu sg.Matud,

Jun 1987, K. Fidilis SAN 119541 (SAN); Keningau, Nabawan, via Tenom, Feb 1989, Jay E.P.

et al. 89-0409 (SING); Tambunan, Sunsuron Km 54 JInTambunan/Penampang, Aug 1989, K.

Fidilis SAN 127872 (SAN); Sipitang, Melakis FR, Oct 1989, S. Awang Amin S 114838 (K, KEP,

L); Sandakan: Lahad Datu, Timbun Mata Is. F.R., Aug 1940, H.G. Keith 34 (SING) & 55500

(KEP); Lahad Datu, S.Pangaruanan, K. Bay, Dec 1959, D. Brand 20061 (L, SING); Labuk

Sugut, Beluran, Sg.Baba, Mar 1980, Aban Gibot SAN 90037 (L, SAN); Sepilok FR, Jun 1981,

T. Sato & S. Dewol SAN 93160 (SAN); Kinabatangan, Keruak FR, Feb 1985, K. Amin et al.

SAN 108160 (SAN); Lahad Datu, Ulu Sg.Segama, Dec 1985, G.C.G. Argent et al. SAN 108225

98 Gard. Bull. Singapore 64(1) 2012

(E, SAN); Lahad Datu, N-trail Danum Valley, Jul 1986, Lee et al. SAN 109039 (SAN); Lahad

Datu, Danum Valley, Mar 1987, K. Amin & R. Joseph SAN 110020 (L, SAN); Sepilok FR, May

1987, Jurimin Ebin SAN 116477 (SAN); Lahad Datu, Kunak, km 79 Tawau/Lahad Datu road,

Jun 1987, E. Suali & Muhamad SAN 117141 (SAN); Lahad Datu, Danum Valley FC, Begonia

Cliff, Feb 1992, E.J.F. Campbell-Gasis 234 (E, SAN); Kinabatangan District, Bukit Garam,

mile 5, Nov 1994, Reza Azmi 68 (SAN); Lahad Datu district, Madai-BaturongFR, Madai Hill,

Jun 1996, S.P. Lim et al. 656 (SAN); Kinabatangan D, Sukau, PangiFR, Temangong Besar Hi,

Sep 1996, S.P. Lim et al.1266 (SAN); Lahad Datu, Tempadong, Borneo Marble Quarry, Aug

1999, R. Kiew 4793 (KEP); Lahad Datu District, Tabin Wildlife Res.near headquarter, Mar

2000, K.H. Kjeldsen 94 (SAN); Lahad Datu, Madai Baturong, Jul 2000, S. Dewol SAN 133042

(KEP, SING); Tawau: Elphinstone prov., 1922-23, A.D.E. Elmer 20864 (L); Luasong Centre

old logging area, Ikea plant, Jul 2001, 1. Postar et al. SAN 144083 (KEP, L, SAN); Tawau Hills

NP, path to Bombalai, May 2005, J.J. Vermeulen in H. Duistermaat 358 (L, SING). Sarawak:

Kuching: Matang, Mar 1955, W.M.A. Brooke 9756 (BM, L); Lundu, Gunong Gading, Sep

1955, J.W. Purseglove & Md.Shah 4522 (K, L, SING); Lundu, Gunong Pueh, Sep 1955, J. W.

Purseglove 4816 (L, SING); Lundu, G.Gading, Sep 1974, James et al. S 35087 (L); Kuching,

Bau, Gg.Meraja, May 2002, S.B. Raymond et al. 3134 (SING); Samarahan: Serian, Bukit

Rawang, Tebakang area, Apr 1983, Dyg. Awa & I. Paie S 45586B (L); Serian, Bukit Majung,

Tebakang, Dec 1998, P. Kessler 221 (L); Sri Aman: Simanggang, Oct 1955, W. Brooke 10730

(BM, L, SING); Ulu Sungai Silantek Kiri, Aug 1980, 7. Paie S 42477 (KEP, L); Betong: up

Layar river, Grajih, Mar 1954, W.M.A. Brooke 8241] (BM, L); Bintulu: Tatau, Bukit Naong, Apr

1995, J. Lai et al. S 71032 (ct; KEP); Ulu Sg.Kakus, Bkt Sarang, Batu Anyi, Jan 2005, S. Lee

S 94752 (KEP); Tatau, Ulu Merirai, Gua Naga, Jul 2005, S. Julia S 95712 (KEP, SING); Tatau,

Bukit Sarang group, Bukit Anyi SE-side, Aug 2005, J.J. Vermeulen in H. Duistermaat 400 (L,

SING); Miri: Baram district, E...River, Mar 1894, C. Hose 370 (BM); Bukit Subis area, Aug

1963, H.P. Fuchs 21281 (L); S slopes of Gg.Subis, nr Sekaloh river, Nov 1966, Sonny Tan & E.

Wright S 27258 (L, SING); Bakam, Oct 1983, A.P. Abang Mohtar S 47201] (KEP, L); Marudi,

Mulu National Park, May 1985, 4.P. Abang Mohtar & I. Othman S 49692 (KEP); Limbang:

Tg.Long Amok, Sg.Ensungei, Sep 1960, R. George et al. S 42352 (L); Kapit: Teneong, Sep

1954, W.M.A. Brooke 912] (BM, L). Kalimantan W: Landak, Ngabang, 1851, /.E. Tevsmann

11551 (BO); ridge SW of G.Bentuang Jul 1989, JS. Burley et al. 3340 (L); Gn Palung NP

Oct 1996, 77G. Laman et al. 97 (E, L). Kalimantan C: Kahayan, Mar 1988, J.S. Burley et

al. 472 (KEP, L, SING). Kalimantan SE: PW. Korthals s.n. (L); Tanahboemboe, G.Mangis,

N v Batoelitjin, Oct 1928, D.F- van Slooten 2184 (L); km 14 Bandjermasin-Martapoera, Oct

1939, B. Polak 503 (L). Kalimantan E: Boelongan, 1914, LZ. Rutten 112 (U); A. Kostermans

5418 (L); E.Kutei, G.Tepian Lobang, Jun 1951, Loa Djanan, W of Samarinda, Apr 1952, A.

Kostermans 6415 (L); Balikpapan distr, Mentawir region, Jul 1954, A. Kostermans 9708 (L);

Kutei Reserve N. of Samarinda, Mentoko, May 1970, Soegeng Reksodihardjo 11 (L); Teluk

Sanggan, W of Malinau, Jul 1981, K. Jwatsuki et al. B 7213 (L); Wartono Kadri, km 3.5 along

Samboja-Semai Rd, Mar 2005, C. Bernard 3 (L); Meratus, Mului, Nov 2005, N. Raes et al.

698 (L).

SULAWESI. Central: Lore Lindu NP, Feb 2008, D. Cicurra 483 (L).

15. Amischotolype mollissima (Blume) Hassk. (Fig. 10D)

Amischotolype mollissima (Blume) Hassk., Flora 46 (1863) 392. — Campelia

mollissima Blume, Enum. P|. Javae (1827) 7; Kunth, Enum. Pl. 4 (1843) 109; Moritz,

Amischotolype in Asia 99

Syst. Verz. (1845-46) 93; Zollinger, Syst. Verz. (1854) 65: Miquel, Fl. Ned. Ind. 3

(1855) 546; Fl. Ned. Ind., Eerste Biyv. (1860) 609. — Forrestia mollis Hassk., Flora 47

(1864) 628, nom. illeg. (see note 1); Commelin. Ind. (1870) 84; Clarke in A.Candolle &

C.Candolle, Monogr. Phan. 3 (1881) 236, p.p.; Hooker f., Fl. Brit. India 6 (1894) 383,

p.p. — Forrestia mollis Hassk. var. blumeana Hassk., Flora 47 (1864) 628; Commelin.

Ind. (1870) 84. — Forrestia mollissima (Blume) Koord., Excursionsfl. Java 1 (1911)

282. — Forrestia hispida A.Rich. var. mollis (Hassk.) Hallier f., Nova Guinea 8 (1913)

906; Beih. Bot. Centralbl. 34 (1916) 47. — Forrestia mollissima Koord. forma typica,

Backer, Handb. Fl. Java I, 3 (1924) 33. — Forrestia mollissima (Blume) Koord.

forma mollissima Backer & Bakh.f., Fl. Java 3 (1968) 15. LECTOTYPE (designated

here): Blume s.n., Java (L, barcode L 0041655). Other syntypes: Blume s.n., in sylvis

montosis Javae insulae (L: 3 sheets); Reinwardt s.n., Tjampea (L).

Forrestia mollis Hassk. var. korthalsii Hassk., Flora 47 (1864) 628; Commelin. Ind.

(1870) 86. TYPE: Korthals s.n., Java (L).

Forrestia mollis Hassk. var. teysmannii Hassk., Flora 47 (1864) 628; Commelin. Ind.

(1870) 86. TYPE: Teysmann s.n., Sumatera, Lubu alang (BO).

Forrestia bicolor Hallier f., Bull. Herb. Boissier 6 (1898) 360, t. XI. SYNTYPES:

Burck s.n., s.d. “Sumatra’ (BO, n.v.), Jaheri s.n., 1895 ‘Deli, Tandjung Gunung’ (BO?,

n.v.).

Stem rhizome absent, erect part 100—160 cm long, basally sometimes with some stilt

roots, simple; internodes glabrous or rarely moderately 0.2 mm long hairy. Sheath

10-19 mm diam., glabrous or with one to few lines of (0.5—)3-4.5 mm long, white

to yellow or brownish hairs, mouth ciliate. Leaf blade 27-44 = 5.7—10.6 cm, 3.4-5.9

times as long as wide, base very gradually to abruptly narrowed into an indistinct

winged pseudopetiole to 4 cm long and 1.5—2 mm wide; lower surface moderately

to densely (0.1—)0.5—1 mm long colourless- to white-hairy or rarely glabrous, upper

surface glabrous (rarely with 0.2 mm long hairs near margin); submarginal hairs on

upper surface, (0.5—)1—1.5 mm long, dense, yellow. Inflorescence on erect stem,

sessile, 3.2-5.7 cm diam., dense with branches obscure, 20—many-flowered. Pedicel

2-3 mm long. Sepals (8.5—)12—15.5 x 2.5-4.5 mm, probably slightly elongating in

fruit, light violet, lilac or red, sparsely 0.5—1.5 mm long ciliate margins and keel or

rarely glabrous, tip hooded. Petals 9-10 = 3 mm, c. as long as sepals, colour unknown,

glabrous, margin minutely fringed at apex. Stamens filament 8—10 mm long, colour

unknown, upper 2 mm with many 3 mm long hairs; anthers 1—1.3 = 0.6—-0.7 mm,

colour unknown, thecae opening by a longitudinal slit. Capsule 6-10 = 4.5-6 mm,

obovoid, shorter than sepals by 3-6 mm, whitish or dark carmine, apex to upper half

0.2-1 mm long hairy; valves free to fused for 1/3 of length; apex depressed, lobes

0.5—1.3 mm long; style remnant 0.5—1.2 x 0.3 mm, finally deciduous or persistent.

Seeds 2 per locule, aril orange.

100 Gard. Bull. Singapore 64(1) 2012

Distribution. Sumatera (Aceh, Sumatera Utara, Sumatera Barat), Java (W: Bogor,

Preanger, Tjisaroea; C: Banjoemas; E: Besuk1).

Ecology. Primary and secondary (riverine) rain forest, along stream, in gorge, on

roadside, on alluvial, fertile soil. Altitude: 60—-600(—1420) m asl.

Notes. 1. Hasskarl (1864), when he transferred species of Amischotolype to Forrestia,

choose for A. mollissima the illegitimate name F’.. mollis (illegal because an autonym

is required in the absence of a preoccupied earlier homonym in the recipient genus),

based on Tradescantia mollis Reinw. in sched., with Campelia mollissima Blume

appearing in the synonymy. Clarke (1881) and Hooker (1894) included in their concept

specimens of A. gracilis, A. hirsuta and A. barbarossa, these only later recognised as

distinct separate species. Ridley (1907, 1924: swb F. mollis) and Turner (1997: sub A.

mollissima) used the name for A. barbarossa exclusively.

2. The species is limited in its distribution to Java and Sumatera. In Java it is

not known above 600 m asl, whereas in Sumatera it occurs up to 1420 m asl. Beaman &

Beaman (1998) cited A. mollissima for Borneo (Daim Andau 566), but this collection

is A. hirsuta.

3. A. mollissima 1s very similar to A. hispida in size and shape of the leaf

blade, inflorescence and flowers. However, A. hispida differs in having long rhizomes,

capsules without apical lobes and with valves fused for 1/4 tol/2 of the length. The two

species are geographically disjunct: A. hispida ranges from Borneo and the Philippines

eastward to New Guinea.

4. [have not seen the type of Forrestia bicolor (it 1s unclear if Jaheri’s collection

in the Hortus Bogoriense ended as a collection in any herbarium). Nevertheless, there

can be no doubt that this is a synonym for A. mollissima, because the combination in

the description of sepals 13-15 mm long, inflorescence opposite the leaves (leaves

thus present), and the capsule apex distinctly depressed (“...apice umbilico profundo

trquetro praedita...”) is unique for this species.

5. The only other species that have apical lobes on the capsule are A. barbarossa

(absent to 0.3 mm long; see there, note 1), 4. we/zeniana (0.5 mm long; sheath, sepals

and capsule glabrous) and A. /obata (1—1.5 mm long; distinctly pseudopetiolate leaf

blades, and capsules exceeding the calyx and bristly-hairy).

Specimens examined: SUMATERA. P.W. Korthals s.n. (L). Aceh: Ketambe, valley of Lau

Alas, May 1972, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 12036 (L); Gn Leuser NR,

Ketambe, valley of Lau Alas, May 1974, H.D. Rijksen 050574 (L); Gng Leuser NR, c. 5 km

S of Ketambe, Alas riv, Jun 1979, W..J.O. de Wilde & B.E.E. de Wilde-Duyfjes 18035(A) (L);

foot of Mt Biak Mentelang Koacane, Feb 1980, Afandi Ma’Roef 371 (L); Kloet NR, along

Krung (=river) Lembang, Jul 1985, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 19908 (L).

Sumatera Utara: Karohoogvlakte bij Lingga, Jan 1919, J.A. Lérzing 6272 (L); Gajo, 1921,

G.C.E. van Daalen 367 (L); Sebolangit, Bukit Senia.., Aug 1921, Md. Nur SF 7365 (K, SING);

Sibolangit NR, Dec 1927, J.A. Lérzing 12747 (L); Biang valley nr Sarinembah, Karo plat nr

Lingga, Oct 1928, J.A. Lérzing 14432 (L); Kaban Djahe, May 1939, 4.H. Batten Pooll s.n.

(SING). Sumatera Barat: Lubu Abang, /.E. Teysmann s.n. (BO); above Telug Kabung, Jun

Amischotolype in Asia 101

1953, J. van Borssum Waalkes 1507 (K, L).

JAVA. C.L. Blume s.n. (L); P.W. Korthals s.n. (L); C.G.C. Reinwardt s.n. (L); J.C. van Hasselt

s.n. (L); J.C. Ploem s.n. (L); Mumutean s.n. (L). Jawa Barat: Salak C.L. Blume 2249 (L):

Vogelberg, Tjampea, J.C. van Hasselt s.n. (L); nr Bogor, foot of mnt Salak, 1821, C.L. Blume

s.n. (L); Tjisaroea, Oct 1870, R.H.C.C. Scheffer s.n. (L); Buitenzorg, Bandongan, Apr 1900,

Soegandiredja 198 (L); G.Karang bij Klappa-Noenggal, NO van Buitenzorg, Dec 1912,

Backer 5968 (L); Bogor, Preanger, Tjisaroea; Preanger, Tjibadak o/d Halimoen, Jan 1920, R.C.

Bakhuizen van den Brink Sr. 3150 (L); Br. Tjiomas, Aug 1922, R.C. Bakhuizen van den Brink

Sr. 1678 (U); Preanger: Z van Radjamandala, May 1924, C.A. Wisse 1089 (L); boschweg van

Tjipannas naar Wijnkoopsbaai, Jun 1932, A. Kloos 371] (L); Bogor, Tjiburial, Feb 1951, Nedi

& Idjan 398 (K, L). Jawah Tengah: Banjoemas, Pasir Salam bij Madjenang, Jan 1915, C.A.

Backer 18755 (L); Jawa Timur: Besuki, Tjoeraman, S.H. Koorders 20664b (L).

16. Amischotolype monosperma (C.B.Clarke) I.M.Turner (Fig. 1OE, 17A—C & 18)

Amischotolype monosperma (C.B.Clarke) I.M.Turner, Novon 6 (1996) 221; Gard.

Bull. Sing. 47 (1997) 516. — Forrestia monosperma C.B.Clarke in Hallier f., Bull.

Herb. Boissier 6 (1898) 359; Ridley, Mat. Fl. Malay. Penins. 2 (1907) 124; Fl. Malay.

Penins. 4 (1924) 360. LECTOTYPE (designated here, see note 1): Curtis s.n., 1890,

Waterloo Estate, Perak; ‘this specimen flowered in the Botanic Garden of Penang’

(SING).

Amischotolype irritans auct., sensu Beaman & Beaman, The Plants of Mount Kinabalu

3 (1998) 109. non Ridley.

Forrestia marginata auct., sensu Hasskarl, Flora 47 (1864) 630, p.p.; Commelin. Ind.

(1870) 90, p.p.; Hooker f., Fl. Br. India 6 (1894) 383, p.p.

Tradescantia sp., Griffith, Not. Pl. Asiat. 3 (1851) 235.

non Forrestia monosperma auct., sensu Cherfils in Lecomte, Fl. Gen. Indo-Chine

6 (1937) 855. — Amischotolype mollissima (Blume) Hassk. var. monosperma

(C.B.Clarke) P.H.H6, Cayco Vietnam 3, | (1993) 478, illus. (“Amischolotype’), nom.

inval., no basionym. (see note 3, and note 2 under Amischotolype).

Stem ascending from creeping and branched, c. 70 cm long rhizome; erect part c. 60

cm long, simple; internodes sparsely to densely 0.1—0.5 mm long red- to redbrown-

hairy. Sheath (7—)12-30 mm diam., reddish green, many (rarely only at front or in

lower half) dense lines of 2.5-4 mm long, orange to orange-brown or red hairs, in

between glabrous to sparsely 0.1 mm long hairy, mouth glabrous to ciliate. Leaf blade

(21—)30-50(-68) = (6.5—)9-16 cm, 2.3—5.1 times as long as wide, bullate, lower

surface usually red to purple, base gradually to rather abruptly narrowed into a winged

pseudopetiole up to 13 cm long and 3—7 mm wide; both surfaces glabrous or rarely

sparsely 0.1mm long hairy; submarginal hairs on upper surface, 0.2—0.7 mm long,

102 Gard. Bull. Singapore 64(1) 2012

rather sparse, colourless or yellow-brown. Inflorescence on rhizome and around knee,

peduncle (0O—)3—20 mm long, (2.5—)3.5—9 cm diam., dense to very lax with branches

obscure or longest 0.54.5 cm long, 20—many-flowered. Pedicel absent. Sepals (8—)

9-14(-18) x 2-4.5 mm, not elongating in fruit, red or reddish brown to purple (in

flower and fruit), rarely creamy yellow at upper half, margins and keel sparsely to

rather densely (0.5—)1—2 mm long orange- to redbrown-hairy, in between glabrous

or 0.3—0.5 mm long hairy, tip hooded. Petals 8—9.5 x 2.5—3 mm, slightly shorter than

sepals, white, glabrous, margin at apex minutely fringed. Stamens with filaments

9—12.5 mm long, white, in upper 2-3 mm with 3.54 mm long hairs; anthers 1—1.5 x

0.9—-1 mm, white, thecae opening by a longitudinal slit. Capsule 7.5—8.5 « 3.54 mm,

obovoid, shorter than sepals by 1.5—5(—8) mm, magenta red or purple, apex to upper

half sparsely to densely 1—2 mm long orange- to red-brown-hairy; valves unknown;

apex depressed, lobes absent; style remnant 0.7 x 0.3 mm, deciduous. Seeds 2 per

locule, 1 rarely abortive, aril orange.

Dicabaner Myanmar (7.v., see note 2), Thailand (Peninsular Thailand: Yala),

Peninsular Malaysia (Kedah, Kelantan, Terengganu, Perak, Pahang, Selangor, Negeri

Sembilan), Borneo (Sabah: West Coast, Interior).

Ecology. Primary forest or forest margin, often on stream or river banks, often in wet

areas and on limestone. Altitude: 30—1000(—1500) m asl.

Notes. 1. The species required lectotypification (see Turner 1996). The original

description mentioned the following locality (Clarke in Hallier 1898): “Aus Perak in

den botan. Garten zu Pinang und von hier in die Garten zu Singapur, Buitenzorg u. s.

w. eingeftihrt.” This is very similar to the annotation on the label of Curtis s.n., 1890,

Waterloo Estate, Perak (SING), which is therefore selected here as the lectotype.

2. The authors who mentioned 7radescantia sp. of Griffth (1851a), including

Hasskarl (1864), Hooker (1894), and Ridley (1907), placed it in A. marginata.

Unfortunately, I did not see Griffith’s material (Merg. Herb. 185, August 1834, in

sylvis Kyouklag), collected in the Mergui Archipelago, Myanmar. Griffith (1851b)

provides no illustration of any Commelinaceae species. However, Griffith’s (1851a)

description (‘infra florifero foliorumque vaginarum reliquis obtectis’—flowering on

the part covered with remnants of leaf sheaths, and ‘sepalis rubro-aurantiaceus dorso

pilosa’—sepals with red-golden hairs on the back) can only refer to 4. monosperma.

It is the only known record for Myanmar.

3. Although the species has been reported for Lao P.D.R. and Vietnam (Cherfils

1937; Ho 1993, Fig. 8427), I doubt its presence there. I have not seen material from

Lao P.D.R. or Vietnam referable to 4. monosperma. Material determined by Cherfils

as F’. monosperma 1s A. divaricata (Harmand 1920, Pierre s.n., and Poilane 186, all

P). Further, the description by Cherfils (1937) cannot refer to 4. monosperma: the leaf

blades are too narrow, the sheaths have white appressed short hairs, and the sepals

are sparsely hairy near the apex only. Lastly, the illustration in Ho (1993) shows a

plant flowering at the erect stem with leaf blades present, and capsules that are much

Amischotolype in Asia 103

longer than the sepals. Both the description of Cherfils (1937) and the illustration in

Ho (1993) could refer to 4. divaricata.

4. In Peninsular Malaysia normally found at low altitudes, only once (in

Perak) at 960 m asl.; in Thailand found only once, and probably in mountainous area.

In Borneo, on the other hand, only found at around 1000 up to 1500 m alt.

Specimens examined: THAILAND. Peninsular Thailand: Yala, waterfall, Oct 1970, Ch.

Charoenphol et al. 4129 (KLU, L, P, SING).

PENINSULAR MALAYSIA. Kedah: Langkawi, Jan 1991, Khairuddin Hj. Itam 9 (KEP).

Kelantan: Gua Batu Goh, 0.5 mile S of Gua Musang, 7.C. Whitmore FRI 4043 (KEP. L):

Gua Panjang, Ulu Kelantan, Aug 1962, UNESCO 334 (SING); Gua Ninik, Gua Musang, May

1990, R. Kiew & S. Anthonysamy 2963 (SING); Gua Musang, Gua Batu Boh, Mar 2008. Chew

M.Y. et al. FRI 60153 (KEP). Terengganu: Ulu Brang, Jul 1937, L. Moysey & Kiah SF 33855

(SING); Hulu Terengganu, Petuang FR, Jul 2006, S.V. Phoon & R. Kiew FRI 51981 (KEP).

Perak: Waterloo Coffee Estate, 18xx, C. Curtis s.n. (SING): Waterloo Perak, May 1890, C.

Curtis s.n. (SING); Cenderiang, 1961, P.M. Castle-Smith 30 (K); Tapah Hills, Along Sg.

Who, Jul 1966, FS.P. Ng FRI 1350 (KEP, SING); Belum, nr Sungei Remai, Apr 1994, 7. M.

Turner 94-49 (SINU). Pahang: Tahan River, 1891, HN. Ridley s.n. (K, SING); Tembeling, Jul

1929, M.R. Henderson SF 21796 (SING); at Gap of Fraser’s Hill Rd., Jul 1960, ME.D. Poore

230 (K, KLU); Gua Musang, Ulu Kelantan, Aug 1962, UNESCO 428 (SING); Ulu Sg. Sat near

Kuala Kelepah, Jul 1970, Md.Shah & Md.Noor 1883 (KEP): Taman Negara, nr Kuala Tahan.,

Gua Rimau limes outcrop, Aug 1971, Chin S.C. 1275A (KLU):; Taman Negara, path from Kuala

Tahan to Batu Linga, Aug 1972, B.C. Stone 10860 (KLU); base of Bt.Terseh, Taman Negara,

Aug 1973, B.C. Stone 11582 (KLU); Batu Luas, Taman Negara, Oct 1984, R. Kiew 1352 (KEP,

SING): Kampong Woh, A. Vogel 960236 (L): Jerantut, Taman Negara, Camp Melantai, May

2008, Chew M.¥. et al. FRI 60260 (L). Selangor: Gua Batu, Apr 1889, H.N. Ridley 4599

(K, SING); Ulu Gombak, Oct 1937, Md. Nur SF 34206 (L, SING); Gombak, Oct 1979, S.

Anthony SA 262 (KEP): Gombak Valley, stream to waterfall, Oct 1981, R. Kiew 1096 (SING);

Kuala Lumpur, Rimba IImu Garden (KLU), May 2005, H. Duistermaat 341 (L, SING); N. of

Kuala Lumpur, Jin Gombak km 36 (KL), Jun 2005, H. Duistermaat 349 (L, SING). Negeri

Sembilan: Pasoh FR, Oct 1987, J. V. LaFrankie 2528 (KEP).

BORNEO. Sabah: West Coast: Kota Belud, Mount Kinabalu, Dallas El., 1931/32, 7. & MS.

Clemens 2614] (BM, K, L); Kota Belud, Mount Kinabalu, Penibukan, Kinitaki R, Jun 1933.

Clemens 50405 (BM); Interior: Tambunan, Tontolob Liwan, Oct 1989, Soinin et al. SAN 124990

(KEP); Tambunan, Crocker Range NP, Oct 1999, A.D. Poulsen et al. 1514 (KEP); Tambunan,

Trusmadi, Loloposon (Gua), Jul 2005, J.J. Vermeulen in H. Duistermaat 397 (L, SING).

17. Amischotolype parvifructa Duist. sp. nov. (Fig. 5 & 13D)

Amischotolype divaricatae similis in foliis supra pilis submarginalibus brevibus

capsulis calyce superantibus, sed vaginis pilis 0.1—0.2 longis, capsulis 7—9 mm longis

albis ad roseis pilis 0.5—1 mm longis satis mollis differt. TYPUS: Chew WL. 837,

October 1963, Peninsular Malaysia, Pahang, Cameron Highlands, Bukit Ruil, 1800 m

a.s.l. (holo SING; iso L).

Stem ascending from a rhizome up to c. 60 cm long; erect part 90-150 cm long and

scrambling or not, simple, at base with few stilt roots; internodes moderately 0.1—

104 Gard. Bull. Singapore 64(1) 2012

0.3 mm long hairy. Sheath 8-10 mm diam., moderately to densely 0.1—0.2 mm long

hairy, mouth ciliate. Leaf blade 18—31.5 = 5.5—8.0 cm, 3+4.4 times as long as wide,

smooth, base rather abruptly narrowed into 3-4 cm long and 1—2 mm wide winged

pseudopetiole; both surfaces glabrous or moderately 0.1 mm long hairy; submarginal

hairs on upper surface, 0.2—0.4 mm long, (rather) sparse, white. Inflorescence on

erect stem, sessile, 2—2.7 cm diam., dense with branches obscure, 10—15-flowered.

Pedicel 0—0.5 mm long. Sepals 7.3—8.5 x 3—3.5 mm, not elongating in fruit, green or

whitish turning purplish, sparsely to rather densely 0.3—0.6 mm long hairy especially

in upper half, tip hooded. Petals c. 7 x 2 mm, slightly shorter than sepals, pale green on

outer surface, white on inner surface, glabrous, margin entire. Stamens with filaments

hairy, anthers white, otherwise unknown. Capsule 7-9 x 4.5—6 mm, obovoid, longer

than sepals by 3—5 mm, white to pink, moderately to rather densely 0.5—1 mm long

hairy; valves fused for 1/4 of length or finally free; apex distinctly depressed, lobes

absent; style remnant 0.5—1.5 x 0.2—0.4 mm, persistent. Seeds 2 per locule, aril colour

unknown.

Distribution. Peninsular Malaysia (Pahang: Cameron Highlands).

Ecology. Hill or (lower) montane forest. Altitude: 1350—1980 m asl.

Notes. The epithet refers to the rather small fruits compared to 4. divaricata with which

it could be confused because of the sometimes scrambling habit, the short submarginal

hairs on the upper surface of the leaf blade and the capsule exceeding the calyx.

Differences between them are mentioned in the key (lead 23). Also, the distribution

is disjunct: A. divaricata has been collected from Myanmar, Cambodia, Lao P.D.R.,

Thailand and Sumatera whereas the present species is endemic to Cameron Highlands

in Peninsular Malaysia.

Specimens examined: PENINSULAR MALAYSIA. Pahang: Ulu Telom, Aug 1931, Jaamat

27271 (KEP); Cameron Highlands: below Robinsons Falls, Oct 1961, H.M. Burkill 2852

(SING); Bukit Ruil, Oct 1963, Chew W.L. 837 (L, SING), Aug 1975, Rao etal. s.n. (SINU); path

no 4 (nr Tanah Rata?), Apr 1980, S. Anthony SA 305 (KEP); Tanah Rata, Robinson waterfall,

Sep 1985, A. Latiff et al. 867 (L), Mar 2010, R. Kiew et al. FRI 70474 (KEP).

18. Amischotolype pedicellata Duist. sp. nov. (Fig. 6)

Amischotolype hispida arte similis, capsulis calyce superantibus, pilis longioribus

setosioribus, capsulae valvis fere ad basin liberis differt. TYPUS: Hallier B2600,

March 1894, Indonesia, S. Kalimantan, S of Banjarmasin, Liang Gagang (holo L).

Forrestia hispida A.Rich. var. typica auct., sensu Hallier f., Beih. Bot. Centralbl. 34

(1916) 46, p.p. (see note); non Hallier f. (= A. hispida).

Forrestia hispida auct., sensu Merrill, J. Straits Branch Roy. Asiat. Soc. (1921) 113,

p.p. (see note); non A. Rich. (= A. hispida).

105

Amischotolype in Asia

WALSMIT SACHS 2012

Fig. 5. Amischotolype parvifructa Duist. A. Habit. B. Fruit with persistent sepals. Drawing by

A. Walsmit Sachs, from Chew W.L. 837 (L).

106 Gard. Bull. Singapore 64(1) 2012

Stem ascending from short to more than 20 cm long rhizome; erect part 70-200 cm

long, simple; internodes glabrous or very sparsely to moderately 0.2—0.3 mm long

hairy. Sheath 9-15 mm diam., with several lines of 3-4 mm long, yellow or brown

hairs, mouth ciliate. Leaf blade 29-47 =< 5—9.1 cm, 4.3—6.2 times as long as wide,

base gradually to rather abruptly narrowed into indistinct to 5 cm long and 2-3 mm

wide winged pseudopetiole; both surfaces glabrous or sparsely 0.2 mm long hairy;

submarginal hairs on upper surface, 1.5—2.5 mm long, dense, yellow. Inflorescence

on erect stem, sessile, 2-4 cm diam., dense with branches obscure to 3 mm long,

20-30-flowered. Pedicel 2-5 mm long. Sepals 8.5—9.5 x 2.5—4 mm, not elongating in

fruit, pink to purple or magenta, glabrous or (very) sparsely 0.2—1.5 mm long ciliate

keel, tip hooded. Petals c. 8 x 2 mm, white or yellow (in bud), glabrous, margin entire.

Stamens filament c. 7 mm long, colour unknown, upper | mm with 0.5—1.5 mm long

hairs; anthers 0.9—1 x 0.4—0.5 mm, colour unknown, thecae opening by a longitudinal

slit. Capsule 7—8.5(—10) x 4—6 mm, ovoid, equalling or exceeding sepals by up to 1.5

mm, bright lilac to purple, at least at apex rather sparsely to moderately 1—2.5 mm long

bristle-hairy, hairs yellow to brown; valves almost free; apex depressed, lobes absent;

style remnant 0.7—1 = 0.3—0.5 mm long, persistent. Seeds 2 per locule, aril orange.

Distribution. Borneo (Sabah: West Coast, see note; Sarawak: Limbang; Kalimantan:

WiC, S):

Ecology. Primary or newly disturbed mixed dipterocarp rain forest, hill forest,

streambanks , on (red) clayey soil. Altitude: 120—150(—1500) m asl.

Notes. The name refers to the presence of a pedicel of at least 2 mm long, a character

this species shares with six more species. The species most closely resembles 4.

hispida, but is different in the capsule exceeding the calyx and with longer and more

bristly hairs, and with valves free almost to the base. It is endemic to Borneo, growing

in lowland rain forest and only on Mount Kinabalu up to 1500 m altitude.

Specimens examined: BORNEO. Sabah: West Coast: Kota Belud, Mount Kinabalu, Penibukan,

Feb 1933, J. & M.S. Clemens s.n. (BM) & 31295 (BM). Sarawak: Limbang, 1/2 day from

Lawas, 1954-55, W.M.A. Brooke 10227 (L). Kalimantan W: Kabupaten Sanggau, Feb 1994,

W. de Jong 743 (L); Serawai, Jan 1995, A.C. Church et al. 1544 (L). Kalimantan C: Bukit

Raya & Upper Katinabatangan R., Upper Samba River, Nov 1982, J.P. Mogea & W.J.J.O. de

Wilde 3681 (L); Tumbang Tubus, Jan 1983, J.) Veldkamp 8193 (L); Sintang, Apr 1994, A.C.

Church et al. 929 (L); Sintang HPH km70-72, SW along logging road, Apr 1994, U.W. Mahyar

et al. 929 (SING). Kalimantan S: S of Banjarmasin, Liang Gagang, Mar 1894, H. Hallier

B2600 (L).

19. Amischotolype rostrata (Hassk.) Duist. comb. nov. (Fig. 10F)

Forrestia rostrata Hassk., Flora 47 (1864) 631, Commel. Ind. (1870) 94; Hallier f., Beih.

Bot. Centralbl. 34 (1916) 48; Merrill, J. Straits Branch Roy. Asiat. Soc. (1921) 113. —

Amischotolype in Asia 107

WALSMILT SACHS 2013

Fig. 6. Amischotolype pedicellata Duist. A. Habit. B. Fruit with persistent sepals. Drawing

by A. Walsmit Sachs, A. from H. Hallier B2600 (L), B. from J.P. Mogea & W.J.J.O. de Wilde

3681 (L).

108 Gard. Bull. Singapore 64(1) 2012

Campelia glabrata auct. 8 acuminatissima Hassk. in Miquel, Pl. Jungh. (1852/53) 155.

— Campelia sp., Zollinger, Syst. Verz. (1854) 65. — Forrestia marginata (Blume)

Hassk. var. B rostrata (Hassk.) C.B.Clarke in A.Candolle & C.Candolle, Monogr.

Phan. 3 (1881) 237; Hooker f., Fl. Brit. India 6 (1894) 384. LECTOTYPE (designated

here): Junghuhn s.n., s.d., Java, Medinie, Ungarang (L, barcode L 0928865). Other

syntypes: Blume s.n., s.d., Java, Preanger, near Lembang (L); Zollinger 503 (B, n.v.; P

in hb Franqueville, 7.v).

Forrestia rostrata Hassk. var. zollingeri Hassk., Commel. Ind. (1870) 96. SYNTYPES:

Zollinger 540 p.p., Java, Bantam prope Tjikoya in silvula humida (G-DC, B, n.v.);

Anon. s.n. Java (L in hb Hasskarl).

Forrestia porrecta Ridl., J. Malayan Branch Roy. Asiat. Soc. 1 (1923) 102.

LECTOTYPE (designated here; see note 3): Ridley s.n., February 1921, Sumatera,

Berastagi Hill Woods (K).

Forrestia distans Ridl., J. Bot., Suppl. 63 (1925) 124; Backer & Bakhuizen van den

Brink, Fl. Java 3 (1968) 15 (see note 4). TYPE: Forbes 612, Java, Mt. Waringin, 5000

ft (holo BM, n.v.).

Stem ascending from a rhizome up to 200 cm long; erect part up to 200 cm long,

scrambling, simple or branched; internodes glabrous or rarely sparsely 0.2—0.3 mm

long hairy. Sheath 3—9 mm diam., 0.2—0.5 mm long hairy, rarely glabrous, mouth

ciliate. Leaf blade 13-23 =< 3-6 cm, 3.54.9 times as long as wide, base gradually

to rather abruptly narrowed into an indistinct or 0.5—1 cm long and 0.5—2 mm wide

winged pseudopetiole; both surfaces glabrous; submarginal hairs on lower surface

or rarely absent, 0.2—0.5(—1.5) mm long, rather sparse. Inflorescence on erect stem,

sessile, 1.3—2.8 cm diam., (rather) dense with branches obscure, 7—10-flowered.

Pedicel absent. Sepals (5—)6—8.5 = 24.5 mm, not elongating in fruit, colour unknown,

glabrous or moderately 0.2—0.7(—1) mm long colourless- to yellowish-hairy in upper

half, tip slightly hooded. Petals 6—7 < 2—2.5 mm, as long or longer than sepals, white,

glabrous, margin at apex minutely fringed. Stamens filament 8—10 mm long, white, in

upper 1—2 mm with 2—2.5 mm long hairs; anthers 0.9—1.6 x 0.6—0.9 mm, white, thecae

opening by a apical pore. Capsule 8.5—11 x (5.5—)7—8.5 mm, ovoid, longer than sepals

by 3-7 mm, colour unknown, entirely to upper 1/3 sparsely to rather densely 0.8—1.5

mm long hairy (longest hairs at apex), hairs colourless to yellowish; valves fused for

(1/4 to) 1/2 of length; apex obtuse, lobes absent; style remnant (0.3—)1(—2.5) x 0.2-0.4

mm, persistent. Seeds 2 per locule, aril colour unknown.

Distribution. Sumatera (Berastagi), Java (West: Priangan, Bogor, Tasikmalaya;

Central: Semarang; East: Madiun, Pasuruan, Besuki), Maluku (unspecified).

Ecology. Moist secondary forest. Altitude: (200—)700—1600 m asl.

Amischotolype in Asia 109

Notes. 1. This species is very similar to the genus Porandra, with the thecae

opening by an apical pore and the scrambling habit often with branched stems (see

Introduction). However, species of Porandra differ from A. rostrata in having purple

anthers, P. ramosa has teardrop-shaped anthers, while P. scandens has sheaths with

a line of at least 1 mm long hairs and submarginal hairs on the upper surface of the

leaf blade. Species of Porandra are hitherto recorded only from mainland Asia, but I

have identified specimens from Sumatera as P. scandens (Lesger 206, Lorzing 6273,

van Steenis 6201). Amischotolype rostrata, on the other hand, is known only from

the Indonesian Archipelago, mainly Java, with only two collections from Sumatera

and one from Maluku. Hasskarl (1864) mentioned 4. rostrata for the island of Pinang

(Peninsular Malaysia). However, the collection he cited (Wallich 8977a) contains a

specimen of A. marginata and a specimen that cannot be identified with certainty from

microfiche (see 4. hookeri, note). Amischotolype rostrata has also been mentioned

for India (Griffith 5486: East Bengal, Mishmee Mountains; Clarke 1881), but this

specimen is P. ramosa.

2. Although I have not seen the types of F. rostrata var. zollingeri, Hasskarl

(1870) described it as different from the species in having more robust stems which

are creeping and then ascending and branched, and larger leaves (but he also describes

smaller ones that are present as well). This variation 1s part of a continuum that does

not merit distinction at any level.

3. Ridley (1923) did not cite specimens when he described F. porrecta, stating

only that it was found in the Berastagi hill woods (Sumatera). His description mentions

the typical long creeping stems lying prostrate on the ground or creeping up the trunk

of a tree, and the remarkably small heads of flowers. I have seen only one collection

of Ridley (K) that fits his description and the locality. | have designated this as the

lectotype of F. porrecta.

4. Ridley (1925) mentioned that F. distans is most nearly allied to F. porrecta

which is a synonym for A. rostrata (see note 3), whereas Backer & Bakhuizen van den

Brink (1968) thought it is identical with F. mollissima forma glabrata (= A. glabrata).

The description of F. distans nicely fits that of A. rostrata. Therefore it is accepted as

a synonym of the present species. Differences between 4. glabrata and A. rostrata

include the number of flowers per inflorescence, the length of the sepals (absolute, and

relative to the length of the capsule), and the opening of the thecae.

Specimens examined: SUMATERA. Sumatera Utara: Berastagi, (1918), C.D. Ouwehand 372

(L); Berastagi Hill Woods, Feb 1921, H.N. Ridley s.n. (K, L).

JAVA. Anon. (in Hb. Hasskarl) s.n. A (L); FW. Junghuhn s.n. (L). Jawa Barat: prope Lembang

haud procula a Bandong, C.L. Blume s.n. (L); Tjibodas, J.G. Boerlage s.n. (L); Limbang, P.W.

Korthals s.n. (L); M.Tjiseroea, Oct 1870, R.H.C.C. Scheffer s.n. (L); m.Gedeh, 1895, H. Hallier

533 / 51 (L); inter Tjibodas et Tjibeureum, 1895, H. Hallier 533/71 (L); Preanger, Tjibodas,

1915, Sapiin 2037A (L, U); Preanger, Tjidadap, Tjibeber, Apr 1917, R.C. Bakhuizen van den

Brink Sr. 2174 (L); Preanger, Pendjaloe, Jul 1917, S.H. Koorders & A. Koorders-Schumacher

44374B (L); Tjibodas, Pantjuran emas, May 1948, D.R. Pleyte 262 (L); Dago (waterval), Feb

1949, S.M. Popta 723/71 (L); Gede, dal van de Tji Bodas, Mar 1950, S.J. van Ooststroom

13206 (L); Jawa Tengah: Medinie, Ungarang, n-helling, FW. Junghuhn s.n. (L); Jawa Timur:

110 Gard. Bull. Singapore 64(1) 2012

Madioen, Ngebel, May 1896, S.H. Koorders 23266B (K, L); Pasuruan, Tangkil, Zuidergeb.,

Jun 1896, S.H. Koorders 23384B (L) & 23887B (L); Besuki, Pantjoer, Idjen, Aug 1897, S.H.

Koorders 28526B (L); Tengger, 1912, J.P. Mousset 431 (L); O.Wilis, 1914, J.A. Lérzing 914

(Gi).

MALUKU. 1859-60, G.H. de Vriese & J.E. Teijsmann s.n. (L).

20. Amischotolype sphagnorrhiza Cowley (Fig. 10G)

Amischotolype sphagnorrhiza Cowley in Coode et al., Checklist Fl. Plants & Gymn.

Brunei Dar. (1996) 352 (‘sphagnorhiza’), 434; Cowley & Furness, Kew Bull. 52

(1997) 465. TYPE: Coode et al. 7062 (holo K, photo; iso BRUN, n.v.)

Stem ascending from a rhizome 19-50 cm long; erect part 10-30 cm long, with up

to 45 cm long aerial roots with Sphagnum-like rootlets, simple; internodes unknown.

Sheath 8-10 mm diam., glabrous, mouth glabrous. Leaf blade 18-25 =< 4.5-7.3

cm, 2.64.8 times as long as wide, smooth, lower surface pale purple or not, base

gradually narrowed into indistinct pseudopetiole; both surfaces glabrous; submarginal

hairs absent. Inflorescence at knee, peduncle 540 mm long, up to 7 cm long, lax

with branches up to 4.5 cm long, 10—25-flowered, at base with Sphagnum-like roots.

Pedicel 2-6 mm long. Sepals 2—10 = 1—5 mm, elongating in fruit, white to greenish-

cream or pink to purple, sparsely to moderately 0.5 mm long redbrown bristly hairy,

tip hooded. Petals 2—6 = ? mm, as long or shorter than sepals, white, glabrous, margin

entire. Stamens with filaments 1-7 mm long, white, upper part with 0.5—1 mm long,

yellow hairs; anthers 1—1.2 x 0.5 mm, yellow, thecae opening by a longitudinal slit.

Capsule 9.5—24 x 5-10 mm, ovoid, longer than sepals by c. 8 mm, deep red-purple or

purple-brown, 0.5 mm long orange-hairy; valves free; apex slightly depressed, lobes

absent; style remnant absent. Seeds 2 per locule, aril unknown.

Distribution. Borneo (Brunei: Belait; Sarawak: Sri Aman: Sri Aman).

Ecology. Low or relatively open mixed dipterocarp forest, river valleys, on yellow

sandy clay. Altitude: 15—230 m asl.

Notes. Avery peculiar species, in habit resembling Palisota spp. from Africa. According

to Cowley & Furness (1997), based on pollen and flower morphology, it is best placed

in Amischotolype. Apart from the Sphagnum-like rootlets, it is very different from the

other species of the genus in the very short erect part of the stem and the very lax and

elongated inflorescence (although those of A. marginata can be quite lax and elongated

as well). Future studies as to the generic placement of this species requires a molecular

analysis, and chromosome count. Described as an endemic for Brunei, it appears to be

present in Sarawak as well (//ias Paie S 42752).

Amischotolype in Asia 111

Specimens examined: BORNEO. Brunei: Tutong, Rambai, Ulu Tutong, Bukit Bahak, M/E.

Coode et al. 7062 (K); Belait, Labi, Sungai Rampayoh, Jan 1994, D.W. Kirkup & M.JE.

Coode 808 (KEP. L, SING); Belait, Bukit Sawat, Jalan Merangking-Buau, Sg.Sindum, May

1997, Suhaili Hj. Zinin et al. BRUN 18020 (SING). Sarawak: Sri Aman, 99 mile Kuching

Simanggang Road, Sep, 1980, . Paie S 42752 (L).

21. Amischotolype strigosa Duist. sp. nov. (Fig. 7)

Amischotolype irritanti similis in bracteis sepalis, petalis cum pilis rubris spinosis, sed

caule repentiore, internodiis vaginisque pubescentibus, foliis capsulisque minoribus,

capsulis pubescentioribus differt. TYPUS: R. Soedarsono 349, February 1973,

Indonesia, Sumatera, N, Bukit Lawang-Bohorok-Langkat (holo SING; iso L).

Stem ascending from short rhizome; erect part c. 100 cm long, simple: internodes

moderately 1.5 mm long yellow-hairy. Sheath 7-10 mm diam., with many lines of

4 mm long, yellow hairs, mouth ciliate. Leaf blade 17-18 = 2.5—3.6 cm, 5—6.8 times

as long as wide, base gradually narrowed into indistinct pseudopetiole; both surfaces

moderately 1.5—3 mm long yellow-hairy; submarginal hairs on upper surface, 1.5

mm long, moderately dense, yellow. Inflorescence on erect stem, sessile, 1.8—2.2

cm diam., very dense with branches obscure, 20—30-flowered. Pedicel 2-5 mm long.

Sepals 7-8 x 1.5—2 mm in flower, elongating to 9 mm long in fruit, colour unknown,

rather densely 2 mm long spiny (dark) red-hairy, tip not hooded. Petals c. 8 x 1.5 mm,

as long to slightly longer than sepals, white, dorsal surface apically with 2.5 mm long

spot of 1—1.5 mm long red spiny hairs, margin entire. Stamens with filaments c. 10

mm long, colour unknown, glabrous; anthers c. 1.2 x 0.7 mm, colour unknown, thecae

opening by a longitudinal slit. Capsule c. 5 x 3.5 mm, obovoid, shorter than sepals

by c. 2 mm, colour unknown, upper 1/3 1 mm long, red spiny-hairy; valves free; apex

slightly depressed, lobes absent; style remnant | x 0.1—0.2 mm, deciduous. Seeds 2 per

locule, aril colour unknown.

Distribution. Sumatera (Sumatera Utara: between Medan and Gunung Leuser).

Ecology. Marshy places in recently logged-over forest. Altitude: 50-200 m asl.

Notes. The epithet refers to the red spiny hairs on the bracts, sepals and petals,

characters it shares with A. irritans. However, the internodes and sheaths are hairy,

the leaf blades and capsules are smaller, and the capsules are more hairy, characters

sufficient to warrant recognition of a new species.

Specimens examined: SUMATERA. Sumatera Utara: Bukit Lawang-Bohorok-Langkat, Feb

1973, R. Soedarsono 349 (L, SING): Sikundur F.R., 75 km NW of Medan, Besitang River, Aug

1979, W.I.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 19490 (KLU, L).

We Gard. Bull. Singapore 64(1) 2012

WALSAIT SACHS 2012

Fig. 7. Amischotolype strigosa Duist. A. Habit. B. Immature fruit with persistent sepals.

Drawing by A. Walsmit Sachs, from R. Soedarsono 349 (L).

Amischotolype in Asia ts,

22. Amischotolype welzeniana Duist. sp. nov. (Fig. 8)

A congeneribus in vaginis laminis sepalis capsulis glabris differt. TYPUS: Maxwell

85-955, Peninsular Thailand, Trang, Khao Chong National Park, 300 m alt., somewhat

disturbed thicket along trail in primary evergreen forest (holo L; iso SINU, E).

Stem ascending from creeping rhizome; erect part 50-100 cm long, simple; internodes

glabrous. Sheath 7-11 mm diam., glabrous, mouth ciliate. Leaf blade 26-30 = 6-8.1

cm, 3.5-4.7 times as long as wide, lower surface green or tinged with purple, base very

gradually narrowed into indistinct pseudopetiole; both surfaces glabrous; submarginal

hairs on upper surface and margin, 1—1.5 mm long, dense, yellow. Inflorescence on

erect stem, sessile, 1.5—3.4 cm diam., dense with branches obscure, 1 5—many-flowered.

Pedicel absent. Sepals 7-13 =< 2.5—3 mm, probably elongating in fruit, whitish (in

flower) turning deep violet (in fruit), glabrous, tip hooded. Petals c. 8 x 2 mm, slightly

longer than sepals, white, glabrous, margins unknown. Stamens with filaments cream-

coloured, at apex with few 0.5 mm long hairs; anthers 2—2.2 x 0.4 mm, cream-coloured,

thecae opening by a longitudinal slit. Capsule (immature) 9 x 4 mm, ovoid, shorter

than sepals by c. 3.5 mm, mauve or green tinted with purple, glabrous; valves fused for

4/5 of length; apex depressed, lobes 0.5 mm long; style remnant | x 0.2 mm, probably

persistent. Seeds 2 per locule, aril orange.

Distribution. Myanmar (Tenasserim: Tavoy), Thailand (C: Krung Thep Maha Nakon

(Bangkok); Peninsular Thailand: Trang, Surat Thani).

Ecology. Evergreen forest, on rocks, or in somewhat disturbed thickets along trails.

Altitude: 210-300 m asl.

Notes. The species has been named after P.C. van Welzen on the occasion of his

inauguration to professor (7ropical Plantbiogeography) on 19 Jan 2009. The plant

is strikingly glabrous throughout (sheaths, leaf blades, sepals and capsules) which

distinguishes it from all other species in the genus. The filaments are almost glabrous.

I have not seen fully ripened fruits; the largest are those of the type specimen and they

still seem to be immature. It is possible that ripe fruits are larger than described here.

Specimens examined: MYANMAR. Tenasserim: Tavoy, Aug 1961, J. Keenan et al. 785 (E).

THAILAND. C: Krung Thep Maha Nakon (Bangkok), Tung Song, Jul 1929, Rahil 96 (L):

Peninsular Thailand: Surat Thani, Kao Nawng, Aug 1927, 4.hG. Kerr 13225 (L); Trang,

Khao Chong National Park, Oct 1985, J.F. Maxwell 85-955 (E, L, SINU).

114 Gard. Bull. Singapore 64(1) 2012

1 cm 1 mm WALSAIT SACHS 2012

Fig. 8. Amischotolype welzeniana Duist. A. Habit. B. Immature fruit with persistent sepals.

Drawing by A. Walsmit Sachs, from J.P. Maxwell 85-955 (L).

Amischotolype in Asia 115

Fig. 9. Flower details. A. Amischotolype glabrata Hassk. Based on F.W. Junghuhn s.n. (L). B.

Amischotolype gracilis (Ridl.) 1.M.Turner. Based on H. Duistermaat 348 (L). C. Amischotolype

griffithii (C.B.Clarke) I.M.Turner. Based on J.S. Burley et al. 1237 (L). D. Amischotolype

hirsuta (Hallier f.) Duist. Based on H. Winkler 2617 (L). E. Amischotolype hispida (A.Rich.)

D.Y.Hong. Based on Lesson ex J.K. Hasskarl s.n. (L). F. Amischotolype hookeri (Hassk.)

H.Hara. Based on J.D. Hooker & T. Thompson s.n. (L). G. Amischotolype irritans (Ridl.)

1.M.Turner. Based on H. Duistermaat 346 (L). All drawings by Walsmit Sachs.

116 Gard. Bull. Singapore 64(1) 2012

Fig. 10. Flower details. A. Amischotolype laxiflora (Merr.) Faden. Based on Hallier f B

3001 (L). B. Amischotolype leiocarpa (Hallier f.) Duist. Based on H. Hallier B3002 (L). C.

Amischotolype marginata (Blume) Hassk. Based on C.L. Blume s.n. (L). D. Amischotolype

mollissima (Blume) Hassk. Based on C.L. Blume s.n. (L). E. Amischotolype monosperma

(C.B.Clarke) I.M.Turner. Based on Md. Nur SF 34206 (L). F. Amischotolype rostrata (Hassk.)

Duist. Based on F.W. Junghuhn s.n. (L). G. Amischotolype sphagnorrhiza Cowley. Based on

D.W. Kirkup & M.J.E. Coode 808 (L). All drawings by Walsmit Sachs.

{mischotolype in Asia 117

Fig. 11. A-B. Amischotolype barbarossa Duist. A. Stem with orange-hairy sheaths and leaf

blades. B. Inflorescence with ripe, red-hairy fruits. From FRI 57441. C—D. Amischotolype

glabrata Hassk. C. Inflorescence with opened fruit showing three seeds each with orange aril.

D. Inflorescence with fully opened flower. From Duistermaat 392. E-F. Amischotolype gracilis

(Ridl.) I.M.Turner. E. Plant with silvery-white variegated leaves. From Duistermaat 353. F.

Inflorescence with opened flower. From Duistermaat 336. All photographs by J.J. Vermeulen,

except E by H. Duistermaat.

118 Gard. Bull. Singapore 64(1) 2012

Fig. 12. Amischotolype gracilis (Ridl.) 1.M.Turner. A. Stem with glabrous sheaths and leaf

blades with red submarginal hairs on the upper surface. Photograph by J.J. Vermeulen from

Duistermaat 328. B. Erect stem with leaves and at each node an inflorescence perforating the

sheath at its base. Photograph by: H. Duistermaat from Duistermaat 328. C. Inflorescence with

ripe purple fruit. Photograph by: H. Duistermaat from Duistermaat 348. D. Inflorescence with

ripe white fruits. Photograph by Chew, M.Y., FRIM from FRI 55597.

Amischotolype in Asia 119

Fig. 13. A. Amischotolype hirsuta (Hallier f.) Duist. Erect stem with hairy leaves and fruiting

inflorescence. Photograph by J.J. Vermeulen from Duistermaat 398. B. Amischotolype hispida

(A. Rich.) D.Y.Hong. Erect leafy stem with inflorescences. Photograph by Serena Lee from

S. Lee 426. C. Amischotolype irritans (Ridl.) 1.M.Turner. Erect stem with glabrous sheaths

and red-spiny-hairy inflorescence. Photograph by H. Duistermaat from Duistermaat 346. D.

Amischotolype parvifructa Duist. Erect stem with leaves and inflorescence with white fruits.

Photograph by H. Duistermaat from FRI 70474.

120 Gard. Bull. Singapore 64(1) 2012

Fig. 14. A-B. Amischotolype laxiflora (Merr.) Faden. A. Stem with hairy sheaths. B.

Inflorescence on leafless rhizome. Photographs by J.J. Vermeulen from Duistermaat 401. C—D.

{mischotolype marginata (Blume) Hassk. C. Erect stem with leaves in the background, the

leafless rhizome with purple inflorescences in the foreground, both indicated with a white

arrow. Photograph by H. Duistermaat from Duistermaat 342. D. Stem with hairy sheaths.

Photograph by J.J. Vermeulen from Duistermaat 400.

Amischotolype in Asia 12]

Fig. 15. Amischotolype marginata (Blume) Hassk. A. Plicate leaf blade with indistinct

submarginal hairs. From Duistermaat 329. B. Inflorescence on rhizome with open flowers. From

Duistermaat 395. C. Ripening purple inflorescences at the nodes of the leafless rhizome. From

FRI 58591. D. Ripening inflorescence. From Duistermaat 395. E. Few-flowered inflorescences

on long peduncles. From Duistermaat 400. F. Fully opened flower showing purple anthers.

From Duistermaat 358. All photographs by J.J. Vermeulen, except A by H. Duistermaat.

122

Gard. Bull. Singapore 64(1) 2012

Fig. 16. Amischotolype marginata (Blume) Hassk. A. Flowering white to pale purple

inflorescences on rhizome. Photograph by Paul K.F. Leong from Duistermaat 329. B.

Inflorescence with purple sepals, white fruits and one opened fruit showing three orange arils

covering the seeds. Photograph by J.J. Vermeulen from Duistermaat 358.

—

(oe)

Amischotolype in Asia

Fig. 17. Amischotolype monosperma (C.B.Clarke) I.M.Turner. A. ‘Rosette’ of leaves at the

apex of the stem. From HBL (Vogel) 960236. B. Inflorescence with ripening fruits. From

Duistermaat 397. C. Young inflorescence with first flower opened. From Duistermaat 397. All

photographs by J.J. Vermeulen.

124 Gard. Bull. Singapore 64(1) 2012

Fig. 18. Amischotolype monosperma (C.B.Clarke) I.M.Turner. Detail of inflorescence showing

opened flower with cream-coloured red-hairy sepals. Photograph by André Schuiteman from

HBL (Vogel) 960236.

Amischotolype in Asia 15

Nomina dubiae

A. mollissima (Blume) Hassk. var. marginata (Blume) R.S.Rao, Maharashtra Vidnyan

Mandir, Patrika 6, 2 (1971) 53, nom. inval.

A. mollissima (Blume) var. glabrata (Hassk.) R.S.Rao, ibid.

Notes. Rao (1971) made two new combinations for the flora of India: A. mollissima

(Blume) Hassk. var. marginata (Blume) R.S.Rao, and A. mollissima (Blume) var.

glabrata (Hassk.) R.S.Rao. He cited Campelia mollissima Blume for both, and

Forrestia mollis Hassk. only for var. marginata. It may therefore be concluded that he

considered this var. marginata the typical variety. Because an autonym is required, the

combination A. mollissima var. marginata (Blume) R.S.Rao is invalid. Furthermore,

both combinations should be considered as nomina confusa. For A. mollissima var.

marginata, C. marginata, C. mollissima and F. hispida are cited, but these are not

known to occur in India. For A. mollissima var. glabrata, both C. glabrata and

Forrestia hookeri Hassk. are cited, which are here considered as distinct species both

occurring in India. Rao (1971) neither gives descriptions, nor does he cite material for

his varieties. However, the combination Amischotolype mollissima (Blume) Hassk.

var. glabrata (Hassk.) R.S.Rao has been made validly.

ACKNOWLEDGEMENTS. The study is based on material from the following herbaria: K,

KEP, KLU, L, P, SAN, SING, SINU, U, and (digital) images of material in B (RGpert 2000),

BO, E and K. The help of the following people is greatly acknowledged: first of all, Dr. Ruth

Kiew, former Keeper of SING, who encouraged me to start on the group; and staff members

of SING (Ms. Serena Lee, Mr. Paul K.F. Leong, Mr. Gwee Aik Teck, Mr. Samsuri Ahmad and

Dr. Jana Leong-Skorniékova), who accepted my company during their field trips which gave

me the opportunity to make some nice collections of Amischotolype and to make very valuable

field observations. Paul helped me enormously to maintain a vital living collection in the SBG

greenhouse. I was able to study some more living collections at the Leiden Hortus Botanicus,

competently cultivated by Art Vogel and Jacco Kruizinga. The Keeper and Curator/Herbarium

Manager at SINU (Prof. Dr. Hugh Tan Tiang Wah, Mr. Chua Keng Soon), KEP (Dr. Saw Leng

Guan, Dr. Richard Chung Cheng Kong) and KLU (Wong Khoon Meng, now Keeper of SING,

and Mr. Yong Kien Thai) are thanked for their hospitality and assistance. Fieldtrips organised

by KEP (Institut Penyelidikan Perhutanan Malaysia-FRIM) for me in Peninsular Malaysia

allowed collection of a number of species not seen in Singapore. Additional thanks are due to

Jana Leong-Skorniékova (SING) for digital images of Amischotolype collections in BO and

E, which made possible several taxonomic decisions. Mr. D. R6pert (B) kindly sent me digital

images of their type material. Dr. Jan-Frits Veldkamp (L) assisted me with nomenclature and

with the Latin diagnoses for the new species. I am grateful to the photographers who allowed

me to use their marvelous pictures: Serena Lee and Paul K.F. Leong (SING), Chew Ming Yee

(FRIM), André Schuiteman (K) and Jaap J. Vermeulen (L). Ben Kieft (L) gave the photos the

final touch for publication. Anita Walsmit Sachs (L) produced the excellent line drawings for

the new species, and flower and fruit details for the other species.

126 Gard. Bull. Singapore 64(1) 2012

References

Babu, C.R. & Dutta, R.M. (1968) Proposal to conserve the generic name 902 Forrestia A.Rich.

(1834) (Commelinaceae) against Forrestia Raf. (1806) (Rhamnaceae). Taxon 17: 597.

Backer, C.A. & Bakhuizen van den Brink, R.C. (1968) Forrestia. Flora of Java 3: 15.

Beaman, J.H. & Beaman, R.S. (1998) The Plants of Mount Kinabalu, 3: Gymnosperms and

Non-orchid Monocotyledons. Malaysia, Sabah: Natural History Publications (Borneo);

UK, Kew: Royal Botanic Gardens, Kew.

Blume, C.L. 1827. Campelia. Enumeratio Plantarum Javae \: 7.

Cherfils, H. (1937) Commeélinacées: Forrestia. In: Lecomte, H. (ed) Flore Générale de |’'Indo-

Chine 6: 852-862.

Clarke, C.B. (1881) Forrestia. In: de Candolle, A. & de Candolle, C. Monographiae

Phanerogamarum 3: 235-238.

Clarke, C.B. (1901) Commelinaceae. In: Thiselton-Dyer, W.T. (ed) Flora of Tropical Africa

Save

Coode M.J.E., Dransfield, J., Kirkup, D.W., Forman, L.L. & Said, I.M. (eds) (1996) A Checklist

of the Flowering Plants and Gymnosperms of Brunei Darussalam. Negara Brunei

Darussalam: Ministry of Industry and Primary Resources.

Cowley, J. (1996) In: Coode, M.J.E., Dransfield, J., Kirkup, D.W., Forman, L.L. & Said, I.M.

(eds) A Checklist of the Flowering Plants and Gymnosperms of Brunei Darussalam, p.

434. Negara Brunei Darussalam: Ministry of Industry and Primary Resources.

Cowley, J. & Furness, C.A. (1997) A new species of Amischotolype (Commelinaceae) from

Brunei: taxonomy and pollen morphology. Kew Bull. 52: 465469.

De Jussieu, A.L. (1820) Forrestia. In: Cuvier, G.-F. (ed) Dictionnaire des sciences naturelles

Me 250:

De Wildeman, E. (1909) Etudes de systematique et de géographie botaniques sur la Flore du

Bas- et du Moyen-Congo. Annales du Musée du Congo, Botanique - Sér. 5(3): 53.

Evans, T.M., Faden, R.B., Simpson, M.G. & Sytsma, K.J. (2000) Phylogenetic relationships in

the Commelinaceae: I. A cladistic analysis of morphological data. Syst. Bot. 25: 668-691.

Evans, T.M., Sytsma, K.J., Faden, R.B. & Givnish, T.J. (2003) Phylogenetic relationships in

the Commelinaceae: II. A cladistic analysis of rhcL sequences and morphology. Syst. Bot.

28: 270-292.

Faden, R.B. (1998) Commelinaceae. In: Kubitzki, K. (ed) The Families and Genera of Vascular

Plants 4:109-128.

Faden, R.B. & Hunt, D.R. (1991) The classification of the Commelinaceae. Jaxon 40: 19-31.

Forbes, F.B. & Hemsley, W.B. (1903) Enumeration of all the plants known from China proper,

Formosa, Hainan, The Corea, The Luchu Archipelago and the Island of Hong Kong;

together with their distribution and synonymy. J. Linn. Soc., Bot. 36: 1-686.

Griffith, W. (1851a) Notulae ad plantas asiaticas. India, Calcutta: Charles A. Serrao.

Griffith, W. (1851b) Icones plantarum asiaticarum III; Monocotyledonous plants. India,

Calcutta: Charles A. Serrao.

Hallier, H. (1898) Neue und bemerkenswerte Pflanzen. Bull. Herb. Boissier 6: 359.

Hallier, H. (1913) Commelinaceae: Forrestia. Nova Guinea 8: 906.

Hallier, H. (1916) Beitraége zur Flora von Borneo. Beihefte zum Botanischen Centralblatt 34:

46-49.

Hara, H. (1966) Commelinaceae: Amischotolype hookeri. The Flora of the Eastern Himalaya

le SY).

Hasskarl, J.K. (1852) Campelia. In: Miquel, F.A.W. (ed) Plantae Junghuhnianae 2: 154-155.

Hasskarl, J.K. (1863) Amischotolype Hassk., eine neue Gattung der Commelinaceae. Flora 46:

391-393.

Amischotolype in Asia 127

Hasskarl, J.K. (1864) Ueber Forrestia A.Rich. Flora 47: 625-631.

Hasskarl, J.K. (1870) Commelinaceae Indicae. Caroli Ueberreuter (M. Salzer), Vindobonae

(Vienna, Austria).

Ho, P. (1993) Amischotolype. An Illustrated Flora of Vietnam 3: 477-479. (In Vietnamese).

Hong, D.Y. (1974) Revisio Commelinacearum Sinicarum. Acta Phytotax. Sin. 12(4): 462-463.

Hong, D.Y. (1997) Amischotolype. In: Wu, Z.Y. (ed), Flora Reipublicae Popularis Sinicae

156); 71:

Hong, D.Y. & DeFilipps, R.A. (2000) Amischotolype, Porandra. In: Wu, Z.Y. & Raven, P.H.

(eds) Flora of China 24: 22-24.

Hooker, J.D. (1864) Forrestia hispida. Bot. Mag. 90: t. 5425.

Hooker, J.D. (1894) The Flora of British India 6: 383.

Hsu, K.-S. (1978) Amischotolype. In: Li, L.H., Liu, T.S., Huang, T.C., Koyama, T. & DeVol,

C.E. (eds) Flora of Taiwan 5: 158, t. 1305.

Hunt, D.R. (1986) Campelia, Rhoeo and Zebrina united with Tradescantia. Kew Bull. 41:

401-405.

Keng, H. (1987) Annotated list of seed plants in Singapore (XI). Gard. Bull. Singapore 40: 124.

Keng, H., Chin, S.C. & Tan, H.T.W. (1998) Amischotolype. The Concise Flora of Singapore

B23:

Kunth, C.S. (1843) Campelia. Enumeratio plantarum 4: 107-109, 116.

Lauterbach, C. (1913) Beitr. Fl. Papuas. 2. Die Commelinaceae Papuasiens. Bot. Jahrb. 50:

54-65.

McKean, D.R. (1988) Catalogue of the names published by Hector Léveillé: XX. Notes Roy.

Bot. Gard. Edinburgh 45: 153-159.

MeNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K.,

Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J. (2006)

International Code of Botanical Nomenclature (Vienna Code) adopted by the Seventeenth

International Botanical Congress Vienna, Austria, July 2005. Regnum Vegetabile 146:

1-568.

Merrill, E.D. (1906) I. New or noteworthy Philippine plants, 1V. Publ. Bur. Sci. Gov. Lab. 35:

5-6.

Merrill, E.D. (1921) A bibliographic enumeration of Bornean plants. J. Straits Branch Roy.

Asiat. Soc. [special number]: 113.

Merrill, E.D. (1925) Forrestia. Enum. Philipp. Fl. Pl. 1: 197-198.

Morton, J.K. (1967) The Commelinaceae of West Africa: A biosystematic survey. J. Linn. Soc.,

Bot. 60: 167-221.

Pichon, M. (1946) Sur les Commeélinacées. Notulae Systematicae. Herb. Mus. Paris 12: 232-

235,

Rafinesque, C.S. (1806) Additions to Michaux’s Flora of North-America. In a Letter from

Mr. Rafinesque, to Dr. Mitchill, dated Palermo, in Sicily, 8° August 1805. The Medical

Repository 2(3): 422.

Rao, R.S., Panigrahgi, G. & Kammathry, R.V. (1960) Cytotaxonomic studies on Indian

Commelinaceae. Proc. Indian Sci. Congr. Assoc., p. 366 (Abstract).

Rao, R.S. (1971) Notes on Indian Commelinaceae nomenclature and distribution. Maharashtra

Vidnyan Mandir, Patrika 6, 2: 52-55.

Richard, A. (1834) Forrestia. In: Dumont d’Urville (ed) Voyage de Découverts de |’Astrolabe

2: 1-5.

Richard, L.C.M. (1808) Démonstrations botaniques, ou analyse du fruit considéré en général,

46. Paris: H.A. Duval.

Ridley, H.N. (1904) New Malayan plants. J. Straits Branch Roy. Asiat. Soc. 41: 37-38.

128 Gard. Bull. Singapore 64(1) 2012

Ridley, H.N. (1907) Materials for a Flora of the Malayan Peninsula, part II. Singapore:

Methodist Publishing House.

Ridley, H.N. (1923) A botanical excursion to Northern Sumatra. J. Malayan Branch Roy. Asiat.

Soc. 1: 102-103.

Ridley, H.N. (1924) Forrestia. Flora of the Malay Peninsula 4: 359-361.

Ridley, H.N. (1925) Dr. H.O.Forbes’s Malayan plants. J. Bot. 63: 124.

RGpert, D. (ed) (2000) Digital specimen images at the Herbarium Berolinense. http://ww2.

bgbm.org/herbarium/ (accessed on 14 Jan. 2011).

Schumann, K. (1897) Commelinaceae africanae. Bot. Jahrb. 24: 344.

Steudel, E.G. von (1840) Nomenclator Botanicus, ed. 2(1). Germany, Stuttgart: J.G. Cotta.

Thitimetharoch, T., Chantaranothai, P. & Faden, R.B. (2003) The genus Porandra

(Commelinaceae) in Thailand. Thai Forest Bull., Bot. 31: 141-148.

Turner, I.M. (1996) Some nomenclatural adjustments in the Commelinaceae, Hypoxidaceae,

Lauraceae, Rubiaceae, Sapotaceae, and Zingiberaceae of the Malay Peninsula. Novon 6:

221.

Turner, I.M. (1997) A catalogue of the vascular plants of Malaya. Gard. Bull. Singapore 47:

515-516.

Walker, E.H. (1976) Flora of Okinawa and the Southern Ryukyu Islands, pp. 298-299. U.S.A.,

Washington: Smithsonian Institute Press.

Wang, J.C., Chen, Y.J. & Peng, C.I. (2000) Commelinaceae. Flora of Taiwan, ed. 2(5): 153—

154.

Yang, Y.-P., Liu, H.-Y. & Lin, T.-P. (2001) Amischotolype. Concise Vascular Flora of Taiwan

5: 457:

Yuan, Q. & Yang, Q.-E. (2006) Chromosomes of four species in three genera of Commelinaceae

from China and their systematic implications. Bot. J. Linn. Soc. 152: 399-403.

129

Amischotolype in Asia

Appendix A. Character synopsis for the species of Amischotolype and Porandra: vegetative characters.

Ry 8 |2

< sy 6 |z3

& ae eee Wes So es = & lie |e

3 = |S s s i=} 3) 3 a 5 a a 3)

2 le le = 4 s | a Sle ia fs |S Jeg |S |o f-s | ]a

s = s c = = iz} 5 =I < A D & = i= aD S

S 3 s = 2 = 3 oS =I Ss S 3 s = BZ Ss ct o rf 5b 3S oo s Ss

el RSs licsy EO SE eS SSS sR | eS er | ice SI (Se cs

FO Soe ER ice ies irs ee ie [ese WES ie! oS ea ee ro ie: lis

a |.2 ey |e || et las = |\-2 me [Les fe ee Te tE ASYM EY Upvete |hree, [10S a ila |e |p

= |e |e || |e a I alice < |< |< |< le le le |< |< |< |< [oe Jo

<i \le Nie [le | |e Nee Neel eS ISS [ISS IS Ns ISS (se |S ISS ES

bg 5 . 5 ° = « o = (=) _ nN foe) Tt al oS > co aD =) = an vt NQ

= a an t+ Vay So > oo D> = = = _ _ ol = = — = nN nN iol aq a

RHIZOME

absent/present

STEM

simple

branched

scrambling

INTERNODE HAIRS

absent

0.1—0.5 mm long

0.6—0.9 mm long

> 1mm long

SHEATH HAIRINESS

glabrous

scattered and < 1 mm long

in lines and 0.5—1 mm long

in lines and > 1 mm long

in ring at node, 0.5—1 mm long

LEAF BLADE SURFACE

smooth

bullate

LOWER LEAF SURFACE

HAIRINESS*

glabrous

hairs < 0.5 mm long

hairs 0.5—1 mm long

hairs > 1 mm long

UPPER LEAF SURFACE

HAIRINESS*

glabrous

hairs < 0.5 mm long

hairs 0.5—1 mm long

hairs > 1 mm long

LEAF SUBMARGINAL

HAIRS - PRESENCE

absent

on upper surface

along margin

on lower surface

LEAF SUBMARGINAL

HAIRS - LENGTH

< 0.5 mm long

>0.5 <1 mm long

1-2 mm long

> 2mm long

LEAF SUBMARGINAL

HAIRS - COLOUR

colourless

white

yellow to yellow-brown

orange to red

*except near and on midvein.

130

Gard. Bull. Singapore 64(1) 2012

Appendix B. Character synopsis for the species of Amischotolype and Porandra: inflorescence,

flower and fruit characters.

3 Ss z 5

a Ss e io 3 <= 3 = =I

3 = Se Sia lisa ome les E s|/§ 1s

Be Ne eal Nes = ta 2 hells fate eels Wale Pe eels: lea)

£/2 |2£ |s |2 |S [8 |S [5S |S le 18 1S [BIZ JS ls ISP IEP IS I8 1S IS

EE POEs eels ie a ies Wee SS Ws Wes WSS We ee ES ee he Ie

ele lo 18 | fe |8 |Z is |2 18 |S |S |e |e le /2 12/2 1812 IE le le

FO ae Ne es En ess lise Ise se Se Se Se [So Ss iso Se se: Se Ss HE! |e

S 5 S 5 : 3 : =) = a on + va) No} ~ oe) a So = a - a

_ N (oa) wr 0 ao] ~ oo a> = = = = = = = = = = Nn a fol a a

INFLORESCENCE POSITION

rhizome + + +

knee 2s (+) Je + + +

erect stem - - _ - - + + a + + + “fe + + 2s + + oe

PEDUNCLE (-), (-)

absent/present = - = 3 = = = = Sr lfesee et Ss Wes |! oi} as] s = = [le i= S|) x |]

PEDICEL

absent - + + + + + + + + +

< 1.5 mm long a: an] se | eee + Han ere |e a

> 1.5 mm long 4: ae de + + a

SEPAL HAIRS -

PRESENCE, COLOUR

absent + + 14/4 + - + . - + | (+) + + + + +

colourless-white-yellow 4 + pi | exe rom (ere ae ae + y Se ae 4. a ti at) | [letu

orange-red-brown a + + + se] |e

SEPAL HAIRS -

HAIR TYPE

soft + + a +. - + + = + + + + + = + - +

bristly/spiny au | eer

PETAL HAIRINESS =

glabrous (-) / hairy (+) = = 5 - |@] - = = + || = i») x 2 = = ? = = me eee bd |)

STAMEN FILAMENT

HAIRINESS

glabrous + + ? ? +

0.5—1 mm long (+)] + ? ay 9 + eaters

| > 1 mm long ee va dk re iy + ae ae 9) 4 ue ay 7) + i.

THECAE OPENING

apical pore + | + ? + + | +

half slit + 9

longitudinal slit ae |) ae |) ge |] au |) 4: ve |} az |f ae fe Wee |] ga |p gh |) py || se se | | ee |! ce

ANTHER LENGTH

<1 mm long se > ) ve 4

I-1.9 mm long + (+)] + | + | + fe |) + | eo el eee |e ? + |e ey +] +

2-3 mm long + re 3. + D > a 4

CAPSULE : SEPAL LENGTH

capsule longer + + + | +] + + + +/+ ]+]+ +

equal re (+) (+) a

capsule shorter + + + + + + (+4)] + | + | + +] +

CAPSULE HAIRINESS

absent (+) Be f + +

soft + + + + = + + + + + + . + +

bristly/spiny rl +4 + + fe a

CAPSULE HAIR LENGTH

<1 mm long + | (4) + + +] + + + | + +] 4+

> 1 mm long + 4 + + + + + | (+) + +74] 4+ ]+ + + = (+)

| CAPSULE HAIR COLOUR

colourless-white-yellow + | + + | + 7 4 + +] +] + + + + orell [ae

orange-red-brown + 4 + (+) ‘ a

| CAPSULE SHAPE

Ovol + + + + + + + + + +

obovoid + fi 4 sts + + + ae i +

| pyriform + + +

CAPSULE APEX

| depresse' + | (+) + + + + + - + > + + + + 7 + +

| obtuse } te i‘ ab + +

acute +

LOBES AT CAPSULE APEX

| absent + + + + + - + + + + + + + + + + + . .

0.30.5 mm long +4 + <i

> 0.5 mm long + +

| CAPSULE VALVES

| free (fused < 1/4 length) + + + + + + + + + + + ? 2 + + =f

| fused > 1/4-half length } + ir 4 9|9 + ot {fie

| fused 4/5 length 9 ; cs

STYLE REMNANT :

persistent + - + (+)] + + + + a ve (?) + +

| (finally) deciduous or absent fi - tu z re fe a 4 ds + + a A

131

Amischotolype in Asia

Appendix C. Character synopsis for the species of Amischotolype and Porandra: distribution.

wynaysou *

wyuypootpad °

tagnd ©

nunedsouow *

VUUISST FOUL”

wyourdueu *

ndanoory ©

WOUND]

Hoyooy *

epidsiy *

wynsaty

mngnyd

siplona

wupunyorfop *

VIVOLIBAIP ©

ussoruqung *

Gardens’ Bulletin Singapore 64(1): 133-137. 2012 133

First record of Taeniophyllum (Orchidaceae) in Myanmar

H. Kurzweil! and S. Lwin?

'Herbarium, Singapore Botanic Gardens, | Cluny Road, Singapore 259569

hubert_kurzweil@nparks.gov.sg

* Myanmar Floriculturist Association, Ahlone Road, Yangon, Myanmar

ABSTRACT. Taeniophyllum Blume was recently discovered in northern Myanmar, a new

generic record for the country. The Myanmar specimens are referred to the widespread species

T. glandulosum Blume, characterised by terete roots, warty inflorescence axes, distichous

bracts, sepals and petals basally fused into a tube about as long as their ovate-lanceolate free

parts, and an ovate-lanceolate lip with a globose spur.

Keywords. Myanmar, Taeniophyllum

Introduction

While undertaking fieldwork near Lake Inndawgyi in Kachin State, Myanmar, the

authors of this article came across a leafless epiphytic orchid and recognised it as

member of the genus 7aeniophyllum Blume. Subsequent studies revealed that this

genus has not yet been recorded in Myanmar (Govaerts et al. 2011, Kress et al. 2003, P.

Ormerod pers. comm.). The genus Zaeniophyllum comprises about 120 species, which

are distributed from Sri Lanka and India throughout tropical and subtropical Asia

eastwards as far as Japan, Australia and several Pacific islands, with a single species

found in tropical Africa. Given the fact that Zaeniophyllum occurs in all surrounding

countries, its newly discovered occurrence in Myanmar is not surprising.

Using recent orchid flora treatments (particularly Jayaweera 1981; Seidenfaden

1988, 1992: Seidenfaden & Wood 1992; Pearce & Cribb 2002; Chen & Wood 2009),

the plants could be positively identified as 7. glandulosum Blume. This species belongs

to a taxonomically difficult complex of several closely related species, and several

botanists have recently pointed out the need for further studies. The complex is most

diverse in the Malay Islands.

Taeniophyllum glandulosum Blume

Bijdr. (1825) 356; Seidenf., Opera Bot. 95 (1988) 23; Comber, Orch. Java (1990) 363;

Seidenf. & Wood, Orch. Pen. Malays. Singap. (1992) 575; Chen & Wood in FI. China

25 (2009) 444. SYNTYPES: Indonesia, Java, Mount Gede and Pangurangu, B/ume

s.n. (repositories not known). Fig. 1 & 2.

Epiphytic herbs, entirely glabrous, leafless in the flowering stage. Roots creeping

directly on the tree bark or in moss growing on it, whitish-green in the dry state,

smooth, terete or semi-terete and hardly flattened, about 2—5(—10) cm long, 0.7—1

mm in diameter. Inflorescences solitary, erect or spreading at the base and arching

134 Gard. Bull. Singapore 64(1) 2012

Fig. 1. Zaeniophyllum glandulosum Blume. A. Whole plant. B. Flower. Kurzweil & Lwin 2790.

Photo credits: Dr. E. Frei.

above, 7-10 mm long; peduncle and rachis brownish green, 0.2—0.4 mm in diameter,

warty; rachis 3—7-flowered with one flower open at a time; floral bracts distichous,

ovate-lanceolate, thickly textured, 0.9-1.2 « 0.7—-0.9 mm when spread out, warts as

on the rachis though somewhat denser. Flowers yellowish green. Pedicel and ovary

indistinguishable, 1.5—2 mm long. Sepals and petals fused in their lower half forming

a tube c. 1.5 mm long, free portions somewhat recurved, dorsally slightly carinate,

apices subacute; free parts of median sepal lorate, 1.25 < 0.4 mm, free parts of lateral

sepals ovate-lanceolate, c. 1.5 x 0.7 mm; free parts of petals broadly ovate-lanceolate,

1.3 x 0.5 mm; lip ovate-lanceolate, navicular with incurved margin, spurred, base

with a septum at the spur entrance, c. 2.2 x 0.75 mm, apex acuminate, with a reflexed

elongate appendage c. 0.5 mm long; spur a globose pouch, c. | x 0.95 mm, glabrous

on the outside, distal part with a minute thickened gland on the inside. Gynostemium

c. 0.5 long, with prominent stelidia. Capsule not seen.

Illustrations. Seidenf., Opera Bot. 95 (1988) 23, Fig. 9a-c; Comber, Orch. Java (1990)

363, photo; Seidenf. & Wood, Orch. Pen. Malays. Singap. (1992) 575, Fig. 259 I-n;

Chen et al. in Fl. China 25, Illustrations (2010) 589, Fig. 589.1-14 & 591, Fig. 591.2—4.

Specimens examined: MYANMAR. Kachin State: Inndawgy1 Wildlife Sanctuary about 115

km SW of Myitkyina, western shore of Lake Inndawgyi, N 25° 10.329’, E 96° 16.815’, 5 May

2010, Kurzweil & Lwin 2790 (SING spirit).

Habitat, abundance and flowering time. The Myanmar specimens were growing as

epiphytes in degraded forest and were locally common on tall trees of Terminalia

bellirica (Gaertn.) Roxb. (Combretaceae). Plants were growing on twigs about | cm

thick. Flowering specimens were found in the beginning of May and the presence of

several plants in the bud stage suggests that flowering would still continue for many

Taeniophyllum glandulosum (Orchidaceae) 135

Fig. 2. Taeniophyllum glandulosum Blume. A. Roots and base of inflorescence. B, C. Cross-

sections of roots. D. Inflorescence apex. E. Sepals and petals flattened out, showing the length

of the floral tube. F. Lip. Scale bars: | mm. Kurzweil & Lwin 2790. Drawn from spirit material

by Evonne Tay.

weeks. This corresponds well with a previous report from China where flowering

was reported as occurring between April and August (Chen & Wood 2009). In other

parts of the distribution area, Taeniophyllum glandulosum is generally reported as

occurring in forest, and in Vietnam the species is common in primary mountain forest

(Averyanov et al. 2003). Elevations differ significantly in this and previous reports.

While the Myanmar plants were collected in a lowland forest about 185 m above

sea level, altitudes were given as 400-1100 m in China (Chen & Wood 2009) and

1000-2500 m in Vietnam (Averyanovy et al. 2003). In Java, this species is found in

high-altitude forest at c. 2280-2700 m, and is sometimes also found growing on rocks

(Comber 1990).

Distribution. India (Assam), Myanmar, Thailand, Vietnam, central and SW China (N

Fujian, Guangdong, Hainan, Hunan, NE Sichuan, S Yunnan), Taiwan, Japan, Korea,

Peninsular Malaysia, Java, Sulawesi, New Guinea.

Notes. Our specimens clearly match the descriptions and illustrations of Taeniophyllum

136 Gard. Bull. Singapore 64(1) 2012

glandulosum Blume that we have consulted (Seidenfaden 1988, Seidenfaden & Wood

1992, Chen & Wood 2009, Chen et al. 2010). Distinctive characters of this species

are the smooth, slender, terete roots; the short and few-flowered inflorescences with

warty peduncles and rachis; the distichously arranged thick warty bracts; the minute

yellowish green flowers; the basal fusion of the sepals and petals which is about as

long as their ovate-lanceolate or lorate free parts; and the ovate-lanceolate lip with

a reflexed elongate appendage near the tip and the basal globose spur. The eastern

Himalayan 7. retrospiculatum (King & Pantl.) King & Pantl. and 7) arunachalense

A.N.Rao & J.Lal share the general habit and the shape of the sepals and petals but

differ in a relatively shorter sepal/petal tube with the free portions 3- or 4-times as long

as the fused portion.

We suspect that Zaeniophyllum glandulosum may be rather widespread

in Myanmar. The orchid flora of this country is generally not well explored and a

comprehensive inventory is not yet available. While searching for orchids in the wild,

the few collectors usually focus on the large-flowered orchids which are horticulturally

valuable and consequently 7. g/andulosum may have been overlooked because of its

small size.

ACKNOWLEDGEMENTS. Weacknowledge the Myanmar Forestry Department for permission

to undertake floristic survey work and to collect material of Orchidaceae. We express sincere

thanks to Mr. Stephen Lasi Bawk Naw and Mr. Brangaung for help in many ways and to U Sein

Tun (Warden of Inndawgy1 Wildlife Sanctuary) for assistance in the field. Furthermore, we are

indebted to Mr. P. Ormerod for useful comments. We also thank Mrs. Evonne Tay (Singapore)

for making the line drawings and Dr. E. Frei (Switzerland) for permission to use his colour

photos here. The second author would also like to acknowledge financial assistance provided

by a Singapore Botanic Gardens Research Fellowship.

References

Averyanov, L.V., Phan Ke Loc, Nguyen Tien Hiep & Harder, D.K. (2003) Phytogeographic

review of Vietnam and adjacent areas of Eastern Indochina. Komarovia 3: 1—83.

Chen, S.-C. & Wood, J.J. (2009) Taeniophyllum Blume. In: Wu, Z.Y., Raven, P.H. & Hong,

D.Y. (eds) Flora of China, vol. 25, pp. 444. Beijing: Science Press & St. Louis: Missouri

Botanical Garden Press.

Chen, S.-C., Liu, Z.J., Zhu, GiEL, Lang Ko-Y¥5 Ts) Zn.) buoy CBs Jin; 2c Cabos

Wood, J.J., Gale, S.W., Ormerod, P., Vermeulen, J.J., Wood, H.P., Clayton, D. & Bell,

A. (2010) Orchidaceae. In: Wu Z.Y., Raven, P.H. & Hong D.Y. (eds) Flora of China,

vol. 25, illustrations, pp. 1-666. Beijing: Science Press & St. Louis: Missouri Botanical

Garden Press.

Comber, J.B. (1990) Orchids of Java. Kew: Royal Botanic Gardens, Kew.

Govaerts, R., Bernet, P., Kratochvil, K., Gerlach, G., Carr, G., Alrich, P., Pridgeon, A.M., Pfahl,

J., Campacci, M.A., Holland Baptista, D., Tigges, H., Shaw, J., Cribb, P.J., George, A.,

Kreuz, K. & Wood, J.J. (2011) World Checklist of Orchidaceae. Facilitated by the Royal

Botanic Gardens, Kew. http://apps.kew.org/wesp/ (accessed on 25 May 2011).

Jayaweera, D.M.A. (1981) Orchidaceae. In: Dassanayake, M.D. & Fosberg, F.R. (eds) A

Taeniophyllum glandulosum (Orchidaceae) iy

Revised Handbook to the Flora of Ceylon, vol. 2, pp. 4-386. New Delhi: Amerind

Publishing Co.

Kress, W.J., DeFilipps, R., Farr, E. & Yin-Yin-Kyi. (2003) A checklist of the trees, shrubs,

herbs and climbers of Myanmar. Contributions from the US National Herbarium 45:

i590:

Pearce, N.R. & Cribb, P.J. (2002) The Orchids of Bhutan. In: Flora of Bhutan 3(3). Edinburgh:

Royal Botanic Garden Edinburgh & Bhutan: Royal Government of Bhutan.

Seidenfaden, G. (1988) Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Bot.

95: 15398.

Seidenfaden, G. (1992) The orchids of Indochina. Opera Bot. 114: 1-502.

Seidenfaden, G. & Wood, J.J. (1992) The Orchids of Peninsular Malaysia and Singapore.

Fredensborg: Olsen & Olsen.

Gardens’ Bulletin Singapore 64(1): 139-169. 2012 139

Diversity of the tree flora in Semenggoh Arboretum,

Sarawak, Borneo

C.Y. Ling’ and S. Julia?

Botanical Research Centre, Sarawak Forestry Corporation,

Km 20, Jalan Puncak Borneo, 93250 Kuching, Sarawak

‘cyling@sarawakforestry.com (corresponding author)

*juliasang@sarawakforestry.com

ABSTRACT. A 4-ha sample plot was established at the Arboretum, Semenggoh Forest Reserve

to document tree species in this lowland mixed dipterocarp forest. The area assessed contains

2837 trees with diameter at breast height > 10 cm belonging to 60 families, 160 genera and 541

species. Euphorbiaceae and Malvaceae (10 genera each) were the most diverse families at genus

level and Dipterocarpaceae (61 species) is most diverse at species level. More than 25% of

trees (720 individuals) were dipterocarps and contributed the highest basal area (cross-sectional

area over-bark at breast height measured in m*) of 16.7 m*/ha. The most abundant species are

Shorea multiflora (21 trees/ha) and Pouteria malaccensis (31 trees/ha) for dipterocarp and

non-dipterocarp species, respectively. Semenggoh Arboretum has a rich and diverse flora and,

being a natural primary forest in the middle of an increasingly developed area, Semenggoh is

important as a genetic reservoir for threatened species (particularly the dipterocarps) and as an

in-situ conservation site for Sarawak’s lowland mixed dipterocarp forest.

Keywords. Borneo, Dipterocarpaceae, Sarawak, Semenggoh Arboretum, tree diversity

Introduction

Floristic composition studies or floristic analyses are a useful tool to understand the

spatial pattern of plant composition and diversity. When combined with ecological,

environmental, geological and historical variables, important information on

mechanisms maintaining high levels of tree species diversity in tropical forest can

be obtained (Slik et al. 2003). For that reason, a floristic composition study is the

prerequisite study, in order to determine the species richness and diversity of a selected

area. Conducting long-term studies is the only way to understand the dynamics of

forest ecosystem due to slow-growing and long-lived trees and numerous long-term

study sites were set up in many forests, including in Asia (Condit et al. 2000, Lee et al.

2004, Primack & Hall 1992).

Borneo, of which Sarawak is a part, is renowned for its rich and diverse flora

comprising of about 3500-5000 tree species (Soepadmo 1995, Soepadmo & Chung

1997). Semenggoh Forest Reserve is Sarawak’s oldest forest reserve constituted in

1920, comprising an area of 653 ha (Forest Department Sarawak 2000). It includes

an arboretum, a wildlife rehabilitation centre and a botanical research centre. The

Semenggoh Arboretum was established in 1951 to preserve in perpetuity a small

140 Gard. Bull. Singapore 64(1) 2012

area of easily accessible natural primary lowland forest located at 1°23’50.0”N to

1°24’5.7”N and 110°18’53.1”E to 110°19°13.0”E, and less than 100 m elevation. It

covers an area of approximately 14 ha and is one of the few least disturbed lowland

mixed dipterocarp forests located about 20 km from Kuching City (Fig. 1A). In April

2000, the Reserve was gazetted as Semenggoh Nature Reserve, which is now legally

protected in Sarawak. The arboretum 1s floristically rich and has attracted a large

number of researchers and naturalists (Forest Department Sarawak 2003).

Generally, Semenggoh is more well-known for its orang-utan rehabilitation

centre rather than its flora as very litte information on the flora of Semenggoh has

been published. Based on herbarium records, many local and overeseas botanists have

collected specimens from the Semenggoh Arboretum from the 1950s to 1970s. To

date, about 700 species (from about 2000 herbarium specimens) from Semenggoh are

represented at the Sarawak herbarium. Among these collections, about 40 species were

described from this locality (e.g., Shorea pubistyla P.S.Ashton, Dipterocarpaceae;

Brownlowia ovalis Kosterm., Malvaceae / Tiliaceae; and Macaranga kingii Hook.f.,

Euphorbiacae) and nine species are endemic to Semenggoh (e.g., Chionanthus rugosus

Kiew, Oleaceae; Xanthophyllum ceraceifolium Meijden, Polygalaceae; and Areca

ahmadii J.Dransf., Arecaceae). A census of trees (girth 45 cm and above) carried out by

the Sarawak Forest Department in the 1980s recorded 12,778 trees from Semenggoh

(unpublished report).

Being one of the reserves located near Kuching City, Semenggoh continues

to face pressure from social and economic development; therefore, it 1s critical to

document the flora of the whole Nature Reserve in order to study its diverisity. This

paper is on the preliminary work carried out in Semenggoh Arboretum to investigate

the tree flora diversity, with the long-term aim of fully documenting the flora of

Semenggoh Nature Reserve. The objectives of this work are to study the tree flora

composition and species richness and to investigate the current conservation value of

Semenggoh Nature Reserve.

Fig. 1. Location of Kuching city (A) and Semenggoh Nature Reserve (B), showing the 4-ha

study area along the eastern side of Masing Trail up to the Arboretum boundary, near the

road leading to the Wildlife Rehabilitation Centre. (Courtesy of GIS Unit, Forest Department

Sarawak.)

Tree diversity at Semenggoh, Sarawak 141

Methodology

Sample plot establishment

Sample plots were established following the method outlined in the “Manual for

Establishment and Enumeration of Permanent Sample Plots in Peat Swamp Forest in

Sarawak” (Tan 2002) with some modification to meet the objectives of the project at

Semenggoh Arboretum. A total of 484 quadrats (each 10 m x 10 m) were established in

an area of approximately 4.0 ha (Fig. 1B). All quadrats were numbered consecutively,

line by line.

Field assessment

A 100% enumeration was carried out for trees with diameter > 10.0 cm. The diameter

of each tree was measured at 1.3 m above the highest ground level for non-buttressed

trees, and 30 cm above the highest buttress for buttressed trees. Leaf samples from

all trees were collected for botanical identification to species level. The collected leaf

samples were identified using the Tree Flora of Sabah and Sarawak volumes I-VI

(Soepadmo & Wong 1995; Soepadmo & Saw 2000; Soepadmo et al. 1996, 2002, 2004,

2006, 2007) and other monographs and publications (e.g., Airy Shaw 1975, Chung

2005, Julia 2005, Sugau 2005, Ashton 2006), as well as by comparison to specimens

kept in the Sarawak Herbarium.

Data analysis

For data analysis, the number of trees, families, genera and species were transformed

into their per ha equivalent. All data analysed were used to determine the floristic

composition, species abundance, and diameter and basal area contribution by all tree

species. Basal area for each tree was calculated by using the formula: Basal area =

(DBH/200)° x zm’, where DBH is the diameter at breast height in cm. To determine the

stand density and basal area contribution, each tree was then grouped into six different

diameter classes (10.0—19.9; 20.0-29.9; 30.0-39.9: 40.0-49.9; 50.0—-59.9; > 60.0 cm).

Species richness was determined by a relative comparison method, comparing the

trees/ha and species/ha of each plot between Semenggoh and four other plots from

different sites established in Sarawak. No statistical method was used to calculate

species richness, as no duplicates of different plot sites or forest types were taken into

consideration in the initial purpose of this study.

Results and discussion

Tree diversity and density

A total of 2837 individuals from 60 families, 160 genera and 541 species were recorded

from the sample plot (Appendix A). Euphorbiaceae and Malvaceae were the most

diverse families at genus level, with 10 genera each. Relative comparison with four

other sites showed that Semenggoh ranked fourth in terms of species richness after

Lambir Hills National Park (NP), Batang Ai NP and Bako NP (Table 1). Batang Ai NP

142 Gard. Bull. Singapore 64(1) 2012

has the highest number of trees recorded, with 813 trees/ha compared with 710 trees/

ha in Semenggoh. Twenty eight families (46.7% of total number of families) were

represented by a single genus and 13 families (21.7% of the total number of families)

were represented by a single species. About 37% of the total species are represented

by a single individual. Within the study area, two families (1.e., Icacinaceae and

Salicaceae), each represented by a single species, had the lowest number of individuals

and also basal area. Both trees are very small in size and may face local extinction from

the plot in the future, due to natural disturbances or anthropogenic disturbances, as the

mortality of only a few individuals will lead to total loss of representation in the plots.

This may result in species turnover in the future. Furthermore, rare species demonstrate

higher mortality rates than common species but, at the same time, contribute to high

diversity in forests in Borneo (Primack & Hall 1992).

Of the 60 families recorded from the plot, Dipterocarpaceae is the main

contributor in terms of total stems (720 individuals), total number of species (61

species), and total stand basal area (16.71 m’/ha) within the study area (Table 2).

Myristicaceae is the most diverse family among non-dipterocarps, with 42 species

recorded. Sapotaceae is the most dominant family with 66 trees/ha, followed by

Myristicaceae (54 trees/ha) and Moraceae (40 trees/ha). In terms of basal area

contribution, Sapotaceae, Moraceae, and Malvaceae contributed a total basal area of

3.32 m’/ha, 1.97 m’/ha, and 1.86 m’/ha respectively, while Fabaceae contributed the

smallest total number of species, trees and basal area among the ten most abundant

families. Dipterocarpaceae is the most dominant family in Semenggoh and other sites

in Sarawak (Table 3). The occurrence of Dipterocarpaceae in Semenggoh is relatively

high when compared with other similar sites in Sarawak (except the 52-ha plot in

Lambir Hills NP) (Table 4).

At the generic level, Shorea (Dipterocarpaceae) contributed the highest

number of trees with 129 trees/ha, followed by Artocarpus (Moraceae) with 38 trees/

ha, Pouteria (Sapotaceae) with 31 trees/ha, and Syzygium (Myrtaceae) with 29 trees/

ha (Table 5). Shorea also contributed the highest basal area with 44.73 m?, followed

by Pouteria (8.75 m?), Hopea (7.51 m*) and Artocarpus (7.29 m*). Tree composition

in Semenggoh Arboretum is dominated by dipterocarp species with more than 25%

of trees belonging to the Dipterocarpaceae. From the total of 2837 trees recorded,

dipterocarp species contributed 180 trees/ha, compared with 529 trees/ha by non-

dipterocarp species. This is a typical pattern for most evergreen rain forests on the

Sunda Shelf region which encompasses the mainland of Peninsular Malaysia, Sumatra,

Java and Borneo (Slik et al. 2003).

Among the dipterocarps, Shorea multiflora (Burck) Symington

(Dipterocarpaceae) is the most abundant species represented by the highest

number of individuals with 84 individuals, followed by Shorea macroptera Dyer

(Dipterocarpaceae) with 74 individuals and Shorea brunnescens P.S.Ashton

(Dipterocarpaceae) with 62 individuals, while Pouteria malaccensis (C.B.Clarke)

Baehni (Sapotaceae) is the most abundant species for non-dipteocarps, followed by

other species as shown in Table 6. Pouteria malaccensis also contributed the highest

basal area (2.19 m’/ha) for the non-dipterocarps. Interestingly, although a low number

Tree diversity at Semenggoh, Sarawak 143

Table 1. Species diversity of the 4-ha plot in Semenggoh compared with other sites.

FDS: Forest Department Sarawak.N: Number of trees. S: Number of species.

Site N S Area (ha) N/ha S/ha

Semenggoh 2837 541 4.0 710 135

Lambir Hills NP (FDS 1986) 1672 428 2.4 697 178

Bako NP (FDS 1996) 1901 390 2.4 792 162

Bukit Mersing (FDS 1996) 1584 266 2.4 660 11]

Batang Ai NP (Diway et al., 2009) 1831 385 225 813 171

Table 2. Ten most abundant families recorded from the 4-ha study area in Semenggoh Arboretum.

S: Number of species. N: Number of trees. BA: Basal Area.

% Trees: N/Total number of trees x 100%.

Family S N Trees/ha % Trees BA (m?/ha)

Dipterocarpaceae 61 (11%) 720 (25%) 180 25.38 16.71 (39%)

Sapotaceae 27 (5%) 262 (9%) 66 9.24 3.32 (8%)

Myristicaceae 42 (8%) 216 (8%) 54 7.61 1.65 (4%)

Moraceae 11 (2%) 160 (6%) 40 5.64 1.97 (5%)

Malvaceae 29 (5%) 125 (4%) 31 4.4] 1.86 (4.32)

Myrtaceae 29 (5%) 114 (4%) 29 4.02 1.43 (3%)

Burseraceae 28 (5%) 106 (4%) LY 3.74 1.28 (3%)

Anacardiaceae 18 (3%) 94 (3%) 24 3.3) 1.69 (4%)

Clusiaceae 38 (7%) 84 (3%) 21 2.96 0.93 (2%)

Fabaceae 15 (3%) 73 (3%) 18 2 2.55 (6%)

Table 3. Five most dominant families in Semenggoh and other sites of Sarawak.

FDS: Forest Department Sarawak.

Bemiedpeoh Lambir Hills NP Bako NP Bukit Mersing Batang Ai NP

(FDS 1986) (FDS 1996) (FDS 1996) (Diway et al. 2009)

Dipterocarpaceae Dipterocarpaceae : Dipterocarpaceae Dipterocarpaceae Dipterocarpaceae

Sapotaceae Euphorbiaceae Myristicaceae Euphorbiaceae Euphorbiaceae

Myristicaceae Burseraceae Anacardiaceae Verbenaceae Myrtaceae

Moraceae Anacardiaceae Sapotaceae Annonaceae Clusiaceae

Malvaceae Myristicaceae Clusiaceae Myristicaceae Burseraceae

144 Gard. Bull. Singapore 64(1) 2012

Table 4. The diversity of Dipterocarpaceae in Semenggoh (4-ha study area) and other sites of

Sarawak. FDS: Forest Department Sarawak.

Site pas netaee Trees/ha

“Semenggoh 7 61 180

Lambir Hills NP (FDS 1986) 7 5D) 180

Lambir Hills NP -52 ha plot (Lee et al. 2004) 8 78 119

Bako NP (FDS 1996) ¢ 49 215

Bukit Mersing (FDS 1986) 5 22 140

Batang Ai NP (Diway et al. 2009) 6 45 267

of trees was recorded for Koompassia malaccensis Maingay ex Benth. (Fabaceae)

and Dyera costulata (Miq.)Hook.f. (Apocynaceae), both species contributed relatively

high basal areas of 1.56 m’/ha and 1.05 m/*/ha, respectively. As for dipterocarps,

although Shorea multiflora contributed the highest number of individuals, Shorea

beccariana Burck (Dipterocarpaceae) with only 6 trees/ha, contributed 2.07 m?’/ha

basal area and was 53.62% higher than Shorea multiflora, followed by Hopea dyeri

F.Heim (Dipterocarpaceae) (1.46 m’/ha), Shorea elliptica Burck (Dipterocarpaceae)

(1.41 m?/ha) and Dipterocarpus genticulatus Vesque (Dipterocarpaceae) (1.07 m?/ha).

Even though Semenggoh Arboretum is dominated by dipterocarp species,

non-dipterocarp species (1.e., Pouteria malaccensis) contributed the highest number

of trees and basal area. This circumstance may relate to the seed dispersal limitations

of Dipterocarpaceae and Sapotaceae (Whitmore 1984). Dispersal limitation plays an

important role in structuring floristic composition over distance of less than five km

(Pyke et al. 2001). Dipterocarp seeds are dispersed by wind, but their heavy weight

limits the seed dispersal range, resulting in their germinating closer to the mother

tree. Sapotaceae seeds are dispersed by animals, and seedlings of Sapotaceae will

eventually establish further from the mother trees, thus reducing competition between

mother trees and seedlings for resources such as light, nutrients and space (Tan et al.

1998). Another possible reason is probably due to the mast-fruiting of Bornean species,

which occurs at five to ten yearly intervals. In dipterocarp mast-fruiting events, a delay

in timing of seedfall after the community fall peak is negatively correlated with seed

survival and seedling establishment (Curran & Webb 2000).

Stand density and basal area by diameter classes

Six diameter classes were considered, ranging from 10.0 cm to = 60.0 cm. The

distribution of trees in the study area followed a normal reverse-J-shaped curve where

the lower diameter classes were dominated by small trees and only few bigger trees

dominated the higher diameter classes. Out of 2837 individuals recorded, 57.41% of

trees were within the 10.0-19.9 cm diameter range (Fig. 2). Non-dipterocarp species

Tree diversity at Semenggoh, Sarawak 145

Table 5. Ten most abundant genera recorded from the 4 ha study area. N: Number of trees. BA:

Basal area. % Trees: N/Total number of trees = 100%.

% BA BA

Genera Family N Trees/ha ee (m2) Gu2fhiad

Shorea Dipterocarpaceae 515 129 18.13 44.73 11.18

Artocarpus Moraceae 150 38 5.28 T29 1.82

Pouteria Sapotaceae 122 ail 4.30 8.75 29

Syzygium Myrtaceae 114 29 4.01 DoS 1.43

Gymnacranthera Myristicaceae 9] 23 3.20 2.94 0.74

Hopea Dipterocarpaceae 86 22 3.03 feat 1.88

Palaquium Sapotaceae 80 20 2.82 2.96 0.74

Teijsmanniodendron Lamiaceae 72 18 2.54 1.24 0.31

Madhuca Sapotaceae 59 15 2.08 es y// 0.39

Xanthophyllum Polygalaceae 33 13 1.87 1.92 0.48

Table 6. Ten most abundant species according to number of trees recorded from the study area.

N: Number of trees. BA: Basal area. % Trees: % Trees: N/Total number of trees x 100%.

Species Family N ithees! ‘és Be ie

ha Trees (m7?) (m?/ha)

Pouteria malaccensis Sapotaceae Ros ib 4.30 Wikio Dio”

Shorea multiflora Dipterocarpaceae 84 21 2.96 3.86 0.96

Shorea macroptera Dipterocarpaceae 74 19 2.61 2.50 0.63

Ee ea Lamiaceae 64 16 «62.26 «= 0.74 —Ss«O0.18

Shorea brunnescens Dipterocarpaceae 62 16 Pag 2.04 0.51

Shorea subcylindrica Dipterocarpaceae 57 14 2.01 2.72 0.68

Hopea dyeri Dipterocarpaceae 55 14 1.94 5.84 1.46

Artocarpus integer Moraceae 49 12 1.73 2.87 0.72

moe Sig Ixonanthaceae 44. 155 194 0.49

Palaquium leiocarpum — Sapotaceae 4] 10 1.45 1723 0.31

dominated the lowest diameter class (10.0—19.9 cm), with 1315 individuals, compared

with only 314 individuals from dipterocarp species in the same diameter class. In

the largest diameter class (> 60.0 cm), dipterocarp species contributed 15 trees more

than non-dipterocarp species (Fig. 3A). The same pattern was observed in basal area

contribution of dipterocarps and non-dipterocarps, where dipterocarps dominated the

highest diameter class and at the same time contributed the highest basal area (28.21

m7’) with c. 30% higher than non-dipterocarps (Fig. 3B). The stand density by diameter

146 Gard. Bull. Singapore 64(1) 2012

classes in the study area corresponds to the condition in natural forests (Faridah-

Hanum & Philip 2006). This is considered as a general characteristic of many uneven-

aged stands and has been observed in Semenggoh Arboretum, where smaller trees

dominate the small diameter classes, and only fewer bigger trees dominated the higher

diameter classes, mostly by Dipterocarpaceae. Only a few non-dipterocarp species will

reach diameters over 60 cm, for instance Dyera costulata (Apocynaceae), Koompassia

malaccensis Maingay ex Benth. (Fabaceae), and Crudia wrayi Prain (Fabaceae).

As for the non-dipterocarps, basal area distribution decreased while diameter

increased, as most non-dipterorcarps will not reach diameter > 60.0 cm. The sudden

increase of basal area in the highest diameter class is due to the non-dipterocarp species

such as Dyera costulata (Miq.)Hook.f. (Apocynaceae), Koompassia malaccensis

(Fabaceae), and Pouteria malaccensis (Sapotaceae) that have diameters over | m, thus

contributing relatively high basal areas, although dipterocarp species dominated the

highest diameter class. Through observation, most of the emergent trees with diameter

over 100 cm are dominated by Dipterocarpaceae, where species such as Shorea

beccariana (Dipterocarpaceae), Shorea elliptica (Dipterocarpaceae), Shorea pubistyla

P.S.Ashton (Dipterocarpaceae), and Upuna borneesnis Symington (Dipterocarpaceae)

were recorded. However, there are some other species such as Dyera costulata

(Apocynaceae), Dialium spp. (Fabaceae), Koompassia malacenssis (Fabaceae) and

Pouteria malaccensis (Sapotaceae) among the non-dipterocarps that contributed to the

high basal area in Semenggoh Arboretum. This is a general characteristic of lowland

evergreen tropical rain forest, including lowland mixed dipterocarp forest (Whitmore

1984).

2000

1629

1500 4

1000 +

Number of Trees

500 +

10.0-19.9 20.0 -29.9 30.0-39.9 40.0 -49.9 $0.0-59.9 > 60.0

Diameter classes (cm)

Fig. 2. Stand density by diameter classes of all recorded trees from the 4-ha study area,

Semenggoh Arboretum.

Tree diversity at Semenggoh, Sarawak 147

Number of Trees

Basal Area (m’)

10.0-19.9 20.0-29.930.0-39.9 -40.0-49.9 $0.0-59.9 > 60.0 100-199 200-299 300-399 -40.0-499

50.0-59.9 260.0

Diameter classes (cm) Diameter classes (cm)

GDipterocarp @Non-Dipterocarp ODipterocarp ®Non-Dipterocarp

Fig. 3. Stand density (A) and basal area contribution (B) of dipterocarps and non-dipterocarps

according to diameter classes (cm).

Furthermore, more than 57% of trees in the study area are in the 10.0—-19.9 cm

diameter class. In tropical forests, the general contribution of trees within this diameter

class is about 50% of the total stand (Whitmore 1984). The high percentage of trees

in the study area suggests that natural regeneration of trees species in Semenggoh is

active. This may be due to the location of the sample plot which is near the roadside,

and is exposed to more or direct overhead light compared to those trees within the

Arboretum where relatively little light reaches the understorey seedlings and saplings.

Besides, natural disturbances such as lightning and fallen dead trees have also created

gaps within the Arboretum and promotes seedling establishment for light-demanding

species.

Diversity of dipterocarp species within the study area

In the study area, most of the trees in the bigger diameter classes are dominated by

dipterocarps. Within this family, seven genera were recorded, of which Shorea is the

dominant genus with 515 trees (71.53% from total number of trees in Dipterocarpaceae),

followed by Hopea with 86 individuals (11.91%). Both Dipterocarpus and Vatica

were represented by almost the same number of trees (1.e., 48 and 47 individuals,

respectively). The least abundant genera are Anisoptera and Upuna, each represented

by two individuals (Fig. 4). Among the 9 genera and about 267 species of dipterocarps

recorded from Borneo, 7 genera and 61 species were represented in the 4-ha study

area, that is, about 70% and 22% of dipterocarps genera and species, respectively.

According to Proctor et al. (1983) and Kochummen et al. (1991), the relative

contribution of dipterocarps within the Sunda shelf forests is 3-10% of species,

8—18% of stem number and 20-56% for basal area. In the study area, the dipterocarps

contributed more than 10% of the total species and more than 18% of the total stem

number, but the basal area contribution is within the normal range, which is about

39% (Table 2). Hence, it can be concluded that the occurrence of dipterocarp species

in the study area is relatively high and Semenggoh is an important area for research

148 Gard. Bull. Singapore 64(1) 2012

600

S15

500

400

300

Number of Trees

200 -

100

Anisoptera Cotylelobium Dipterocarpus Hopea Shorea Upuna Vatica

Fig. 4. Relative commonness of Dipterocarpaceae genera in the 4-ha study area.

on this family. Moreover, Dipterocarpaceae 1s the most important family economically

in mixed dipterocarp forest, because the most valuable timbers belong to this family

(Whitmore 1984).

The reverse-J-shaped distribution could only be observed in Shorea species

while other genera had rather random distributions among the six diameter classes.

Shorea species dominated both in the number of trees and basal area in all six diameter

classes (Fig. SA and 5B). The highest number of trees represented by this genus is in

the 10.0-19.9 cm diameter range. However, for basal area distribution, Shorea species

contributed the highest basal area in the diameter class = 60.0 cm. Overall, the basal

area of dipterocarp species increased with diameter class. This may be due to the

different growth rates and shade tolerance among the species in other genera in the

Dipterocarpaceae (Loewenstein et al. 2000). The distribution of Shorea species within

the study area indicated that regeneration and recruitment of Shorea species into larger

diameter classes were satisfactory (Maliondo et al. 2005). The diversity of dipterocarp

species in the study area is also important to serve as a seed source for reforestation

in the future. The Seedbank at Semenggoh is responsible for collecting seeds from the

Seed Production Area established in the Arboretum. The seeds collected are important

for supporting plantation, restoration, enrichment planting, urban forestry and greenery

projects.

Tree diversity at Semenggoh, Sarawak 149

Number of Trees

10.0-19.9 20.0 -29.9 30.0 -39.9 40.0-49.9 50.0-59.9 > 60.0

Diameter classes (cm)

WAnisoptera WCotylelobium ODipterocarpus S8Hopea Shorea SUpuna BVatica

16 B

Basal Area (m°)

10.0-19.9 20.0-29.9 30.0-39.9 40.0-49.9 50.0 -59.9 > 60.0

Diameter classes (cm)

@Anisoptera @Cotylelobium ODipterocarpus QHopea OShorea SUpuna OBVatica

Fig. 5. Stand distribution (A) and basal area contribution (B) of dipterocarp species according

to diameter classes in the study area.

150 Gard. Bull. Singapore 64(1) 2012

Conservation value of Semenggoh

Semenggoh is an important habitat for many threatened, rare and endemic species. The

conservation status of all tree species recorded from the study area was determined by

using the JUCN Red List of Threatened Species 2010 ((UCN 2010). Out of 541 species

recorded from the study area, about 11% of the species are considered threatened, with

20 species listed as vulnerable (VU), 14 species endangered (EN), 24 species critically

endangered (CR) and | species extinct. In addition, one species (Shorea cuspidata

P.S.Ashton), which is considered extinct in the IUCN Red List, was recorded in the

plot. Within the 4-ha study area, two individual trees of Shorea cuspidata with a basal

area of 0.067 m* were documented.

In terms of endemism, 24 species out of the total species recorded are endemic

to Sarawak, with 13 species considered rare and three species endemic to Semenggoh.

Theseincludespeciessuchas Melanochylaborneensis(Ridl.) Ding Hou(Anacardiaceae),

Actinodaphne semengohensis S. Julia (Lauraceae) and Xanthophyllum ceraceifolium

Meijden (Polygalaceae) which are restricted to Semenggoh. Approximately 20% of

the species recorded are endemic to Borneo, where 19 species are consireded rare and

only occurs in less than five localities in Sarawak; for instance, the Bornean monotypic

species, Upuna borneensis (locally known as penyau) is the largest tree documented,

with only two trees recorded in the study site. Other interesting findings include

the presence of 10 species that are uncommon in Sarawak, for example, Kibatalia

maingayi (Hook.f.) Woodson (Apocynaceae), Sindora velutina Baker (Annonaceae),

Pouteria malaccensis, Madhuca sericea (Miq.) H.J.Lam (Sapotaceae), Crudia wrayi

Prain, and Madhuca pallida (Burck) Baehni (Sapotaceae).

Besides the high endemicity of tree species in the study plot, the primary

lowland mixed dipterocarp forest of Semenggoh is in the vicinity of Kuching city, so

this area is easily accessible by land transport and numerous herbarium collections have

been made during earlier years by many taxonomists. In addition, more than 40 species

with their type localities at Semenggoh have been described. During this study, 19

species with type specimens collected from Semenggoh were recorded. These include

Swintonia minutalata Ding Hou (Anacardiaceae), Hopea kerangasensis P.S.Ashton

(Dipterocarpaceae), Shorea cuspidata P.S.Ashton, Shorea subcylindrica P.S.Ashton

(Dipterocarpaceae), Actinodaphne semengohensis, and Gonystylus micranthus Airy

Shaw (Thymelaeaceae).

Due to the uniqueness and richness of the tree species in the Semenggoh

Arboretum, more detailed studies (e.g., phenology, plant development, flowering

and fruiting onset) should be carried out in order to better understand its flora and to

propose a proper conservation management of species and habitats.

Conclusion

With 541 species and 710 trees/ha contributing to a 43.04 m*/ha basal area, the 4-ha

plot in Semenggoh can be considered a floristically rich area. As the only remaining

primary lowland mixed dipterocarp forest in the vicinity of Kuching city, this area is

Tree diversity at Semenggoh, Sarawak 151

an important site for threatened, endemic and rare species; therefore, it is critical to

protect and maintain the Arboretum in order to conserve this diverse primary mixed

dipterocarp forest into the future. At the same time, due to its high level of endemism,

the Arboretum can be an important site for future research into forest demographics

and dyamics of endangered species, which will further enhance the conservation value

of the area.

ACKNOWLEDGEMENTS. We would like to thank the Flora Malesiana Foundation and

8th Flora Malesiana secretariat for partially funding the first author to attend the symposium.

We would like to extend our gratitutde to the people who have helped in plot establishment

and enumeration work, especially Jemree Sabli, Andy Samuel, Hazlinah Suod, Tinjan Kuda,

Shalih Rebi and Army Kapi. This work would not have been possible without the support and

assistance of AFSID (Conservation and Biodiversity Research Unit) staff in fieldwork and

botanical identification in the herbarium. We would also like to acknowledge the supervision

from Ms. Lucy Chong, Mr. Sylvester Tan and Mr. Malcom Demies throughout this study, for

their comments, editing and information for this paper. Special thanks to Dr. Ruth Kiew and

Dr. Paul Chai for editing and giving constructive comments on this paper. Also thanks to GIS

Unit of the Forest Department Sarawak for providing the map of Semenggoh Forest Reserve.

This research was funded by SFC.

References

Airy Shaw, H.K. (1975) The Euphorbiaceae of Borneo. Kew Bulletin Additional Series IV, pp.

1-245. Kew: Royal Botanic Gardens, Kew.

Ashton, P.S. (2006) New Syzygium (Myrtaceae) from northern Borneo. Kew Bull. 59: 87—94.

Chung, R.C.K. (2005) Revision of Grewia (Malvaceae-Grewioidea) in Peninsular Malaysia

and Borneo. Edinburgh J. Bot. 62(1&2): 1—27.

Condit, R., Ashton, P.S., Baker, P., Bunyavejchewin, S., Gunatilleke, S., Gunatilleke, N.,

Hubbell, S., Foster, R., Itoh, A. & LaFrankie, J. (2000) Spatial patterns in the distribution

of tropical tree species. Science 288: 1414-1418.

Curran, L.M. & Webb, C.O. (2000) Experimental tests of the spatiotemporal scale of seed

predation in mast-fruiting Dipterocarpaceae. Ecol. Monogr. 70(1): 129-148.

Diway, B.M., Chai, P.K. & Blum, M. (2009) A study on the vegetation of Batang Ai National

Park, Sarawak, Malaysia (Part 2). In: /7TO Project PD 288/04 Rey. 2 (F) Development of

Lanjak Entimau Wildlife Sanctuary as a totally protected area, Phase IV (Final Phase),

47 p. International Tropical Timber Organisation, Forest Department Sarawak & Sarawak

Forestry Corporation.

Faridah-Hanum, I. & Philip, L. (2006) Tree species composition and stand attributes of Ayer

Hitam Forest Reserve, Selangor in Peninsular Malaysia. Malaysian Forester 69(2): 191—

224.

Forest Department Sarawak. (1986) Research Plot Inventory Data. Unpublished report.

Forest Department Sarawak. (1996) Research Plot Inventory Data. Unpublished report.

Forest Department Sarawak. (2000) Annual Report 2000, 62 p.

Forest Department Sarawak. (2003) Semenggoh Nature Reserve. www. forestry.sarawak. gov.

my (accessed on 2 August 2010).

152 Gard. Bull. Singapore 64(1) 2012

IUCN (2010) IUCN Red List of Threatened Species. Version 2010.2. www.iucnredlist.org.

(accessed on 2 August 2010).

Julia, S. (2005) A synopsis of the genus Actinodaphne Nees (Lauraceae) in Sabah and Sarawak,

Malaysia. Gard. Bull. Singapore 57: 69-100.

Kochummen, K.M., Manokaran, N. & LaFrankie, J.V. (1991) Floristic composition of Pasoh

Forest Reserve, a lowland rainforest of Peninsular Malaysia. J. Trop. Forest Sci. 3: 1-13.

Lee, H.S., Tan, S., Davies, S.J., LaFrankie, J.V., Yamakura, T., Itoh, A., Okhubo, T. & Harrison,

R. (2004) Lambir Forest Dynamics Plot, Malaysia. In: Losos, E.C. & Leigh, J.E.G., (eds)

Forest Diversity and Dynamism: Findings From a Large-scale Plot Network, pp. 527—

539. Chicago: Chicago University Press.

Loewenstein, E.F., Johnson, P.S. & Garrett, H.E. (2000) Age and diameter structure of a

managed uneven-aged oak forest. Canad. J. Forest Res. 30: 1060-1070.

Maliondo, S., Abeli, W., Ole Meiludie, R.E.L., Migunga, G., Kimaro, A. & Applegate, G.

(2005) Tree species composition and potential timber production of a communal Miombo

Woodland in Handei district, Tanzania. J. Trop. Forest Sci. 17(1): 104-120.

Primack, R.B. & Hall, P. (1992) Biodiversity and forest change in Malaysian Borneo.

BioScience 42(11): 829-837.

Proctor, J., Anderson, J.M., Chai, P. & Vallack, H.W. (1983) Ecological studies in four

contrasting rain forests in Gunung Mulu National Park, Sarawak. I. Forest environment,

structure and floristics. J. Ecol. 71: 237-260.

Pyke, C.R., Condit, R., Aguilar, S. & Lao, S. (2001) Floristic composition across a climatic

gradient in a neotropical lowland forest. J. Veg. Sci. 12: 553-566.

Slik, J.W.F., Poulsen, A.D., Ashton, P.S., Cannon, C.H., Eichhorn, K.A.O., Kartawinata,

K., Lanniari, I., Nagamasu, H., Nakagawa, M., Nieuwstadt, M.G.L.V., Payne, J.,

Purwaningsih, Saridan, A., Sidiyasa, K., Verburg, R.W., Webb, C.O. & Wilkie, P. (2003)

A floristic analysis of the lowland dipterocarp forests of Borneo. J. Biogeogr. 30: 1517—

Soepadmo, E. (1995) Plant diversity of the Malesian tropical rainforest and its phytogeographical

and economic significance. In: Primack, R.B. & Lovejoy, T.E. (eds) Ecology Conservation

and Management of SE Asian Rainforest, pp. 19-40. New Haven & London: Yale

University Press.

Soepadmo, E. & Chung, R.C.K. (1997) Tree Flora of Sabah and Sarawak project — a botanical

inventory of tree resources. In: Dransfield, J., Coode, M.J.E. & Simpson, D.A. (eds) Plant

Diversity in Malesia III, pp. 285-294. Kew: Royal Botanic Gardens, Kew.

Soepadmo, E. & Saw, L.G. (eds) (2000) Tree Flora of Sabah and Sarawak, vol. 3. Malaysia:

Sabah Forestry Department, Forest Research Institute Malaysia, & Sarawak Forestry

Department.

Soepadmo, E. & Wong, K.M. (eds) (1995) Tree Flora of Sabah and Sarawak, vol. 1. Malaysia:

Sabah Forestry Department, Forest Research Institute Malaysia, & Sarawak Forestry

Department.

Soepadmo, E., Wong, K.M. & Saw, L.G. (eds) (1996) Tree Flora of Sabah and Sarawak, vol.

2. Malaysia: Sabah Forestry Department, Forest Research Institute Malaysia, & Sarawak

Forestry Department.

Soepadmo, E., Saw, L.G. & Chung, R.C.K. (eds) (2002) Tree flora of Sabah and Sarawak, vol.

4. Malaysia: Sabah Forestry Department, Forest Research Institute Malaysia, & Sarawak

Forestry Department.

Soepadmo, E., Saw, L.G. & Chung, R.C.K. (eds) (2004) Tree Flora of Sabah and Sarawak, vol.

5. Malaysia: Sabah Forestry Department, Forest Research Institute Malaysia, & Sarawak

Forestry Department.

Tree diversity at Semenggoh, Sarawak 153

Soepadmo, E., Julia, S., Soh, W.K. & Yahud, W. (2006) Tree Flora of Sabah and Sarawak —

Species Diveristy and Endemism. In: Ho, Y.F. & Sarifah, K.A. (eds) Highlights of FRIM’s

Non-IRPA Projects 2006, pp. 35-52. Kepong: Forest Research Insitute Malaysia.

Soepadmo, E., Saw, L.G., Chung, R.C.K. & Kiew, R. (eds) (2007) Tree Flora of Sabah

and Sarawak, vol. 6. Malaysia: Sabah Forestry Department, Forest Research Institute

Malaysia, & Sarawak Forestry Department.

Sugau, J.B. (2005) Twelve new species of Adinandra (Pentaphylacaceae) from Borneo.

Sandakania 16: 1-27.

Tan, K.H., Zubaid, A. & Kunz, T.H. (1998) Food habits of Cynopterus brachyotis Muller

(Chiroptera: Pteropodidae) in Peninsular Malaysia. J. Trop. Ecol. 14: 299-307.

Tan, S. (2002) Sustainable Management of Peat Swamp Forest of Sarawak with Special

Reference to Ramin. Manual for Establishment and Enumeration of Permanent Sample

Plots in Peat Swamp Forest in Sarawak. Joint Working Group Malaysia - The Netherlands

Project.

Whitmore, T.C. (1984) Tropical Rain Forests of the Far East, 2nd ed. Oxford: Oxford

University Press.

Appendix A. Composition, stand density and basal area (BA) contribution of tree families

from the 4-ha study site, Semenggoh.

No. Family Genus Species Total BA (m°)

1 Achariaceae 3 1] 44 1.98

2 Anacardiaceae 9 18 94 6.74

3. Anisophylleaceae l 3 15 0.54

4 Annonaceae 5 11 49 LES

5 Apocynaceae 3 + 28 4.85

6 Araliaceae l l 2 0.04

7 Burseraceae 3 28 106 Spl?

8 Cardiopteridaceae | l 18 0.39

9 Celastraceae 3 3) 1] 0.24

10 Chrysobalanaceae 3 4 6 0.10

11 Clusiaceae 4 38 84 3.74

12 Combretaceae | ] 2 0.16

13. Cornaceae ] 2 D, 0.05

14. Crypteroniaceae l | l 0.04

15 Ctenolophonaceae l l 3 0.32

16 Dilleniaceae | 5 21 0.65

17 Dipterocarpaceae di 61 720 66.86

18 Ebenaceae | 16 39 1.84

19 Elaeocarpaceae 1 6 7 0.20

20 Euphorbiaceae 10 16 61 1.41

21 Fabaceae a 15 73 10.20

22 Fagaceae 3 8 14 N79

154 Gard. Bull. Singapore 64(1) 2012

No. Family Genus Species Total BA (m’)

23 Icacinaceae ] ] ] 0.01

24 Ixonanthaceae D, 2 50 DAS

25 lLamiaceae D, 6 73 1.26

26 Lauraceae V 18 49 2.30

27 Lecythidaceae l 2, di 0.41

28 Loganiaceae l l 37 3.28

29 Magnoliaceae 2 3 0.35

30 Malvaceae 10 29 125 7.46

31 Melastomataceae 2 2 1] 0.16

32 Meliaceae 7 23 55 1.62

33. Memecylaceae 1 6 9 0.22

34 Moraceae 3 1] 160 7.88

- 35 Myristicaceae 4 42 216 6.58

36 Myrtaceae l 29 114 18

37 Ochnaceae 1 l 1 0.04

38 Olacaceae 2 3 1] 0.55

39 Oleaceae l 2 4 0.06

40 Oxalidaceae 1 l 4 1.45

41 Peraceae 2 D, 13 0.37

42 Phyllanthaceae y 16 42 0.64

43 Polygalaceae ] 2 35) E92

44 Proteaceae ] l 2 0.05

45 Putranjivaceae | 4 10 0.30

46 Rhizophoraceae 2 2 3 0.07

47 Rosaceae ] 3 4 0.07

48 Rubiaceae 6 6 17 0.25

49 Rutaceae 3 3 10 0.22

50 Salicaceae ] | ] 0.03

51 Sapindaceae 4 7 14 1.00

52 Sapotaceae 4 27 262 W329

53 Simaroubaceae l ] 3 0.39

54 Staphyleaceae | 2 2 0.02

55 Stemonuraceae 2 3 3 0.21

56 Symplocaceae l 3 J 0.04

57 Theaceae 3 4 8 0.32

58 Thymelaeaceae 2 3) 14 0.43

59 Ulmaceae l 2 35 179

60 Unidentified ] | 1 0.04

Tol “e** Ye0 oe 2837 172.16.

Tree diversity at Semenggoh, Sarawak L55

Appendix B. Checklist of trees from the 4-ha study site in Semenggoh Arboretum. (Only

species that were listed in The IUCN Red List of Threatened Species 2010 are specially indicated

with conservation status in the notes.)

Achariaceae

Hydnocarpus anomala (Merr.) Sleumer

Hydnocarpus borneensis Sleumer

Hydnocarpus kunstleri (King) Warb. var. tomentosa (King) Sleumer

Hydnocarpus pinguis Sleumer

Hydnocarpus polypetala (Sloot.) Merr.

Hydnocarpus subfalcata Mert.

Hydnocarpus tenuipetala Sleumer

Hydnocarpus woodii Merr.

Ryparosa hirsuta J.J.Sm.

Ryparosa hullettii King

Trichadenia philippinensis Mert.

Anacardiaceae

Androtium astylum Stapf

Buchanania arborescens (Blume) Blume

Campnosperma squamatum Ridl.

Drimycarpus luridus (Hook.f.) Ding Hou Notes: Rare in Sarawak (only from two other

localities in Tatau & Belaga districts).

Gluta beccarii (Engl.) Ding Hou

Gluta oba (Merr.) Hou. Notes: Endemic to Borneo.

Gluta wallichii (Hook.f.) Ding Hou

Mangifera griffithii Hook.f. Notes: Uncommon in Sarawak.

Mangifera parvifolia Boerl. & Koord. Notes: Uncommon in Sarawak, usually from peat

swamp forest.

Melanochyla angustifolia Hook.f. Notes: Uncommon in Sarawak.

Melanochyla beccariana Oliv. Notes: Endemic to Borneo but uncommon in Sarawak.

Melanochyla borneensis (Ridl.) Ding Hou Notes: Endemic to Sarawak and so far only known

from Kuching districts.

Melanochyla elmeri Mert. Notes: Endemic to Borneo but uncommon in Sarawak.

Melanochyla fulvinervis (Blume) Ding Hou Notes: Uncommon in Sarawak.

Parishia insignis Hook.f. Notes: Not endemic but uncommon in Sarawak.

Parishia maingayi Hook.f.

Swintonia glauca Engl.

Swintonia minutalata Ding Hou Notes: Endemic to Borneo and uncommon in Sarawak

(Semenggoh & Bako only); type specimen from Semenggoh.

Anisophylleaceae

Anisophyllea beccariana Baill Notes: Not endemic but in Sarawak, only found in Kuching &

Bintulu district.

Anisophyllea corneri Ding Hou

Anisophyllea ferruginea Ding Hou Notes: Uncommon in Sarawak (only known from 4 districts

in Sarawak).

156 Gard. Bull. Singapore 64(1) 2012

Annonaceae

Alphonsea javanica Blume

Alphonsea johorensis J. Sinclair

Cyathocalyx biovulatus Boerlage

Cyathocalyx carinatus (Ridl.) J.Sinclair

Mezzettia parviflora Becc.

Polyalthia hookeriana King

Polyalthia lateriflora King

Xylopia caudata Hook.f. & Thomson

Xylopia elliptica Maingay ex Hook.f.

Xylopia ferruginea Baill.

Xylopia malayana Hook.f. & Thomson

Apocynaceae

Alstonia angustifolia Wall. ex A. DC.

Alstonia angustiloba Miq. Notes: Not endemit but only found in western part of Sarawak.

Dyera costulata (Miq.) Hook.f.

Kibatalia maingayi (Hook.f.) Woodson Notes: Not endemic but very localised in Sarawak

(Semenggoh & Bau only).

Araliaceae

Arthrophyllum diversifolium Blume

Burseraceae

Canarium apertum H.J.Lam

Canarium dichotomum (Blume) Miq.

Canarium littorale Blume forma littorale

Canarium littorale Blume forma rufum (A.W.Benn.) Leenh. Notes: Less common in Sarawak.

Canarium megalanthum Mert.

Canarium merrillii H.J.Lam Notes: Endemic to Borneo.

Canarium odontophyllum Miq.

Canarium pilosum A. W. Benn. subsp. pilosum

Dacryodes costata (A. W. Benn.) H.J.Lam

Dacryodes incurvata (Engl.) H.J.Lam Notes: From 4 districts in Sarawak.

Dacryodes longifolia (King) H.J.Lam Notes: Less common in Sarawak.

Dacryodes macrocarpa H.J.Lam var. patentinervia Leenh

Dacryodes nervosa (H.J.Lam) Leenhouts Notes: Not endemic but in Sarawak, only found in

Tubau, Bako, Bintulu & Sadong.

Dacryodes rostrata (Blume) H.J.Lam forma cuspidata (Blume) H.J.Lam

Dacryodes rugosa (Blume) H.J.Lam var. rugosa

Dacryodes rugosa (Blume) H.J.Lam var. virgata (Blume) H.J.Lam

Dacryodes sp. |

Santiria apiculata A.W. Benn. var. apiculata

Santiria apiculata Benn. var. pilosa (Engl.) Kalkman

Santiria griffithii (Hook.f.) Engl.

Santiria laevigata Blume forma glabrifolia (Engl.) H.J.Lam

Santiria megaphylla Kalkman Notes: Endemic to Borneo but uncommon in Sarawak

(Semenggoh & Dulit).

Tree diversity at Semenggoh, Sarawak 157

Santiria mollis Eng|. Notes: Endemic to Borneo but uncommon in Sarawak (Semenggoh &

Lambir, Niah, Bukit Raya).

Santiria oblongifolia Blume

Santiria sarawakana Kochummen Notes: Endemic to Sarawak and only from Miri & Sabal.

Santiria tomentosa Blume

Santiria sp. |

Santiria sp. 2

Cardiopteridaceae

Gonocaryum minus Sleumer

Celastraceae

Bhesa paniculata Arn.

Kokoona littoralis Laws. var. longifolia Kochummen

Lophopetalum glabrum Ding Hou Notes: Endemic to Borneo.

Chrysobalanaceae

Atuna nannodes (Kosterm.) Kosterm. Notes: Not Endemic but uncommon in Sarawak (Bau

& Kuching only).

Atuna sp. |

Kostermantus heteropetalus Notes: Sarawak only recorded from 4 districts.

Licania splendens (Korth.) Prance

Clusiaceae

Calophyllum biflorum M.R.Hend. & Wyatt-Sm.

Calophyllum castaneum P.F.Stevens

Calophyllum elegans Ridley

Calophyllum lanigerum Mig. var. austrocoriaceum (Whitmore) P.F.Stevens

Calophyllum molle King

Calophyllum pulcherrinum Wall. ex Choisy

Calophyllum roseocostatum P.F.Stevens Notes: Type specimen from Semenggoh.

Calophyllum soulattri Burm.f. Notes: LR/Ic ver 2.3

Calophyllum tetrapterum Miq. Notes: LR/Ic ver 2.3

Calophyllum teysmannii Zoll. ex Planch. & Tiana

Calophyllum woodii P.F.Stevens

Calophyllum sp. |

Cratoxylum arborescens (Vahl) Blume Notes: LR/Ic ver 2.3

Cratoxylum formosum (Jack) Dyer Notes: LR/Ic ver 2.3

Cratoxylum maingayi Dyer Notes: LR/Ic ver 2.3

Garcinia beccarii Pierre

Garcinia blumei Pierre

Garcinia calophyllifolia Ridl.

Garcinia caudiculata Ridl.

Garcinia celebica L.

Garcinia cuspidata King

Garcinia drvobalanoides Pierre

Garcinia maingayi Hook.f. Notes: LR/Ic ver 2.3

Garcinia mangostana L.

158 Gard. Bull. Singapore 64(1) 2012

Garcinia myristicaefolia Pierre

Garcinia nervosa Miq.

Garcinia nitida Pierre

Garcinia parvifolia (Miq.) Miq.

Garcinia vidua Ridl.

Garcinia sp. 39°

Garcinia sp. ‘AZ’

Mesua beccariana (Baill.) Kosterm.

Mesua borneensis P.F.Stevens

Mesua congestiflora P.F.Stevens

Mesua elmeri (Merr.) Kosterm.

Mesua macrantha (Baill.) Kosterm.

Mesua myrtifolia (Baill.) Kosterm.

Mesua variabilis P.F.Stevens

Combretaceae

Terminalia foetidissima Griff.

Cornaceae

Alangium javanicum (Blume) Wangerin var. javanicum

Alangium javanicum var. meyeri (Metr.) Berhaman

Crypteroniaceae

Crypteronia griffithii C.B.Clarke

Ctenolophonaceae

Ctenolophon parvifolius Oliv.

Dilleniaceae

Dillenia beccariana Martelli

Dillenia excelsa (Jack) Gilg

Dillenia excelsa (Jack) Gilg. var. tomentella (Martelli) Masamune

Dillenia reticulata King

Dillenia sumatrana Miq.

Dipterocarpaceae

Anisoptera grossivenia Sloothen Notes: EN Alcd+2ced ver 2.3; endemic to Borneo.

Cotylelobium melanoxylon (Hook.f.) Pierre Notes: EN Alcd+2cd ver 2.3

Dipterocarpus confertus Slooten Notes: Endemic to Borneo.

Dipterocarpus geniculatus Vesque subsp. geniculatus Notes: Endemic to Borneo.

Dipterocarpus globosus Vesque Notes: CR Alcd+2cd, B1+2c ver 2.3; endemic to Borneo.

Dipterocarpus lowii Hook.f. Notes: CR Aled+2ced, B1+2c ver 2.3

Dipterocarpus sarawakensis F.G.Browne ex Slooten Notes: Type specimen from Semenggoh.

Hopea beccariana Burck Notes: CR Alcd+2ed, B1+2c ver 2.3

Hopea bracteata Burck

Hopea dyeri F.Heim

Hopea griffithii Kurz Notes: VU Alct2c ver 2.3

Hopea kerangasensis P.S.Ashton Notes: CR Alcd+2cd, B1+2c ver 2.3; type specimen from

Semenggoh.

Tree diversity at Semenggoh. Sarawak 159

Hopea latifolia Symington Notes: CR Alc, B1+2c ver 2.3

Hopea longirostrata P.S.Ashton Notes: CR Aled, B1+2c, C1, D ver 2.3: endemic to Sarawak.

Hopea pachycarpa (F.Heim) Symington Notes: VU Alcd+2cd, B1+2c ver 2.3

Hopea sphaerocarpa (F.Heim) P.S.Ashton Notes: CR Alc, B1+2c ver 2.3; endemic to Sarawak.

first and second Division only.

Hopea tenuinervula P.S.Ashton Notes: CR Alc, B1+2c ver 2.3: endemic to Borneo.

Shorea atrinervosa Symington

Shorea beccariana Burck Notes: Endemic to Borneo.

Shorea bracteolata Dyer Notes: EN Alcd+2cd ver 2.3

Shorea brunnescens P.S.Ashton Notes: EN Alcd+2cd, C2a ver 2.3; endemic to Borneo.

Shorea crassa P.S.Ashton Notes: Endemic to Borneo; type specimen from Semenggoh.

Shorea cuspidata P.S. Ashton Notes: EX ver 2.3; endemic to Borneo, type specimen from

Semenggoh.

Shorea dasyphylla Foxw. Notes: EN Alcd ver 2.3

Shorea elliptica Burck Notes: CR Alcd, C2a ver 2.3; endemic to Borneo, uncommon in

Sarawak (Kuching & Lundu only).

Shorea falcifera Dyer ex Brandis Notes: EN Alcd ver 2.3

Shorea flemmichii Symington Notes: CR Alcd, C2a ver 2.3; endemic to Sarawak.

Shorea isoptera P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo.

Shorea kunstleri King Notes: CR Alcd ver 2.3

Shorea ladiana P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo, type specimen

from Semenggoh.

Shorea leprosula Miq. Notes: EN Alcd ver 2.3

Shorea lunduensis P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo; uncommon

in Sarawak (only from Lundu, Bau & Kuching Marudi districts).

Shorea macrobalanos P.S.Ashton Notes: CR Aled, C2a ver 2.3; endemic to Borneo, uncommon

in Sarawak (only from Kapit , Bintulu & Lundu districts).

Shorea macrophylla (de Vries) P.S.Ashton Notes: VU Alcd ver 2.3; endemic to Borneo.

Shorea macroptera Dyer

Shorea maxwelliana King Notes: EN Alc ver 2.3

Shorea multifiora (Burck) Symington Notes: LR/Ic ver 2.3

Shorea myrionerva Sym. ex P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo.

Shorea obscura Meijer Notes: EN Alcd ver 2.3; endemic to Borneo.

Shorea ovalis (Korth.) Blume subsp. sarawakensis P.S.Ashton Notes: Endemic to Borneo.

Shorea parvifolia Dyer subsp. velutina P.S.Ashton

Shorea pubistyla P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo, type specimen

from Semenggoh.

Shorea quadrinervis Slooten Notes: EN Alcd ver 2.3: endemic to Borneo.

Shorea richetia P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo, uncommon in

Sarawak (Kuching & Lundu only).

Shorea rubra P.S.Ashton Notes: Endemic to Borneo.

Shorea scaberrima Burck Notes: Endemic to Borneo.

Shorea scabrida Symington

Shorea seminis (de Vriese) Slooten Notes: CR Alcd ver 2.3

Shorea slootenii Wood ex P.S.Ashton Notes: CR Alcd, C2a ver 2.3; endemic to Borneo.

Shorea subcylindrica Slooten Notes: CR Alcd, C2a ver 2.3; endemic to Borneo, type specimen

from Semenggoh.

Upuna borneensis Symington Notes: EN Alcd, C2a ver 2.3: endemic to Borneo.

160 Gard. Bull. Singapore 64(1) 2012

Vatica badiifolia P.S.Ashton Notes: EN Alcd ver 2.3; endemic to Borneo.

Vatica coriacea P.S.Ashton Notes: CR Alc, C2a ver 2.3; endemic to Borneo, uncommon (in

Sarawak known from Lundu & Kuching districts).

Vatica dulitensis Symington Notes: Endemic to Borneo.

Vatica granulata Slooten subsp. granulata Notes: Endemic to Borneo.

Vatica havilandii Brandis Notes: CR Alc, C2a ver 2.3

Vatica maingayi Dyer Notes: CR Alcd, C2a ver 2.3

Vatica micrantha Slooten Notes: Endemic to Borneo.

Vatica nitens (King) Kosterm.

Vatica oblongifolia Hook.f. subsp. oblongifolius Notes: Endemic to Borneo.

Vatica pedicellata Brandis Notes: EN Alc ver 2.3; endemic to Sarawak, in Kuching, Lundu,

Simunjan only.

Ebenaceae

Diospyros borneensis Hiern

Diospyros curranii Mert.

Diospyros dictyoneura Hiern

Diospyros ferruginescens Bakh. var. ferruginescens Notes: Endemic to Borneo.

Diospyros lunduensis Ng Notes: Endemic to Borneo, uncommon in Sarawak (Lundu &

Semenggoh only).

Diospyros maingayi (Hiern) Bakh.

Diospyros mindanaensis Mert.

Diospyros neurosepala Bakh. Notes: Endemic to Borneo.

Diospyros pendula Hasselt ex Hassk.

Diospyros pilosanthera Blanco var. pilosanthera

Diospyros puncticulosa Bakh. Notes: Endemic to Borneo.

Diospyros ridleyi Bakh.

Diospyros stvraciformis King & Gamble

Diospyros venosa Wall. ex A. DC. var. venosa

Diospyros sp. |

Diospyros sp. 2

Elaeocarpaceae

Elaeocarpus beccarii DC.

Elaeocarpus clementis Merr.

Elaeocarpus cristatus Coode

Elaeocarpus dolichobotrys Mert.

Elaeocarpus griffithii (Wight) A.Gray

Elaeocarpus petiolatus (Jack) Wall.

Euphorbiaceae

Blumeodendron calophyllum Airy Shaw

Blumeodendron tokbrai (Bl.) Kurz

Cephalomappa beccariana Baill. var. beccariana Notes: Endemic to Sarawak.

Croton argyratus Blume

Endospermum diadenum (Miq.) Airy Shaw

Macaranga bancana (Miq.) Muell.Arg.

Macaranga conifera Muell.Arg.

Tree diversity at Semenggoh, Sarawak 161

Macaranga lowii King ex Hook.f.

Macaranga recurvata Gage

Macaranga triloba (Thunb.) Muell.Arg.

Mallotus griffithianus (Muell.Arg.) Hook.f.

Mallotus penangensis Muell.Arg.

Neoscortechinia kingii (Hook.f.) Pax & Hoffm.

Pimeleodendron griffithianum (Muell.Arg.) Benth.

Ptychopyxis costata Miq. var. oblanceolata Airy Shaw

Sebastiania borneensis Pax & Hoffm.

Fabaceae

Archidendron borneense (Benth.) I.C. Nielsen

Archidendron microcarpum (Benth.) I.C. Nielsen

Crudia wrayi Prain Notes: Not endemic but uncommon in Sarawak (Kuching only).

Dialium indum L. var. bursa (de Wit) Rojo

Dialium indum L. var. indum

Dialium kunstleri Prain var. kunstleri

Dialium platysepalum Baker

Dialium sp. |

Dialium sp. 2

Koompassia malaccensis Maing. ex Benth.

Parkia sumatrana Miq.

Pithecellobium kunstleri Prain

Sindora irpicina de Wit

Sindora leiocarpa Backer ex de Wit

Sindora velutina Baker

Sindora sp. |

Fagaceae

Castanopsis costata (Blume) A. DC.

Castanopsis hypophoenicea (Seemen) Soepadmo Notes: Endemic to Borneo.

Castanopsis motleyana King

Lithocarpus andersonii Soepadmo Notes: Endemic to Borneo.

Lithocarpus coopertus (Blanco) Rehder

Lithocarpus gracilis (Korth.) Soepadmo

Lithocarpus pulcher (King) Markgr. Notes: Endemic to Borneo.

Quercus kerangasensis Soepadmo Notes: Endemic to Borneo; uncommon in Sarawak (in Ulu

Baram, Kuching, Simunjan).

Icacinaceae

Platea latifolia Blume Notes: Endemic to Sarawak (4 localities only).

Ixonanthaceae

Allantospermum borneense Forman subsp. borneense Notes: Type specimen from Semenggoh.

Ixonanthes petiolaris Blume

Lamiaceae

Clerodendrum villosum Blume Notes: Endemic to Sarawak (6 localities only).

162 Gard. Bull. Singapore 64(1) 2012

Teijsmanniodendron bintuluensis Moldenke

Teijsmanniodendron havilandii (Ridl|.) G.Rusea Notes: Endemic to Sarawak (Sadong &

Semenggoh only).

Teijsmanniodendron sarawakanum (H.H.W.Pearson) Kosterm. Notes: Endemic to Borneo.

Teijsmanniodendron simplicifolium Merr. Notes: Endemic to Borneo.

Teijsmanniodendron subspicatum (H.Hallier) Kosterm.

Lauraceae

Actinodaphne macrophylla (Blume) Nees

Actinodaphne semengohensis S.Julia Notes: Endemic to Sarawak; only from Semenggoh, type

specimen from Semenggoh.

Beilschmiedia kunstleri Gamble

Beilschmiedia maingayi Hk.f.

Cryptocarya ferrea Blume var. ferrea

Cryptocarya ferrea Blume var. scortechinii (Gamble) Ng

Cryptocarya griffithiana Wight var. crassinervia (Miq.) Ng

Endiandra clavigera Kosterm. Notes: Endemic to Sarawak.

Endiandra coriacea Mert.

Endiandra rubescens (Blume) Miq.

Lindera lucida (Blume) Boerl.

Litsea accedens (Blume) Boerl. var. oblanceolata (Gamble) Ng

Litsea gracilipes Hook.f.

Litsea grandis Hook.f.

Litsea lancifolia (Roxb. ex Wall.) Hook.f.

Litsea resinosa Blume

Notaphoebe sp. |

Notaphoebe sp. 2

Lecythidaceae

Barringtonia lanceolata (Ridl.) Payens Notes: Endemic to Borneo.

Barringtonia sarcostachys (Blume) Miq.

Loganiaceae

Norrisia maior Soler.

Magnoliaceae

Magnolia ashtonii Dandy ex Noot.

Magnolia borneensis Noot.

Talauma beccarii Ridl.

Malvaceae

Brownlowia ovalis Kosterm. Notes: Type specimen from Semenggoh.

Durio grandiflorus (Most.) Kosterm. Notes: VU Alc ver 2.3

Durio graveolens Becc.

Durio kutejensis (Hassk.) Becc. Notes: VU Alc ver 2.3

Durio lanceolatus Mast.

Durio malaccensis Planch ex Ma

Durio oblongus Mast.

Tree diversity at Semenggoh, Sarawak 163

Durio testudinarius Becc. Notes: VU Alc ver 2.3

Grewia gracilis (Stapf ex Ridl.) P.S. Ashton

Heritiera albiflora Kosterm.

Heritiera aurea (Miq.) Kosterm.

Heritiera javanica (Blume) Kosterm.

Heritiera simplicifolia (Mast.) Kosterm.

Heritiera sumatrana (Miq.) Kosterm.

Microcos cinnamomifolia Burret Notes: Endemic to Borneo.

Microcos triflora (Blanco) R.C.K.Chung var. triflora

Neesia piluliflora Becc.

Pentace borneensis Pierre

Pentace curtisii King

Pentace polyantha Hassk.

Pentace rigida Kosterm.

Scaphium affine (Mast.) Pierre

Scaphium macropodum (Miq.) Beumée ex Heyne

Schoutenia accrescens (Mast.) C.H.Curtis

Schoutenia accrescens (Mast.) C.H.Curtis subsp. ste//ata Roekmowati-Hartono

Sterculia oblongata R.Br.

Sterculia parvifolia Wall. ex R.Br.

Sterculia scortechinii King

Sterculia sp. |

Melastomataceae

Lijndenia laurina Zoll. & Moritzi

Pternandra hirtella (Cogn.) Nayar

Meliaceae

Aglaia argentea Blume

Aglaia exstipulata (Griff.) W.Theob. subsp. brunneostellata Pannell

Aglaia forbesii King

Aglaia hiernii King

Aglaia leucophylla King

Aglaia malaccensis (Ridl.) J.A.R.Anderson

Aglaia meliosmoides Craib

Aglaia odoratissima Blume

Aglaia ramotricha Pannell Notes: Endemic to Borneo.

Aglaia rubiginosa (Hiern) Pannell

Aglaia sexipetala Griff.

Aglaia speciosa Blume

Aglaia squamulosa King

Aglaia sp. |

Chisocheton pentandrus (Blanco) Merr. subsp. paucijugus (Miq.) Mabb.

Dysoxylum densiflorum (Blume) Miq.

Dysoxylum magnificum Mabb.

Dysoxylum pachyrhache Mert. Notes: Endemic to Borneo.

Pseudoclausena chrysogyne (Miq.) T.P.Clark

Reinwardtiodendron humile (Hassk.) Mabb.

164 Gard. Bull. Singapore 64(1) 2012

Sandoricum borneense Mig. Notes: Endemic to Borneo.

Sandoricum dasyneuron Baill. Notes: Endemic to Borneo.

Walsura sp. A

Memecylaceae

Memecylon amplexicaule Roxb.

Memecylon argenteum K.Bremer

Memecylon campanulatum C.B.Clarke

Memecylon durum Cogn.

Memecylon garcinioides Blume

Memecylon scolopacinum Ridl.

Moraceae

Artocarpus anisophyllus Miq.

Artocarpus integer (Thunb.) Merr.

Artocarpus kemando Miq.

Artocarpus longifolius Becc. Notes: Endemic to Borneo and rare.

Artocarpus nitidus Tréc.

Artocarpus obtusus Jarrett Notes: Endemic to Borneo and rare; in Sarawak only from 4

locations, type specimen from Semenggoh.

Artocarpus odoratissimus Blanco Notes: Endemic to Borneo.

Artocarpus peltatus Merr. Notes: Endemic to Borneo.

Artocarpus rigidus Blume

Parartocarpus venenosus (Zoll. & Moritzi) Becc. subsp. borneensis (Becc.) Jarrett Notes:

Endemic to Borneo.

Prainea frutescens Becc. Notes: Endemic to Borneo and uncommon, Sarawak only from west

& northern parts (4 localities only).

Myristicaceae

Gymnacranthera bancana (Miq.) J. Sinclair

Gymnacranthera contracta Warb. Notes: Endemic to Borneo.

Gymnacranthera farquhariana (Hook.f. & Thomson) Warb. var. eugeniifolia (A.DC.)

R.T.A.Schouten

Gymnacranthera farquhariana (Hook.f. & Thomson) Warb. var. farquhariana

Gymnacranthera forbesii (King) Warb. var. crassinervis (Warb.) J. Sinclair Notes: Endemic

to Borneo.

Gymnacranthera ocellata Schouten Notes: Endemic to Borneo.

Horsfieldia androphora W.J. de Wilde Notes: Endemic to Borneo.

Horsfieldia borneensis W.J. de Wilde Notes: Endemic to Borneo; type specimen from

Semenggoh.

Horsfieldia brachiata (King) Warb.

Horsfieldia grandis (Hook.f.) Warb.

Horsfieldia laticostata (J.Sinclair) W.J.de Wilde Notes: Endemic to Borneo.

Horsfieldia nervosa W.J.de Wilde Notes: Endemic to Sarawak and rare.

Horsfieldia pallidicaula W.J.de Wilde var. pallidicaula Notes: Endemic to Borneo.

Horsfieldia polyspherula (Hook.f. emend. King) J. Sinclair var. maxima W.J.de Wilde Notes:

Endemic to Borneo.

Horsfieldia punctatifolia J.Sinclair

Tree diversity at Semenggoh, Sarawak 165

Horsfieldia tenuifolia (J.Sinclair) W.J.de Wilde Notes: Endemic to Borneo.

Horsfieldia wallichii (Hook.f. & Thomson) Warb.

Knema curtisii (King) Warb. var. arenosa

Knema curtisii (King) Warb. var. curtisii

Knema galeata J.Sinclair Notes: Endemic to Borneo.

Knema glaucescens Jack

Knema /atericia Elmer subsp. albifolia (Sinclair) W.J.de Wilde Notes: Endemic to Borneo.

Knema latericia Elmer subsp. ridleyi (Gand.) W.J.de Wilde

Knema latifolia Warb.

Knema lunduensis (Sinclair) W.J.de Wilde Notes: Endemic to Borneo.

Knema membranifolia H.J.P.Winkler Notes: Endemic to Borneo.

Knema pedicellata W.J.de Wilde Notes: Endemic to Borneo.

Knema pericoriacea Sinclair forma sarawakensis Notes: Endemic to Borneo.

Knema rufa Warb. Notes: Endemic to Borneo.

Knema stenophylla (Warb.) J.Sinclair subsp. longipedicellata (Sinclair) W.J.de Wilde Notes:

Endemic to Borneo.

Knema viridis W.J.de Wilde Notes: Endemic to Sarawak.

Knema sp. |

Knema sp. 2

Knema sp. 3

Myristica beccarii Warb.

Myristica borneensis Warb. Notes: Endemic to Borneo.

Myristica cinnamomea King

Myristica crassa King

Myristica gigantea King

Myristica iners Blume

Myristica malaccensis Hook.f. subsp. papillosa W.J.de Wilde Notes: Endemic to Borneo.

Myristica villosa Warb. Notes: Endemic to Borneo.

Myrtaceae

Syzygium accuminatissinum A. de Camdolle

Syzygium adenophyllum Merrill & Perry Notes: Endemic to Borneo.

Syzygium borneense (Miq.) Miq.

Syzygium brachyrachis Merr. & Perry Notes: Endemic to Borneo.

Syzygium chloranthum (Duthie) Merrill & Perry

Syzygium durifolium Merrill & Perry Notes: Endemic to Borneo and rare, in Sarawak from

Semenggoh only.

Syzygium glabratum Veldkamp

Syzygium gladiatum (Ridl.) Merrill & Perry Notes: Endemic to Borneo and rare; in Sarawak

from Sabal & Semenggoh.

Syzygium glanduligerum (Ridl.) Merrill & Perry Notes: Endemic to Borneo.

Syzygium grande (Wight) Walpers

Syzygium hoseanum (King) Merrill & Perry

Syzygium incarnatum (Elmer) Merrill & Perry

Syzygium inophyllum A. de Candolle

Syzygium kunstleri (King) Bahadur & R.C. Gaur

Syzygium leptostemon (Korth.) Merrill & Perry

Syzygium longiflorum Pres.

166 Gard. Bull. Singapore 64(1) 2012

Syzygium napiforme (Koord. & Valeton) Merrill & Perry

Syzygium nemestrinum (Henderson) I.M.Turner

Syzygium oligomyrum Diels Notes: Endemic to Borneo.

Syzygium pendens (Duthie) I.M.Turner

Syzygium polyanthum (Wight) Walpers

Syzygium pyrifolium (Blume) A.de Candolle

Syzygium remotifolium (Ridl.) Merrill & Perry Notes: Endemic to Borneo.

Syzygium ridleyi (King) P. Chantaranothai & J. Parn

Syzygium rostratum (Blume) A.de Candolle

Syzygium rugosum Korth.

Syzygium urceolatum Merrill & Perry ssp. kuchingense (Merrill) P.S.Ashton Notes: Endemic

to Borneo.

Syzygium urceolatum Merrill & Perry ssp. palembanicum P.S.Ashton

Syzygium sp. |

Syzygium sp. 2

Ochnaceae

Gomphia serrata (Gaertn.) Kanis

Olacaceae

Anacolosa frutescens (Blume) Blume

Strombosia ceylanica Gardner

Oleaceae

Chionanthus havilandii Kiew Notes: Endemic to Sarawak (from 6 localities only).

Chionanthus lucens Kiew

Oxalidaceae

Sarcotheca diversifolia (Miq.) Hallier f.

Peraceae

Chaetocarpus castanocarpus (Roxb.) Thw.

Trigonopleura malayana Hook.f.

Phyllanthaceae

Aporosa bullatissima Airy Shaw Notes: Endemic to Borneo; type specimen from Semenggoh.

Aporosa elmeri Merr. Notes: Endemic to Borneo.

Aporosa illustris Airy Shaw Notes: Endemic to Sarawak.

Aporosa lucida (Miq.) Airy Shaw var. lucida

Aporosa nitida Merr. Notes: Endemic to Borneo.

Aporosa subcaudata Mert.

Baccaurea macrophylla (Muell.Arg.) Muell.Arg.

Baccaurea maingayi Hook.f.

Baccaurea minor Hook.f.

Baccaurea pyriformis Gage

Baccaurea racemosa (Reinw. ex Blume) Muell.Arg.

Baccaurea reticulata Hook.f.

Baccaurea sarawakensis Pax & Hoffm.

Tree diversity at Semenggoh, Sarawak 167

Baccaurea sumatrana (Miq.) Muell.Arg.

Cleistanthus coriaceus Airy Shaw Notes: Endemic to Sarawak, type specimen from Semenggoh

Cleistanthus pseudopodocarpus Jab).

Polygalaceae

Xanthophyllum brevipes Meijden Notes: Endemic to Borneo; from 6 localities in Sarawak

(Type specimen from Semenggoh)

Xanthophyllum ceraceifolium Meijden Notes: Endemic to Sarawak; from Semenggoh only

(Type specimen from Semenggoh)

Xanthophyllum ecarinatum Chodat Notes: Endemic to Borneo.

Xanthophyllum ellipticum Miq.

Xanthophyllum ferrugineum Meijden Notes: Endemic to Borneo.

Xanthophyllum flavescens Roxb.

Xanthophyllum griffithii A.W.Benn. var. angustifolium Ng

Xanthophyllum parvifolium Meijden Notes: Endemic to Borneo.

Xanthophyllum rufum A.W.Benn.

Xanthophyllum stipitatum A.W.Benn.

Xanthophyllum trichocladum Chodat Notes: Endemic to Borneo; uncommon in Sarawak

(Kapit & Kuching only).

Xanthophyllum velutinum Chodat Notes: Endemic to Borneo.

Proteaceae

Helicia petiolaris Benn.

Putranjivaceae

Drypetes crassipes Pax & Hoffm.

Drypetes eriocarpa Airy Shaw Notes: Endemic to Sarawak, from Bukit Raya, Lambir, Sabal

& Semenggoh.

Drypetes longifolia Pax & Hoffm.

Drypetes macrostigma J.J.Sm. Notes: Endemic to Borneo.

Rhizophoraceae

Carallia brachiata (Lour.) Merr.

Pellacalyx lobii (Hook.f.) Schimp.

Rosaceae

Prunus arborea (Blume) Kalkman

Prunus arborea (Blume) Kalkman var. stipulacea (Blume) Kalkman

Prunus lamponga (Miq.) Kalkman

Rubiaceae

Canthium didymum Ridl.

Diplospora singularis Korth.

Pleiocarpidia capituligera (Ridl.) Bremek.

Porterandia anisophylla (Jack ex Roxb.) Ridl.

Tarenna winkleri Valeton

Timonius eskerianus W.W.Sm.

168 Gard. Bull. Singapore 64(1) 2012

Rutaceae

Acronychia pendunculata (L.) Miq.

Maclurodendron porteri (Hook.f.) T.G.Hartley

Melicope incana T.G.Hartley

Salicaceae

Flacourtia rukam Zoll. & Mor.

Guioa diplopetala (Hassk.) Radlk.

Sapindaceae

Nephelium cuspidatum Blume var. ophioides (Radlk.) Leenh. subvar. beccarianum (Radlk.)

Leenh. Notes: Endemic to Borneo.

Nephelium lappaceum L. var. pallens (Hiern) Leenh

Nephelium macrophyllum Radlk. Notes: Endemic to Borneo.

Pometia pinnata J.R.Forst & G.Forst.

Xerospermum laevigatum Radlk. subsp. laevigatum

Sapotaceae

Madhuca barbata T.D.Penn. Notes: Endemic to Sarawak.

Madhuca engleri (Merr.) Vink Notes: Endemic to Borneo and uncommon in Sarawak (only

from Semenggoh).

Madhuca erythrophylla (King & Gamble) H.J.Lam Notes: Not endemic but uncommon in

Sarawak (Semenggoh & Segan only).

Madhuca kingiana (Brace ex King & Gamble) H.J.Lam

Madhuca korthalsii (Pierre ex Burck) H.J.Lam

Madhuca kuchingensis Yii & P.Chai Notes: Endemic to Sarawak and confined to the western

and central parts of Sarawak.

Madhuca lancifolia H.J.Lam Notes: Endemic to Borneo but uncommon; in Sarawak from

Semenggoh only.

Madhuca oblongifolia (Merr.) Merr. Notes: Not endemic but in Sarawak only from Semenggoh

& Bintulu.

Madhuca pallida (Burck) Baehni Notes: Not endemic but uncommon in Sarawak.

Madhuca proxila (Pierre ex Dubard) Yii & P.Chai

Madhuca sarawakensis (Pierre ex Dubard) H.J.Lam Notes: Endemic to Sarawak; Retsricted

to Kuching Division.

Madhuca sericea (Miq.) H.J.Lam

Madhuca sp. |

Palaquium beccarianum (Pierre) P-Royen Notes: Endemic to Borneo, uncommon in Sarawak

(Kuching Division only).

Palaquium calophyllum (Teijsm. & Binn.) Pierre

Palaquium cryptocariifolium P.Royen Notes: Endemic to Borneo; type specimen from

Semenggoh.

Palaquium decurrens H.J.Lam Notes: Endemic to Borneo.

Palaquium gutta (Hook.f.) Baill.

Palaquium herveyi King & Gamble Notes: Endemic to Borneo; in Sarawak from Semenggoh

only.

Palaquium leiocarpum Boerlage

Palaquium pseudorostratum H.J.Lam

Tree diversity at Semenggoh, Sarawak

169

Palaquium ridleyi King & Gamble

Palaquium rigidum Pierre ex Dubard Notes: Endemic to Sarawak and confined to Matang

Semenggoh & Baram only.

Palaquium rufolanigerum P.Royen Notes: Endemic to Borneo, type specimen from Semenggoh.

Palaquium walsurifolium Pierre ex Dubard

Payena obscura Burck subsp. havilandii (King & Gamble) J.T.Pereira Notes: Endemic to

Borneo: from first and second Division only.

Pouteria malaccensis (C.B.Clarke) Baehni Notes: Not endemic but uncommon in Sarawak

(Semenggoh & Kelabit Highland only).

Simaroubaceae

Quassia borneensis Noot.

Staphyleaceae

Turpinia sphaerocarpa Hassk. var. sphaerocarpa Hassk.

Turpinia sp. |

Stemonuraceae

Cantleya corniculata (Becc.) R.A.Howard

Stemonurus grandifolius Becc. Notes: Endemic to Sarawak.

Stemonurus secundiflorus Blume var. lanceolatus (Becc.) Sleum Notes: Endemic to Sarawak.

Symplocaceae

Symplocos goodeniacea Noot. Notes: Endemic to Borneo and uncommon in Sarawak.

(Semenggoh & Usun Apau only)

Symplocos henschelii (Moritzi) Benth. ex C.B.Clarke var. maingayi (Benth. ex C.B. Clarke)

Noot.

Symplocos rubiginosa Wall. ex DC.

Theaceae

Adinandra sarosanthera Mig.

Gordonia havilandii Burkill

Gordonia sarawakensis Keng

Ploiarium alternifolium (Vahl) Melchior

Thymelaeaceae

Aetoxylon sympetalum (Steenis & Domke) Airy Shaw Notes: Endemic to Borneo.

Gonystylus affinis Radlk. var. elegens Airy Shaw Notes: Endemic to Borneo, from Kuching

Lundu and Marudi District in Sarawak.

Gonystylus maingayi Hook.f.

Gonystylus micranthus Airy Shaw Notes: Type specimen from Semenggoh

Gonystylus stenosepalus Airy Shaw Notes: VU Alc+2c ver 2.3

Ulmaceae

Gironniera nervosa Planch.

Gironniera subaequalis Planch.

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Gardens’ Bulletin Singapore 64(1): 171-193. 2012 1

Podocarpus orarius (Podocarpaceae), a new species

from the Solomon Islands and a taxonomic clarification

of Podocarpus spathoides from Malaysia

R.R. Mill’ and M. Whiting

Royal Botanic Garden Edinburgh,

EH3 5LR, Scotland, U.K.

"R.Mill@rbge.ac-uk

ABSTRACT. Podocarpus spathoides de Laub. (Podocarpaceae) is revised and is restricted to

material from Malaysia where the type was collected. An emended description is given because

the protologue was based on a mixture of different taxa. Plants from the Solomon Islands,

previously described as Podocarpus spathoides var. solomonensis Silba, are here raised to

species rank as the new species Podocarpus orarius R.R.Mill & M.Whiting. This is currently

believed to be endemic to the Solomon Islands where it has been wild-collected on Choiseul, San

Jorge and Guadalcanal: cultivated material, apparently originating from the wild, has also been

seen from the island of New Georgia. Similar plants occur on neighbouring islands of Vanuatu

but require proper evaluation before they can be assigned to the new species. Illustrations

of the habit and reproductive characters of Podocarpus orarius are provided. Material from

Morotai in the Moluccas that has in the past been assigned to Podocarpus spathoides is also

morphologically distinct from the type but is insufficient for formal naming. The leaf cuticle

micromorphology of Podocarpus spathoides and P. orarius is described and illustrated.

Keywords. conservation assessments, cuticle micromorphology, Malaysia, new species,

Podocarpaceae. Podocarpus, Solomon Islands, taxonomy

Introduction

Podocarpus spathoides de Laub. (Podocarpaceae) was originally described by de

Laubenfels (1985) on the apparent basis of a single specimen, de Laubenfels 600,

from Gunung Ledang (Mt. Ophir), Peninsular Malaysia, that was cited as its holotype

(Fig. 1A). Three years later, in his treatment of Podocarpus for Flora Malesiana, de

Laubenfels (1988) expanded the published concept and distribution of Podocarpus

spathoides to encompass specimens from a range of scattered islets including Morotai

in the Moluccas, Rossel Island in the Louisiades Archipelago between eastern Papua

New Guinea and the Solomons, and the Solomon Islands themselves: a distribution that

has been repeatedly quoted by later authors, most recently by Farjon (2010a, 2010b).

However, examination of specimens received on loan from the Leiden Herbarium (L)

has revealed that de Laubenfels had been using the name *Podocarpus spathoides’ on

determinavit slips as early as 1979, on specimens from all the above areas. Therefore,

although the protologue (de Laubenfels 1985) appears to be based on a single collection

from Mt. Ophir, de Laubenfels’s description as given in it was actually based on all,

AZ Gard. Bull. Singapore 64(1) 2012

FLORA OF THE SOLOMON ISLANDS:

BOWAL ANDEN CCU

Fig. 1. Type specimens of Podocarpus spathoides de Laub. from Malaysia and of some

Podocarpus species occurring in the Solomon Islands. A. Holotype of P. spathoides de Laub.

(Malaysia: de Laubenfels P600, L). B. Holotype of P. orarius R.R.Mill & M.Whiting (Solomon

Islands, Pitisopa et al. 7, E). C. Isotype of P. salomoniensis Wasscher (Solomon Islands, Brass

2881, L). D. Holotype of P. insularis de Laub. (Sudest Island, Brass 27987, L).

Podocarpus orarius sp. nov. from Solomon Islands 173

or nearly all, the material he had seen prior to 1985. This becomes evident when one

compares it with his corresponding description in the later Flora Malesiana account

(de Laubenfels 1988). In both descriptions of P. spathoides tree height is given as 3-20

m and bud length as 2-6 mm.

However, in his Flora Malesiana account de Laubenfels (1988) added a note

that “In Malaya and Rossel I. the foliage buds are no more than 3 mm long, the others

are twice as long. In Malaya the trees grow in a summit scrub and are no more than 3-4

m high, elsewhere collectors report 12—20 m high trees” and acknowledged that “more

than one similar taxon may be involved here”. Female material was also described in

the protologue of Podocarpus spathoides (de Laubenfels 1985). Curiously, however,

the later Flora Malesiana account of that species (de Laubenfels 1988) did not include

any female reproductive characters in its description. The label of de Laubenfels P598

from Mt. Ophir said that the specimen was taken from a female tree but no cones are

present on the sheet or in the packet and no other female material from Mt. Ophir could

be located. The holotype of Podocarpus spathoides (de Laubenfels P600, L: Fig. 1A),

from the same locality, was said to be male but again no cones are present on that sheet

and none are visible on the images of the isotypes at RSA and US; the K isotype is

also sterile (A. Farjon, pers. comm. 26 Jan 2011). In the description below, therefore,

we state that both pollen cone and seed cone characters are not reliably known for

Podocarpus spathoides at the present time, although we have given a translation of the

brief description of female cones given by de Laubenfels (1985) in his protologue.

Some years later, Silba (2000) described Podocarpus spathoides var.

solomonensis Silba from easternmost Choiseul (Solomon Islands), distinguishing it

from typical Malayan P. spathoides by its larger bud scales and being a relatively

large tree up to 20 m tall, in comparison to the Malayan plants that are typically only

34 m tall. Silba’s description was extremely brief and all of it is summarised above.

The holotype of this variety (Whitmore BSIP 5247 at L) has been examined by us

and there is no evidence, such as a determinavit slip, that it was actually seen by

Silba. It was originally identified as Podocarpus neriifolius D.Don and later, by de

Laubenfels in 1967, as Podocarpus rumphii Blume and subsequently as P. spathoides

by de Laubenfels in 1979.

In October 2008 a joint expedition team from the Royal Botanic Garden

Edinburgh and the Ministry of Forests, Honiara, Solomon Islands collected a large

suite of specimens from most of the members of the Podocarpaceae known from the

Solomon Islands. This included specimens of Silba’s taxon Podocarpus spathoides

var. solomonensis which was found growing on the islands of Choiseul (including

at the locality where Whitmore collected the specimen that Silba made the type of

his variety) and San Jorge, off Santa Isabel, where a similar plant had previously

been collected by E.J.H. Corner (see specimen citations). It was noted that, in stark

contrast to the exposed mountain ridge-top habitat at altitudes above 1000 m above

sea level (asl) favoured by Podocarpus spathoides in its locus classicus of Mount

Ophir, the Solomon Islands plants located on this expedition were always found at

very low altitudes (from | to 35 m asl) along the coastline. Examination of their gross

morphology revealed many differences between these Solomon Islands plants and

174 Gard. Bull. Singapore 64(1) 2012

Malay P. spathoides and it became obvious that, as surmised by de Laubenfels (1988),

they were not the same species. This was later confirmed by examination of the cuticle

micromorphology of the two taxa by Whiting (2009). Accordingly, these Solomon

Islands specimens are described here as the new species Podocarpus orarius R.R.Mill

& M.Whiting.

De Laubenfels (1985) divided Podocarpus L’ Her. ex Pers. into two subgenera,

each of which had nine sections. The two subgenera have consistently been recovered

with strong support in molecular phylogenies of Podocarpaceae (Conran et al. 2000,

Sinclair et al. 2002, Biffin et al. 2011, Knopf et al. 2012). However, the present limits of

the sections are, with one or two exceptions, not supported by molecular phylogenetics

and they have not been recognised in the most recent world treatment of Podocarpus

(Farjon 2010a). Consequently, until a new, more meaningful sectional classification is

available, the new species is only assigned to subgenus.

Materials and methods

The taxonomic study reported here is based on the examination of twenty-four different

collections, eighteen of which belong to the new species Podocarpus orarius described

below. Four collections represent Podocarpus spathoides from the type locality and

one each represent taxonomically unassigned specimens from Morotai (Moluccas)

and Rossel I. (Louisiades Archipelago, New Guinea) as discussed below. Duplicates of

all specimens gathered by Pitisopa et al. in 2008 and cited below as being conserved

at E will be distributed to BSIP with all but one also duplicated at BISH. The intended

allocations are listed in the specimen citations.

The cuticle micromorphology of Podocarpus spathoides from the type

locality and of the new species P. orarius from the Solomon Islands was examined

using scanning electron microscopy. Specimens used are indicated by an asterisk (*) in

the specimen citations within the species accounts. The cuticle was isolated following

the method of Alvin and Boulter (1974) with changes made to suit Podocarpaceae

based on work by Kershaw (1997), Stockey et al. (1998) and Stark Schilling (2004).

From each specimen, six leaf slices of around 0.8 cm? were placed in glass vials with

6 ml of 20% aqueous chromium trioxide solution. The vials were sealed and kept at

room temperature for 96 hours. Isolated cuticles were removed from the solution,

washed with distilled water and left to dry on filter paper. Cuticles were checked under

the light microscope to check for homogeneity. Two cuticle slices were mounted on

an aluminium specimen stub using carbon adhesive discs (Agar Scientific Ltd.). The

external surface of the cuticle was observed on untreated leaf slices and leaf slices were

placed in chloroform to remove wax. These samples were mounted on to a specimen

stub as above. Specimens were sputter coated with 60% gold and 40% palladium for

two minutes in an Emscope SC500 sputter coater. Examination of all cuticles was

done using a Hitachi S-4700 II scanning electron microscope (SEM) at the University

of Edinburgh. Accelerating voltage was 5 kilovolts (kV), working distance varied

from 12.8 to 16.5 mm and it was found that optimal images were obtained by setting

Podocarpus orarius sp. nov. from Solomon Islands (75

LensMode to Analysis, instead of Normal. For the external surface of the cuticle,

images were taken of the abaxial and adaxial surface with and without wax at x250

magnification. Close up pictures were taken of the stomatal complex with and without

wax removed, a group of stomata and the guard cells. Images of the internal surface

abaxial and adaxial surfaces of the cuticle were taken at 250 magnification. All stubs

are deposited at the Royal Botanic Garden Edinburgh.

Conservation assessments have been applied to both species using the IUCN

guidelines (IUCN 2001, IUCN Standards and Petitions Working Group 2010).

Taxonomy

Podocarpus spathoides de Laub., Blumea 30 (1985) 267, descr. emend. hoc loco.

Subgen. Foliolatus de Laub. TYPE: Malaya, G. Ledang (Mt. Ophir), 3500 ft. [1067

m], 27 Jul 1978, small tree 4 m in mountain thicket, D.J. de Laubenfels P600 [‘‘male”

according to field label; holo L (no cones seen!); iso RSA, image seen (no cones

visible!), US, image seen (no cones visible!)]. (Fig. 1A)

Small tree up to 4 m tall. Characters of bark or branching not currently known. Twigs

of first and second years greenish brown, of third year not seen. Terminal buds globose

or depressed-globose, c. 3.5 x c. 3 mm, protected by at least 8 decussate scales in at

least three series and at most equalling bud diameter; outermost scales c. 2.5 « 0.5—0.3

mm, lanceolate or ovate, middle ones c. 3.5 = 0.8 mm, longer and slightly narrower

than the inner which are c. 2.7 x 1 mm; all scales lacking keels, outer ones purplish

tinged, inner ones brownish, outer ones acute with recurved or reflexed tips, inner

ones obtuse and erect, their margins entire, hyaline in distal half, the laminar part of

the scale smooth. Scale leaves absent in reproductive zones. Foliage leaves spirally

arranged, adult ones 3-8 mm apart and diverging from axis at 45—60°, juvenile ones

5—14 mm apart and diverging at 50—85°, all petiolate; petiole 5-8 mm, not distinctly

twisted, decurrent; lamina crimson or purplish and glaucous when flushing, turning

deep green (drying brownish green) and glossy above, paler and matt beneath with

dark midrib, narrowly elliptic, elliptic, narrowly oblong-elliptic or oblong-elliptic,

adult ones (30—)55—85 =x 9.5—13 mm, juvenile ones rather longer and wider, 65—100 x

14-20 mm, all straight (not falcate), thick and coriaceous, stiff, shallowly transversely

convex or flat adaxially; margin not thickened (or only slightly so, when young)

nor revolute, normally not undulating unless as a response to insect damage; midrib

relatively broad (1.3—2 mm wide), with striate band on either side beneath, raised both

above and beneath, on upper (adaxial) surface impressed in a broad channel equivalent

to the striate bands of the lower (abaxial) surface, on abaxial surface darker than rest of

lamina; apex obtuse or broadly rounded, not ending in a drip tip, sometimes blackening:

base cuneate or shortly attenuate. Pollen and seed cones not seen. “Female cones on

2—6 mm peduncles; basal foliola |.5 mm long; receptacle 5 mm long; seed 7 x 5 mm”

(translated by R.R.M. from the Latin protologue of de Laubenfels 1985: 267).

176 Gard. Bull. Singapore 64(1) 2012

Distribution. Podocarpus spathoides is here regarded as endemic to Peninsular

Malaysia and known only from the type locality. Records from the Solomon Islands

belong to the new species Podocarpus orarius R.R.Mill & M.Whiting described below.

Other records from east of Wallace’s Line (Rossel I., Morotai, Kepulauan Talaud)

are considered in the Discussion but excluded from the species. A specimen (Paie

32883, K) has also been collected under the name P. spathoides from Lawas, Sarawak

but is currently missing from the Kew herbarium so cannot be considered further at

present; according to Farjon (pers. comm., 26 Jan. 2011) it was from a tree 18 m tall

bearing yellow (presumably unripe) female cones. If it is correctly named, it would

truly extend the range of P. spathoides to Sarawak as stated by Farjon (2010a, 2010b).

Habitat and ecology. At the type locality P. spathoides occurs in low shrubby vegetation

and stunted forest on exposed ridges from c. 1000 m asl to the summit (1276 m asl),

above the altitudinal limit of Podocarpus ridleyi N.E.Gray (Farjon 2010b).

IUCN conservation assessment. In the present state of knowledge, Podocarpus

spathoides is best regarded as DD (Data Deficient), the category assigned to it by

Farjon (2010a, 2010b) albeit for different reasons. Farjon’s assessment was based on

the assumption that, given the very scattered distribution of the species as circumscribed

by him, it could perhaps be more widespread. The much more restricted distribution

accepted here paints a totally different picture about the possible threat to the species

and the species is more likely to fall within one of the threatened categories. However,

at present DD is still appropriate since more data concerning its current status at the

type locality, the level of threat there, and the identity of the putative collection from

Sarawak, are all required.

Local names. None recorded.

Other specimens seen (* denotes specimen utilised for examination of leaf cuticle).

MALAYSIA. G. Ledang (Mt. Ophir), 4000 ft [1219 m]., 27 Jul 1978, “sprouts 2 m”, D.J. de

Laubenfels P596 (L; juvenile); ibid., 4000 ft. [1219 m], 27 Jul 1978, “young plant 2 m”, D.J.

de Laubenfels P597 (L); ibid., 4000 ft. [1219 m], 27 Jul 1978, D.J. de Laubenfels P598 (L*;

“female” according to label but no cones present).

Note. If the description of the female cones of Podocarpus spathoides given in the

protologue by de Laubenfels (1985) can be relied upon (as noted above, no cones

from the type locality have been seen to verify their morphology), the Malay plant

differs from P. orarius in having considerably shorter peduncles and foliola, smaller

receptacles and shorter, narrower seeds. These differences, should they hold true, are

additional to those given below in the diagnosis of P. orarius.

Podocarpus orarius R.R.Mill & M.Whiting, sp. nov. Subgen. Folio/latus de Laub.

A Podocarpo spathoide de Laub. habitu multo altiore (usque ad 35 m haud 4 m tantum)

Podocarpus orarius sp. nov. from Solomon Islands vag

gemmis terminalibus in ramulis duplo longioribus anguste conicis non subglobosis

squamis diametro gemmae multo longioribus haud aequalibus interioribus caudatis

vel longe attenuatis nec exterioribus nec interioribus reflexis, foliis novellis pallide

viridibus roseo tincto (haud purpurascentibus) herbaceis demum subcoriaceis tantum

(non coriaceis) costa nec supra nec infra elevata inferne quam lamina distincte

pallidiore (haud atriore), ramulis cum foliis inferis persistentibus (haud caducis)

differt. A Podocarpo insulari de Laub. foliis multo longioribus latioribusque et squamis

gemmarum terminalium exterioribus haud valde reflexis insignis. A Podocarpo

salomoniensi Wasscher foliis multo latioribus (12-19 mm non 6.5—8 mm) oblongo-

ellipticis vel oblongis (haud anguste lineari-lanceolatis) ad marginem planis (haud

valde revolutis) costa superne indistincto (haud prominenti) et non sulcato, receptaculo

fructifero e bracteis duabus vel tribus (haud quattuor decussatis) composito facile

distinguitur. Podocarpo rubenti de Laub. foliis novellis rubentibus similis a qua

tandem foliis multo majoribus et receptaculo maturo rubro haud purpureo distincta.

Podocarpo neriifolio D.Don foliis novellis rubentibus et foliis longis etiam similis a quo

perulis gemmarum primariis longioribus 4-8 mm longis, amentis masculis pedicellatis

haud sessilibus, receptaculo majore et seminibus subglobosis differt. TYPE: Solomon

Islands, Choiseul Province, Choiseul, Loloko District, mainland opposite Bembalama

Island, 07° 21’ 11.4"S 157° 33’ 39.6"E, 20 m, 4 Oct 2008, F: Pitisopa, M.F- Gardner, S.

Herrington, P. Kosui, R. Olisae & P. Tofu 7 (ripe female: holotype, E; isotypes, to be

distributed to BISH and BSIP). (Fig. 1B, 2 & 3)

Synonyms: Podocarpus spathoides de Laub. var. solomonensis Silba, J. Int. Conifer

Preserv. Soc. 7:1 (2000) 39. — P. spathoides de Laub. subsp. solomonensis (Silba)

Silba, J. Int. Conifer Preserv. Soc. 17:1 (2010) 19. TYPE: Solomon Islands, 7°C.

Whitmore BSIP 5247 (holo L, iso K).

Erect or sometimes leaning single- or less commonly multi-stemmed dioecious tree to

35 m tall, up to 20 cm dbh. Crown rather narrow and ellipsoid (Fig. 2A). Bark (Fig.

2B) smooth, flaking, shallowly fissured; outer bark medium brown, inner bark pinkish

brown, wood straw-coloured. Twigs of first year pale grey green and herbaceous,

of second year pale green or brownish green; growing tip purplish. Terminal buds

narrowly conical, 4-8 x c. 1 mm, protected by about 8 decussate, narrowly lanceolate

scales in two series and much longer than bud diameter; outer scales 4-8 = 0.2—0.3

mm, longer and slightly broader than the inner which are 3—5S.5 = 0.1—0.25 mm; all

scales lacking keels, purplish at base (purplish tinge fading with age), green towards

tip, long-caudate or long-attenuate, obtuse at extreme tip, erect or the outer ones only

slightly recurved, their margins hyaline in distal half, the laminar part of the scale

rugose. Scale leaves present in reproductive zones with an abrupt transition to foliage

leaves. Foliage leaves present on penultimate and lower axes as well as ultimate ones,

those on the penultimate and lower axes long-persistent on young trees but caducous

on older ones; juvenile and adult foliage leaves similar in shape and size; leaves

spirally arranged, those of adult trees 6-25 mm apart and diverging at 60—85°, of

juvenile trees 3-15 mm apart and diverging at 60—85°, all leaves petiolate; petiole 4—6

178 Gard. Bull. Singapore 64(1) 2012

Fig. 2. Podocarpus orarius R.R.Mill & M.Whiting: vegetative features. A. Habit (Pitisopa et

al. 78, in field). B. Bark (Pitisopa et al. 6, in field). C. Mature foliage, Choiseul (Pitisopa et al.

6, in field). D. Flushing leaves, Choiseul (Pitisopa et al. 6, in field). E. Flushing leaves, Santa

Isabel (Pitisopa et al. 78, in field). (Photos: M.F. Gardner).

1 80 Gard. Bull. Singapore 64(1) 2012

mm, twisted, not or scarcely decurrent; lamina light green tinged pink on both surfaces

when flushing (Fig. 2C & 2D) but not glaucous, turning deep glossy green above but

much paler beneath (Fig. 2E), younger leaves held suberect but older ones becoming

horizontally spreading, (sometimes narrowly) elliptic, oblong-elliptic or oblong, (S0—

)70-195 x (10.5—)12—20 mm with little difference in size between juvenile and adult

leaves, straight, falcate throughout or not, thin, flexible and herbaceous at first becoming

thicker, stiff and subcoriaceous with age, shallowly concave adaxially; margin scarcely

thickened, very slightly undulating or not, not revolute; midrib narrow (0.2—0.3 mm

wide), with indistinct striate band on either side beneath, not or only slightly raised

on both surfaces, not impressed on upper (adaxial) surface, on lower (abaxial) surface

markedly paler than rest of lamina; apex acute or subacute, not ending in a drip-tip,

sometimes blackening; base cuneate or shortly attenuate. Pollen cones shedding pollen

at same time as leaf flushing, shedding from base to tip, lateral, subtended by a scale

leaf and arising on current growth just above bud scales of previous season, solitary or

paired, up to 12 together but individual groups composed of not more than 3 (Fig. 3A);

common peduncle absent; individual pollen cones (Fig. 3B) pedicellate, the pedicel

3-9 mm, shorter than cone, straight, erecto-patent, with spreading pedicel scales and

surrounded by basal scales; basal scales c. 4, light brown, suberect and encircling base

of pedicel, not keeled, ovate-lanceolate and rather narrow (outer ones c. 3 x 1.5 mm,

inner ones c. | x | mm), acute, muticous, with very narrow entire scarious hyaline

margins; scales on pedicels decurrent at base, 5 or usually 6, spirally arranged, c.

1.3—-1.7 x 1 mm, pinkish-brown, subadpressed to erecto-patent, not keeled, oblong-

ovate, subacute, muticous, with crenulate scarious hyaline margins; pollen cones erect,

suberect or spreading outwards, 20-36 x 2—3.5 mm, cream or lemon-yellow, lighter

than and contrasting with foliage, tapering slightly distally, narrowly cylindrical,

curved from base or in distal half; microsporophyll phyllotaxis 13/34 with c. 32 short

spirals of 3-4 microsporophylls and c. 26 long spirals of 5—6 microsporophylls per

cone; microsporophylls c. 250 per cone, c. | mm; microsporophyll lamina greenish

at base with pinkish tip, the free part at apex broadly triangular or deltate, c. 0.25

x 0.2 mm, not up-turned, with crenulate, somewhat hyaline and fimbriate margin,

not scarious, truncate at apex; microsporangia slightly oblique and divergent, free,

cream, ellipsoid, c. 0.7 x 0.5 mm, the stomium elliptic when open with hood at top, the

microsporangial walls scarious-margined; pollen milky-white or hyaline. Seed cones

borne on current growth, paired, lateral on a specialized reproductive shoot subtended

by a leaf or bract; receptive cones (Fig. 3C & 3D) borne among leaves but presented

on long peduncles (up to 16 mm, longer than both the receptive cone complex and the

receptacle); subtending scale 1, c. | x 0.7 mm, light brown, not keeled, + rectangular

with a narrower, subacute muticous tip and very narrow scarious margin, falling

before cone is ripe; peduncles horizontal both when receptive and when ripe, ridged,

compressed, broadening distally; ripe cones (Fig. 3E & 3F) horizontal, the peduncle

on cones seen 10-11 mm and now shorter than both the whole cone complex and

the receptacle; prophylls (foliola) light green tipped purplish, narrowly lanceolate, c.

2.5 x 0.2 mm, erecto-patent at receptivity, soon caducous, if present when ripe then

deflexed, straight throughout their life with acute, + cucullate apex; receptacle fleshy

Podocarpus orarius sp. nov. from Solomon Islands 181

when ripe, composed of | fertile and | or 2 sterile bracts, asymmetrical, obovoid and

infundibular at receptivity, rectangular-ellipsoid at maturity and then 10—13.5 mm

along longest edge, 8.5—10.5 mm along shortest edge, 5.5—7.5 mm wide at top, greenish

at receptivity (Fig. 3C & 3D), turning yellow and finally deep vermilion at maturity

(Fig. 3E & 3F), not glaucous at any stage; fertile bract longer than both ovule and seed,

erect at receptivity, with median longitudinal groove; receptacular scales all connate

and + wholly fused, with short, broad, lip-like free tips that are erect in vivo, slightly

bent outwards and 2—3 x 4-5 mm (fertile bract) and 1—2.5 x 2—3 mm (sterile bract)

in sicco, swollen and fleshy at maturity; seed asymmetrically placed on receptacle,

subglobose, 11—12 « 9-9.5 mm, laterally compressed, not crested at topographically

distal (chalazal) end, without a notch or beak at micropylar end, the micropyle arch-

like at receptivity, pinkish brown, 2-pronged; seed coat and epimatium olive green

without purplish tinge even when fully ripe, glaucous at receptivity, wrinkled and

rugose when dry; cotyledons unknown.

Phenology. Male and young female cones present in early October (and possibly

earlier); new leaves flushing at same time. Ripe female cones (of previous year)

also present in early October together with slightly unripe ones; therefore time from

pollination to ripening is c. 12 months.

Distribution. At present only known with certainty from the Solomon Islands

archipelago and therefore currently regarded as endemic there. However, images of

similar material have been seen from Vanuatu (islands of Erromango and Aneityum)

that are superficially similar to Podocarpus orarius. These sheets have been determined

as Podocarpus insularis by de Laubenfels (in sched.) but are not that species, the type

of which has much smaller leaves as noted above; nor are they P. neriifolius which

was the only Podocarpus species recorded from those islands by Schmid (1975). They

have the large, broad leaves of Podocarpus orarius but examination of the actual

specimens will be necessary to determine whether they are conspecific with it or not.

If they are conspecific, they would extend the geographic range of Podocarpus orarius

to the neighbouring Vanuatu archipelago.

Habitat and ecology. Podocarpus orarius occurs on steep slopes of primary coastal

rainforest, often with species of Gymnostoma L.A.S.Johnson (Casuarinaceae); also in

secondary rainforest; 1—60(—460) m asl. Recent collections on the Solomons have all

been by the coast below 40 m asl but, as indicated in the specimen citations below, it

has in the past been collected on Guadalcanal up to an altitude of just over 450 m asl.

The species should therefore also be searched for at higher altitudes and in the interior

of the islands; on Guadalcanal and New Georgia at least, there is evidence (from the

information on the labels of Lipageto BSIP 3321 and Waterhouse 209) that trees of the

species do (or did) occur away from the coast.

IUCN conservation assessment. NT (Near Threatened). There is no substantial evidence

of decline although in some coastal areas there has been some forest destruction with a

182 Gard. Bull. Singapore 64(1) 2012

few mutilated trees of P. orarius. Local information suggests that it is likely that these

largely intact forests will be targeted for their timber in the near future, therefore this

species will be vulnerable to any change in land use. The nearby nickel mines on the

island of San Jorge are also a potential future threat. Previously listed as DD, under

the name Podocarpus spathoides, by Pippard (2008, App. 2 p. 5) based on a 1998

assessment.

Notes. The type of Podocarpus orarius was collected at the same site as the type

locality of P. spathoides var. solomonensis. We have chosen to describe a new species

rather than raise Silba’s varietal epithet to species rank for two reasons: the similarity

of ‘solomonensis’ to ‘salomoniensis’ which is already in use for a completely different

species of Podocarpus, and the brief and unsatisfactory nature of Silba’s protologue

which contained only the bare minimum of information required to validate the name.

The type of Silba’s name was also a sterile specimen, whereas both male and female

specimens (the females at two different stages) as well as more juvenile specimens

were collected in 2008, allowing the preparation of a much more complete description

from these and the other available material.

Local names. Locally known on Choiseul and Guadalcanal as Dengali in the Kwara’ae

language (Whitmore 5247, Lipageto BSIP 3321) and on Santa Isabel as Graoragota

(female) in the Maringe dialect of the Cheke Holo language (Pitisopa et al. 72, 73,

90, 91, 92, 95, 103, 104, 105). Graoragota 1s also used for Podocarpus salomoniensis

according to the notes on Pitisopa et al. 96 (E).

Etymology. The epithet orarius is Latin for ‘coastal’ and alludes to what appears to be,

from the majority of collections seen, the typical habitat of the species.

Other specimens examined (* denotes specimen utilised for examination of leaf cuticle):

SOLOMON ISLANDS. Choiseul Province: Choiseul, ultrabasic hill on coast opposite

Bembalama Island, 3 Mar 1964, 7.C. Whitmore BSIP 5247 (L, sterile, holotype of P. spathoides

var. solomonensis Silba; K, isotype of P. spathoides var. solomonensis); Loloko District,

mainland opposite Bembalama Island, 07° 21’ 11.4”S 157° 33’ 39.6"E, 20 m, 4 Oct 2008, F

Pitisopa, M.F: Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 5 (E, BSIP, BISH; male);

ibid., F! Pitisopa, M.F: Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 6 (E, BSIP, BISH;

immature female). Isabel Province: Santa Isabel, Turungurungu Island, San Jorge, sea level,

25 Sep 1965, E.J.H. Corner 2737 (K, L); Bughotu District, San Jorge, Turunghu Island, east

of the village of Talise, 08° 27’ 57.2"S 159° 38’ 21.7"E, 35 m, 16 Oct 2008, F Pitisopa, M.F-

Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 72 (E, BSIP, BISH); ibid., F) Pitisopa,

M.F: Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 73 (E, BSIP, BISH); Bughotu

District, San Jorge, Tupilla Island, 08° 27’ 0.68”S 159° 38’ 28.8” E, 1 m, 16 Oct 2008, F

Pitisopa, M.F: Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 78 (E*, BSIP; immature

female: cuticle of both young flushing leaf and adult leaf examined); Bughotu District, San

Jorge, Ramahale Point, 08° 29’ 34.8”S 159° 38’ 59.8”E, 15 m, 17 Oct 2008, F: Pitisopa, MF:

Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 83 (E*, BSIP, BISH; sterile: cuticle of

adult leaf examined); Bughotu District, San Jorge, Gobu Bay, 08° 29’ 37.7"S 159° 38’ 47.5"E,

Podocarpus orarius sp. noy. from Solomon Islands 183

5 m, 17 Oct 2008, F Pitisopa, M.F: Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 90

(E, BSIP, BISH); Bughotu District, San Jorge, Gobu Bay, 08° 29’ 37.7”S 159° 38’ 47.6"E, 6

m, 17 Oct 2008, F Pitisopa, M.F- Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 91 (E,

BSIP, BISH; young female); Bughotu District, San Jorge, Gobu Bay, 08° 29’ 37.5"S 159° 38’

47.7"E, 6 m, 17 Oct 2008, F Pitisopa, MF: Gardner, S. Herrington, P. Kosui, R. Olisae & P.

Tofu 92 (E, BSIP, BISH; immature female); Bughotu District, San Jorge, Simia River c. 0.5

km from mouth of river, 08° 32’ 31.7”S 159° 38’ 45.4"E, 12 m, 18 Oct 2008, F. Pitisopa, MF.

Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 102 (E, BSIP, BISH); Bughotu District,

San Jorge, Kogaruta Bay, 08° 30’ 14.4” S 159° 41’ 1”E, 1 m, 18 Oct 2008, F Pitisopa, MF:

Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 103 (E, BSIP, BISH); Bughotu District,

San Jorge, Tanegula, 08° 29’ 51.1” S 159° 41' 4.8"E, 1 m, 18 Oct 2008, F Pitisopa, MF:

Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 104 (E, BSIP, BISH); Bughotu District,

San Jorge, Turunghu Island, east of the village of Talise, 08° 27'57”S 159° 38’ 1.7"E, 1 m, 18

Oct 2008, F Pitisopa, M.F- Gardner, S. Herrington, P. Kosui, R. Olisae & P. Tofu 105 (E, BSIP,

BISH). Guadalcanal Province: Guadalcanal, Eastern slopes of Mt. Gallego, 1500 ft. [457

m], 08 Aug 1965, 7. Whitmore 2079 (K). Rere River, c. 3 miles inland, 200 ft. [61 m], 19 Nov

1963, Z. Lipaqeto BSIP 3321 (K). Western Province: New Georgia, grown in garden on coast,

originating from interior (Kusage), 26 Jul 1929, J.H.L. Waterhouse 209 (K, 2 sheets: cited by

Gray, 1958 under P. neriifolius).

Cuticle micromorphology

Podocarpus spathoides (Fig. 4A—H). External surface. Adaxial epidermal cells (Fig.

4A) indistinctly visible, abaxial ones (Fig. 4B) clearly visible on external surface

of cuticle. Stomatal plugs present, forming a popcorn-like amorphous, very porous

interconnected network (Fig. 4C inset). Stomata without Florin rings but surface

upraised (Fig. 4C). Internal surface: epidermal cells. Adaxial epidermis cells (Fig.

4D & 4E) polygonal, square to elliptic or rectangular (either longitudinally or less

commonly transversely); walls slightly undulating, no cavities along wall flanges;

periclinal surfaces fibrous-granular with numerous small holes. Abaxial epidermal

cells (Fig. 4F & 4G) narrowly rectangular to rectangular or oblong-polygonal; those

adjoining stomata trapezoidal or arcuate, those within stomata rows transversely

elliptic / rectangular or polygonal; walls straight or curved, not buttressed though

edges irregularly frilled, cavities along wall flanges indistinct or absent; periclinal

surfaces rather coarsely granular. Internal surface: stomatal arrangement. Stomata

in discontinuous rows on abaxial surface (hypostomatic), parallel to long axis of

leaf with + no deflection of individual stomata; stomatal chains developed (Fig. 4F).

Stomata (Fig. 4G & 4H) brachyparacytic, close together within same row, separated

by | or 2 epidermal cells; adjacent rows widely spaced, separated by (1—)2—3(—5) rows

of epidermal cells; horizontal stomata rows absent. Stomatal apparatus (including

subsidiary cells) suborbicular (Fig. 4G & 4H). Stoma (excluding subsidiary cells)

suborbicular to + square, the opening elliptic and often wide (Fig. 4G & 4H). Subsidiary

cells: 2 most common, 3 frequent, 4 rare; polar subsidiary cells absent; cuticle on outer

flanges thick, with no groove but cuticle with a deep crease that is hidden by lateral

184 Gard. Bull. Singapore 64(1) 2012

Tee eee ea

BIOSEM SOKV 15.7mmx250 SEL) 07/02/09 16:33

Moa &

Fig. 4. Podocarpus spathoides de Laub.: cuticle micromorphology (all from de Laubenfels

P598, L). A. Adaxial external surface. B. Abaxial external surface; composite image showing

(left half) untreated cuticle with wax plugs covering stomata, (right half) cuticle treated with

chloroform, stomata with wax plugs removed. C. external view of stoma; (inset) detail of wax

plug before treatment. D. Adaxial epidermal surface. E. Adaxial epidermal cells. F. Abaxial

epidermis with stomatal rows and epidermal cells. G. Group of stomata. H. Stomatal complex.

Scale bars: A, B, D, F, 200 micrometres (um); C, 20 um; E, 40 um; G, 50 um; H, 30 pm.

(Photos: M. Whiting).

Podocarpus orarius sp. nov. from Solomon Islands 185

flange extensions (Fig. 4G, top left); margins of subsidiary cell wall flanges irregularly

denticulate, the teeth incurving; thinning of cuticle present in an arc midway across the

subsidiary cell; texture fibrous-granular. Cuticle flange between guard cells smooth

towards outside of flange, slightly more rugose towards stoma, apparently not inrolled,

not standing out laterally (Fig. 4H). Polar extensions always present, ribbon-like, thin

and broad, wholly fused with subsidiary cells, longer than broad, with a central ridge,

flaring outwards from the base (Fig. 4H). Cuticle on guard cell surfaces rugose.

Podocarpus orarius (Fig. 5A-H & 6A-F). External surface. Abaxial and adaxial

epidermal cells clearly visible on external surface of cuticle (Fig. SA, 5B, 6A & 6B).

Stomatal plugs present (Fig. 5A & 6A), composed of dendroid or coralloid fibrous

rodiets (Fig. SC inset & 6C), porous. Stomata without Florin rings but surface

somewhat upraised (Fig. 5C), especially when young (Fig. 6C & 6D). Developing

lateral subsidiary cells forming ridges either side of stomata, these ridges continuing

along the stomatal rows on young leaves, connecting one stoma with the next (Fig.

6A & 6B), this feature becoming much less evident on mature leaves (Fig. 5B, right).

Internal surface: epidermal cells. Adaxial epidermis cells (Fig. 5D & 5E) rectangular,

trapezoidal, square or transversely rectangular; walls straight or curved, not buttressed

though edges irregularly frilled, cavities along wall flanges present, more distinct

than in P. spathoides; periclinal surfaces granular with numerous small, rectangular

pits. Abaxial epidermal cells (Fig. SF & SG) narrowly rectangular to rectangular

or oblong-polygonal, those adjacent to stomata arcuate-rectangular or trapezoidal,

those within stomata rows transversely elliptic or suborbicular-polygonal; walls

straight, curved, or slightly undulating, not buttressed though edges irregularly frilled,

distinct small cavities present along wall flanges; periclinal surfaces granular with

scattered small + rectangular pits. Internal surface: stomatal arrangement. Stomata

in discontinuous rows on abaxial surface (hypostomatic), parallel to long axis of leaf

with some noticeably deflected; short stomatal chains sometimes developed. Stomata

(Fig. SG & 5H) brachyparacytic, close together within same row and separated by

(1—)2(—3) epidermal cells; adjacent rows widely spaced and separated by (2—)3—6 rows

of epidermal cells; horizontal stomata rows absent. Stomatal apparatus (including

subsidiary cells) transversely oblong or elliptic. Stoma (excluding subsidiary cells)

rectangular, the opening elliptic (Fig. 5H). Subsidiary cells: 2 most common, 3 or

4 occur; polar subsidiary cells absent; cuticle on outer flanges thick, with no visible

groove or crease (Fig. 5G & 5H); margins of subsidiary cell wall flanges irregularly

denticulate, the teeth incurving; thinning of cuticle present in an arc midway across

the subsidiary cell (Fig. 5H); texture fibrous-granular. Cuticle flange between guard

cells thick, smooth towards edge, finely rugose towards stoma, slightly inrolled or not,

not standing out laterally. Polar extensions always present, ribbon-like, wholly fused

with subsidiary cells, longer than broad, with a rather indistinct central ridge, flaring

outwards only distally (Fig. SH). Cuticle on guard cell surfaces rugose.

The cuticle micromorphology of Podocarpus spathoides (Fig. 4) and P. orarius

(Fig. 5 & 6) at first sight appears very similar and indeed there are many characters

186 Gard. Bull. Singapore 64(1) 2012

eas

"€

a :

BioSEM 5.02M.16.1mm x1 .00k SE(L

Fig. 5. Podocarpus orarius R.R.Mill & M.Whiting: cuticle micromorphology (adult leaves,

fully developed, Pitisopa et al. 78 & 83). A. Adaxial external surface. B. Abaxial external

surface; composite image showing (left half) untreated cuticle with wax plugs covering stomata,

(right half) cuticle treated with chloroform, stomata with wax plugs removed. C. External view

of stoma; (inset) detail of wax plug before treatment. D. Adaxial epidermal surface. E. Adaxial

epidermal cells. F. Abaxial epidermis with stomatal rows and epidermal cells. G. Group of

stomata. H. Stomatal complex. Scale bars: A, B, D, F, 200 tm; C, 40 um; E, G, 50 um; H, 30

um. (Photos: M. Whiting).

Podocarpus orarius sp. nov. from Solomon Islands 187

J y

ei fee BE

BIoSEM 16.0kV'13,3mim «250 SIL) 2}

Fig. 6. Podocarpus orarius R.R.Mill & M.Whiting: cuticle micromorphology (very young,

flushing leaf, Pitisopa et al. 78). A. external surface, abaxial cuticle, before treatment. B.

external surface, abaxial cuticle, after treatment with chloroform to remove wax. C. Stoma

before treatment, showing wax plug apparently sitting on guard cells within peristomatal

chamber. D. Stoma after treatment with chloroform to remove wax; guard cells closed. E.

Another stoma with wax removed showing peristomatal chamber. F. External surface, adaxial

cuticle. Scale bars: A, B, D, F, 200 um; C, 20 um; D, 10 um; E, 30 um. (Photos: M. Whiting).

188 Gard. Bull. Singapore 64(1) 2012

in common. However, closer examination of the images reveals some significant

differences that, when combined with the gross morphological differences and the

major disjunctions in both distribution and altitude, support separation of P. orarius.

On the external adaxial cuticle, no obvious sculpturing is visible in P. spathoides (Fig.

4A) but P. orarius has a pattern of ridges and hollows that corresponds to the shapes

of the adaxial cells as revealed from the inner cuticle surface (Fig. SA, 5D & 6F).

This pattern is particularly evident on young leaves (Fig. 6F). The sculpturing pattern

of the abaxial external cuticle of P. orarius is also very prominent, especially after

treatment with chloroform to remove wax (Fig. 5B, right); P. spathoides also has some

sculpturing on this surface but it is different and less distinct (Fig. 4B). The internal

cuticle also reveals differences between the Malaysian (P. spathoides) and Solomon

Islands (P. orarius) specimens. In P. spathoides the guard cell ‘wings’ are fairly large

and often cover a fairly deep and porous groove on the lateral subsidiary cells, whereas

in P. orarius the guard cell ‘wings’ are usually absent or relatively small and the lateral

subsidiary cells of the stomatal complex have only a slight groove or porous area. Small

cavities (more distinct on the abaxial surface than the adaxial) are present alongside the

epidermal cell walls on both leaf surfaces in P. orarius (Fig. SE & 5G) but these were

not observed in P. spathoides (Fig. 4E & 4G). The periclinal surfaces of the adaxial

epidermal cells have a more coarsely granular texture in Podocarpus spathoides (Fig.

4E) than in P. orarius (Fig. 5E). Finally, the margins of the lateral subsidiary cells are

noticeably more irregular and ‘jagged’ in appearance in P. spathoides and are clearly

outspread (Fig. 4H) compared with P. orarius in which they are almost smooth and

even tend to curl inwards (Fig. 5H).

Examination of the cuticle of an immature leaf of Podocarpus orarius shows

that in this species the wax plug/layer appears to sit on the guard cells within the

peristomatal chamber early in development (Fig. 6C & 6D). The lateral subsidiary

cells at this point form a narrow ridge either side of the guard cells (Fig. 6C & 6D).

These ridges continue beyond the stomata to link all the stomata in the row but the

feature gradually becomes less evident as the leaf becomes fully developed (Fig. 5B,

right). In other Podocarpus species Whiting (2009) found that the wax plug sits high

in the peristomatal chamber with a relatively large gap between the wax plug and the

guard cells. It was also found that the base of the wax plug often reflected the shape

of the guard cells somewhat (Whiting 2009). The shape of the wax plug therefore

indicates that it was once seated on the guard cells (as seen in Fig. 6C—E) although

the latter have changed in shape or position. It 1s possible that as the subsidiary cells

develop and expand, they push or rotate the guard cells downwards. The layer of wax

occurring on the guard cells may be displaced, leaving a wax layer fixed between the

subsidiary cells and with a gap between the wax plug and the guard cells. Another

explanation would be that the wax plug has been lifted and compressed by the lateral

subsidiary cells causing an overall change in shape of the wax plug. There may be an

upward movement of the wax plug as the subsidiary cells expand upwards, particularly

in Podocarpus subgenus Podocarpus in which there is a Florin ring. This topic will be

discussed in more detail elsewhere (Whiting 2009, Whiting & Mill in prep.).

Podocarpus orarius sp. nov. from Solomon Islands 189

Discussion

There are three other Podocarpus species that have been definitely recorded from the

Solomon Islands: Podocarpus glaucus Foxw.. P. insularis de Laub. and P. salomoniensis

Wasscher (Wasscher 1941, de Laubenfels 1988). Podocarpus neriifolius D.Don has

also been recorded (Gray 1958, two specimens cited; Whitmore 1966, Farjon 2010a)

but many specimens from the Solomons purporting to be that species have been

misidentified and we have not so far seen correctly identified material from there.

Waterhouse 209 at K, cited by Gray (1958) under P. neriifolius, bears an annotation

slip by Buchholz saying, “Under study — probably juv. form of P salomoniensis

Wasscher’, and a much later determinavit slip by A. Farjon who identified it as P

insularis. It is none of those three species and in fact belongs to P. orarius of which it

represents one of the few inland collections seen.

Podocarpus glaucus is a small-leaved (8—18 mm long: Farjon 2010a) species

from relatively high altitudes (normally above 1000 m asl) that cannot be confused

with P. insularis, P. salomoniensis or P. orarius. Previous records of P. salomoniensis

have been from San Cristoval (the type locality and others on that island) and San

Jorge (Corner 2717, K). It has also been recorded from Bougainville (Foreman 1971);

this island is politically part of New Guinea although phytogeographically it forms,

together with the Solomon Islands (other than the Santa Cruz group), a well-defined unit

that has been called the Solomon Islands rain forests ecoregion (ecoregion AAO119:

WWF 2001). Until the 2008 expedition, PR salomoniensis had not been recorded

from Choiseul but on that expedition it was collected there and also re-collected on

San Jorge (M.F. Gardner, pers. comm. 15 Nov 2010). During the 2008 expedition,

specimens of P. salomoniensis were found at very low altitudes (as low as c. 10 m asl)

whereas previously the known altitudinal range had been quoted as being 400-900 m

asl (Silba 1986). Podocarpus salomoniensis has very narrow leaves (Fig. 1C) and is a

very distinct, unmistakable species that cannot be confused with any other Podocarpus

on the Solomons archipelago or indeed anywhere in Malesia.

Podocarpus insularis de Laub. was based on Brass 27987 (Fig. 1D) from Mt.

Riu on Sudest Island; this is the largest island of the Louisiades Archipelago (New

Guinea, Milne Bay District) and is now known as Vanatinai. De Laubenfels (1988)

gave its distribution as “New Hebrides [ Vanuatu] and all Solomon Islands; in Malesia,

New Guinea and adjacent islands: Rossel, Sudest, Misima, Woodlark, Fergusson, and

New Britain.” The leaves of the holotype (Fig. 1D) and isotype of P. insu/aris measure

only 45—70 x 4—7 mm, much shorter and narrower than those of P. orarius (Fig. 1B) and

smaller also than those of Malaysian P. spathoides (Fig. 1A). Farjon in 2007 determined

Whitmore BSIP 5247 at K as Podocarpus insularis. Since that specimen is the type

of Podocarpus spathoides var. solomonensis, that name (and the later combination P.

spathoides subsp. solomonensis) would become synonymous with P. insularis were

Farjon’s identification proved to be correct. However, neither the Leiden nor the Kew

examples of Whitmore BSIP 5247 match the type of Podocarpus insularis—they have

much larger leaves and belong to P. orarius described above—and consequently we

disagree with Farjon’s determination. It is possible that the statement by de Laubenfels

190 Gard. Bull. Singapore 64(1) 2012

(1988) that P. insularis occurs on “all Solomon Islands” is at least partly wrong and

that some material of the new species here described as P. orarius was included in his

original concept of P. insularis as well as his circumscription of P. spathoides. Until

P. insularis is revised (a topic beyond the scope of this paper), its account in Flora

Malesiana, like that of P. spathoides, needs to be treated with caution.

Podocarpus neriifolius has large leaves that could perhaps be confused with

those of P. orarius. However, even in its widest senses as interpreted by Wasscher

(1941) and Farjon (2010a), it can be distinguished from P. orarius by its shorter,

spreading outer bud scales normally not exceeding 5 mm, its shorter receptacles of the

female cones (8-10 mm, as opposed to 10—13.5 mm in P. orarius), and its purplish-

tinged ripe seeds that are more oblong or ovoid than the subglobose seeds of P. orarius,

being (8—)10—15 x 7—8 mm including the epimatium rather than 11—12 x 9—9.5 mm as

in P. orarius.

Podocarpus spathoides has also been recorded from the island of Morotai

in the north Moluccas, off the north tip of Halmahera (de Laubenfels 1988). The

specimen upon which this record was based has been examined and again was found

to differ from the Malayan type but although it bears immature male cones it 1s a rather

poor specimen that is here regarded as insufficient as the basis for a formal description.

No other material identified as P spathoides has yet been seen from that island.

Therefore, formal segregation of the Morotai plant from P. spathoides must await the

examination of further material, either by discovery of more material in herbaria or

by collecting. Similarly, another record of P. spathoides from the Moluccas, from the

islet of Kepulauan Talaud (Silba 1986) also needs re-evaluation but the material upon

which it was based is not known. It is possible that it might have been an error for the

Morotai record although Kepulauan Talaud is in a different region of the Moluccas,

NE of Sulawesi close to the Philippine Trench.

De Laubenfels (1988) also recorded P. spathoides from Rossel Island in the

Louisiades Archipelago (New Guinea). These islands lie between New Guinea and

the Solomons; Rossel (now known as Yela) 1s the easternmost of the archipelago. One

specimen from Rossel Island has been seen; it was determined in 1979 by de Laubenfels

as Podocarpus spathoides and is presumably the basis for his record of that species

from Rossel Island in Flora Malesiana (de Laubenfels 1988). Earlier determinations

on this sheet were as Podocarpus rumphii (on the sheet’s label by an unknown person,

possibly the collector) and P. polystachyus R.Br. ex Endl. (by de Laubenfels in 1968).

De Laubenfels (1988) himself noted that the terminal buds of the Rossel I. specimen

were half the size of those from the Solomons, being instead similar in size to those

of the type of P. spathoides from Mt. Ophir. Also, the leaves are concave adaxially as

in Podocarpus polystachyus rather than flat as in P. orarius. The habitat, “inner edge

of mangroves’, is one favoured by Podocarpus polystachyus in other areas (Wasscher

1941, Farjon 2010b); it also likes rocky shores (Turner et al. 2000) and limestone

karst where it can be very dwarfed (Donnelly et al. 2003). All these are different from

the habitats favoured by both Podocarpus spathoides and P. orarius. De Laubenfels

(1988) restricted P. polystachyus to a range extending only from the Malay Peninsula

through islets off Sumatra to Borneo (especially Sabah, Brunei and Sarawak but also a

Podocarpus orarius sp. nov. from Solomon Islands 19]

few localities in W Kalimantan), the Philippines and western New Guinea (especially

the Vogelkop Peninsula): Wasscher (1941) gave a similar range except that he did not

include any material from New Guinea. Further study of this Rossel Island plant is

therefore needed but it does not belong to either Podocarpus spathoides as delimited

here or to P. orarius.

Henderson 186 (K) from the Solomon Islands (Malaita, Maluu), seen only as

an image, has much more tapering and narrower leaves than is typical for Podocarpus

orarius. Its original label bears the identification Podocarpus insularis but the specimen

more resembles P. salomoniensis than either P. insularis or P. orarius. However it is

not typical of P. salomoniensis either and examination of the actual specimen is needed

before a definite identification can be made.

The work reported here clearly demonstrates that Podocarpus spathoides as

delimited by de Laubenfels (1985, 1988) was a mixture of at least three taxa and that P

insularis may well have been a mixture too. In the case of P. spathoides, de Laubenfels

himself acknowledged the possibility that his concept of that species might encompass

more than one taxon. Despite that, however, the apparently very disjunctly scattered

distribution of Podocarpus spathoides sensu de Laubenfels (1988) has been regarded

as following a biogeographic pattern by Heads (2001, 2003). In the second of those

papers (Heads 2003, Fig. 87) this pattern was graphically illustrated by two lines,

one connecting the Malay and Rossel I. populations and the other linking Morotai

with the Solomons, based on the comments by de Laubenfels (1988) concerning bud

size. The present paper has shown that the Malay, Morotai, Rossel I. and Solomons

plants all belong to different taxa, not all of which can at present be given formal

names due to the insufficiency in quality and/or quantity of the Morotai and Rossel

I. material for description or identification. Consequently, the ‘pattern’ identified by

Heads (2003) is in this instance false although in his earlier paper (Heads 2001) he did

give other examples of species that show a disjunction between the Malay Peninsula

and the Solomons. The present paper emphasises how larger patterns and conclusions,

such as in forming biogeographic patterns, might be better based on data generated

in unambiguous situations. Particularly in a currently poorly known family such as

Podocarpaceae, many of the apparently very wide disjunctions in distribution may

turn out to be fictional, based on incompletely understood taxonomy, as in the case of

P. spathoides that has in part been unravelled here. This also has implications when

assessing the conservation status of species such as Podocarpus spathoides and P.

insularis, and highlights that sound taxonomic research is also crucial in underpinning

such conservation studies.

ACKNOWLEDGEMENTS. We wish to acknowledge the help from Fred Pitisopa and his

staff of the Ministry of Forests, Honiara, Solomon Islands for their logistic support which

helped Royal Botanic Garden Edinburgh (RBGE) researchers access Podocarpus species in

the Solomon Islands. The curator of L is thanked for the loan of specimens, including the

type of Podocarpus spathoides, and for permission to undertake cuticle micromorphological

examination of the latter. Martin Gardner (RBGE) took the photographs of Podocarpus orarius

in the field that form Fig. 2A—E, 3A, 3E and 3F and also did its IUCN conservation assessment.

192 Gard. Bull. Singapore 64(1) 2012

Paulina Hechenleitner V. kindly photographed several specimens at Kew. Chris Jeffree

(University of Edinburgh) helped with isolation of the cuticle and he together with Stephen

Mitchell (University of Edinburgh) also assisted with the SEM work. Michelle Whiting’s work

was undertaken while she was in receipt of a NERC Studentship. Prof. David de Laubenfels is

thanked for helpful comments. The RBGE is supported by the Scottish Government Rural and

Environment Science and Analytical Services Division (RESAS).

References

Alvin, K.L. & Boulter, M.C. (1974) A controlled method of comparative study for Taxodiaceous

leaf cuticles. Bot. J. Linn. Soc. 69: 277-286.

Biffin, E., Conran, J.G., & Lowe, A.J. (2011) Podocarp evolution: a molecular phylogenetic

perspective. In: Turner, B.L. & Cernusak, L.A. (eds.), Ecology of the Podocarpaceae in

Tropical Forests, pp. 1-20. Smithsonian Contributions to Botany, No. 95. Washington

D.C.: Smithsonian Institution Scholarly Press.

Conran, J.G., Wood, G.G., Martin, P.G., Dowd, J.M., Quinn, C.J., Gadek, P.A., & Price,

R.A. (2000) Generic relationships within and between the Gymnosperm families

Podocarpaceae and Phyllocladaceae based on an analysis of the chloroplast gene rbcL.

Austral. J. Bot. 48: 715-724.

de Laubenfels, D.J. (1985) A taxonomic revision of the genus Podocarpus. Blumea 30: 251—

278.

de Laubenfels, D.J. (1988) Coniferales. In: van Steenis, C.G.G.J. & de Wilde, W.J.J.O. (eds)

Flora Malesiana 10(3). Dordrecht, Boston & London: Kluwer Academic Publishers.

Donnelly, R., Neville, D. & Mous, P.J. (eds.) (2003) Report on a rapid ecological assessment

of the Raja Ampat Islands, Papua, Eastern Indonesia, held October 30—November 22,

2002. Final Draft November 2003. http://www.coraltrianglecenter.org/downloads/

RajaAmpatREA FinalDraft%20Screen.pdf#page=101 (accessed on 24 Jan. 2011).

Farjon, A. (2010a) A Handbook of the World’s Conifers. Leiden & Boston: E.J. Brill.

Farjon, A. (2010b) Podocarpaceae. In Kiew, R., Chung, R.C.K., Saw, L.G., Soepadmo, E. &

Boyce, P. (eds.), Flora of Peninsular Malaysia. Series II: Seed Plants, volume 1, pp.

171-201. Selangor, Kepong: Forest Research Institute Malaysia.

Foreman, D.B. (1971) A Checklist of the Vascular Plants of Bougainville with Descriptions

of Some Common Forest Trees, Botanical Bulletin (No. 5). Lae: Department of Forestry.

Gray, N.E. (1958) A taxonomic revision of Podocarpus XI. The South Pacific species of section

Podocarpus, subsection B. J. Arnold Arbor. 39: 424-477.

Heads, M. (2001) Birds of paradise, biogeography and ecology in New Guinea: a review. J.

Biogeogr. 28: 893-925.

Heads, M. (2003) Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology.

Telopea 10: 311-449.

International Union for Conservation of Nature (IUCN). (2001) IUCN Red List Categories and

Criteria: Version 3.1. Gland, Switzerland & Cambridge, U.K.: IUCN Species Survival

Commission.

International Union for Conservation of Nature (IUCN) Standards and Petitions Working Group.

(2010) Guidelines for Using the IUCN Red List Categories and Criteria. Version 8.0.

Prepared by the Standards and Petitions Working Group of the IUCN SSC Biodiversity

Assessment Sub-Committee in March 2010. http://intranet.iucn.org/webfiles/doc/SSC/

RedList/RedListGuidelines.pdf (accessed on 11 Jun. 2010).

Podocarpus orarius sp. nov. from Solomon Islands 193

Kershaw, D.A. (1997) An investigation into the relationships of Nageia s.l. (Gymnospermae,

Podocarpaceae). Unpublished M.Sc. Thesis. University of Edinburgh & Royal Botanic

Garden Edinburgh, Edinburgh, Scotland, U.K.

Knopf, P., Schulz, C., Little, D.P., Stiitzel, T., & Stevenson, D.W. (2012) Relationships within

Podocarpaceae based on DNA sequence, anatomical, morphological, and biogeographical

data. Cladistics 28: 271-299.

Pippard, H. (2008) The Pacific islands: an analysis of the status of species as listed on the 2008

IUCN Red List of Threatened Species™. IUCN. http://cmsdata.iucn.org/downloads/the_

pacific islands an analysis of the status of species as listed on the 2008 iucn r.

pdf & http://www.sprep.org/att/IRC/eCOPIES/Pacific_Region/483.pdf (both accessed on

16 Nov. 2010).

Schmid, M. (1975) La flore et la végétation de la partie méridionale de |’ Archipel des Nouvelles

Hébrides. Philos. Trans., Ser. B 272: 329-342.

Silba, J. (1986) Encyclopaedia Coniferae, Phytologia Memoirs 8. Oregon, Corvallis: Harold N.

Moldenke & Alma L. Moldenke.

Silba, J. (2000) Variation geographic et populations isole de les gymnospermes rarissime. JJ.

Int. Conifer Preserv. Soc. 7(1): 17-40.

Sinclair, W.T., Mill, R.R., Gardner, M.F., Woltz, P., Jaffré, T., Preston, J., Hollingsworth,

M.L., Ponge, A., & Méller, M. (2002) Evolutionary relationships of the New Caledonian

heterotrophic conifer, Parasitaxus usta (Podocarpaceae), inferred from chloroplast

trnL-F intron/spacer and nuclear rDNA ITS2 sequences. P/. Syst. Evol. 233: 79-104.

Stark Schilling, D.M. (2004) Taxonomic studies of Caribbean and Central American species

of Podocarpus subgenus Podocarpus: A multi-disciplinary approach. Unpublished

M.Sc. Thesis. University of Edinburgh & Royal Botanic Garden Edinburgh, Edinburgh,

Scotland, U.K.

Stockey, R.A., Frevel, B.J., & Woltz, P. (1998) Cuticle micromorphology of Podocarpus,

subgenus Podocarpus, section Scytopodium (Podocarpaceae) of Madagascar and South

Africa. Int. J. Pl. Sci. 159: 923-940.

Turner, I.M., Xing, F. & Corlett, R.T. (2000) An annotated check-list of the vascular plants of

the South China Sea and its shores. Raffles Bull. Zool. Suppl. 8: 23-116.

Wasscher, J. (1941) The genus Podocarpus in the Netherlands Indies. Blumea 4(3): 359-481.

Whiting, M. (2009) Cuticular micromorphology of Podocarpus as a_ systematic tool.

Unpublished M.Sc. Thesis. University of Edinburgh & Royal Botanic Garden Edinburgh,

Edinburgh, Scotland, U.K.

Whitmore, T.C. (1966) Guide to the Forests of the British Solomon Islands. Oxford: Oxford

University Press for British Solomon Islands Protectorate.

World Wildlife Fund (WWE). (2001) Solomon Islands rain forests (AAO119). http://www.

worldwildlife.org/wildworld/profiles/terrestrial/aa/aa0119_full-html (accessed on 19

Aug. 2010).

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Gardens’ Bulletin Singapore 64(1): 195-200. 2012 195

Distichochlamys benenica (Zingiberaceae),

a new species from Vietnam

Quoc Binh Nguyen! and Jana Leong-Skorniékova?

‘Vietnam National Museum of Nature,

18 Hoang Quoc Viet Street, Cau Giay, Hanoi, Vietnam

>The Herbarium, Singapore Botanic Gardens,

1 Cluny Road, 259569 Singapore

jana_skornickova@seznam.cz

ABSTRACT. Distichochlamys benenica (Zingiberaceae) from north Vietnam is described.

Colour plates are provided and the key to Distichochlamys species is updated.

Keywords. Distichochlamys, Vietnam, Ben En National Park

Introduction

The small ginger genus Distichochlamys M.F.Newman is endemic to Vietnam. The

type species of the genus, D. citrea M.F.Newman, was based on collections from central

Vietnam, Bach Ma National Park (Newman 1995). Since then, another two species, D.

orlowii K. Larsen & M.F.Newman (Larsen & Newman 2001) from central Vietnam,

Gia Lai Province, and D. rubrostriata W.J.Kress & Rehse (Rehse & Kress 2003) from

northern Vietnam, Cuc Phuong National Park, have been described. Morphologically,

Distichochlamys resembles Scaphochlamys Baker but differs by the distichous

arrangement of the bracts and tubular bracteoles (vs. spiral bracts and open bracteoles

in Scaphochlamys). Larsen & Newman (2001) suggested that placing Distichochlamys

in synonymy under Scaphochlamys could not be ruled out yet as early molecular

analyses did not seem to be conclusive (Searle & Henderson 2000; Ngambriabsakul

2001). In the more recent studies of Kress et al. (2002) and Ngamriabsakul et al.

(2004), the two genera appear to be distinct sister-clades. The geographic disjunction

of over 1000 km between Distichochlamys and Scaphochlamys, which is confined to

Malaysia and S. Thailand, provides additional support for retaining them as separate

genera.

During recent exploration of northern Vietnam, we encountered a new

Distichochlamys species, which is described and illustrated below. The key to the

species of Distichochlamys given by earlier authors (Larsen & Newman 2001; Rehse

& Kress 2003) is modified to include the latest addition.

Distichochlamys benenica Q.B.Nguyen & Skorni¢k., sp. nov. (Fig. 1 & 2)

Distichochlami rubrostriatae similis, robustior ad 60 cm alta (contra ad 30 cm),

staminodiis lateralibus luteis (contra luteis maculis duabus linearibus atrorubris),

labello in parte dimidia basali macula rubella (contra clare lutea) differt. TYPE:

196 Gard. Bull. Singapore 64(1) 2012

Vietnam, Thanh Hoa Province, Nhu Thanh District, Ben En National Park, altitude

c. 120-150 m, mixed evergreen forest, limestone, 5 April 2011, Nguyen Q.B.

VNM-B0001352 (holo VNMN; iso E, HN, P, SING).

Terrestrial rhizomatous herb to 60 cm tall. Rhizome sympodially branching, c. 7-13

mm in diam., light brown externally, cream white to ochraceous internally with

prominent citrol smell, sheathing bracts not seen (decaying very soon and leaving

only scars on mature rhizomes), root tubers fusiform, 3—8 cm long, 7-10 mm in

diam., light brown externally, cream white to ochraceous internally. Leafy shoots with

a single leaf, tightly arranged forming clumps. Sheathing bracts |—3, inner one to 6

x c. 2 cm wide, outer ones gradually smaller, papery and decaying fast by the time of

flowering, glabrous. Leaf sheaths 3—5 cm, deep purple, glabrous. Ligule bilobed, c. 3

mm long, papery, glabrous, soon decaying. Petiole 8—25 cm long, canaliculate, dark

red-purple, glabrous. Lamina oval to obovate, unequal, 15—28 x 10—14.5 cm, plicate,

green above, lighter green with purple shading at the apex beneath, apex subacute

to nearly rounded, base rounded to slightly cordate. Inflorescence to 15 cm, partly

hidden in leaf sheath. Peduncle 3—6 cm, c. 4 mm in diam., glabrous, with 2-3 sterile

bracts, the outer ones papery, to 6 cm long, decaying fast, the inner ones red purple, c.

3—5 cm long, glabrous, 2 cm broad, apex acuminate. Spike 6—10 cm long, consisting

of 8-13 bracts. Bracts ovate with acute apex; with more or less pink-red tinge, 1.8—3.2

x ]—2.2 cm, glabrous, enclosing cincinnus of 2—3 flowers. Bracteoles 13—23 mm long,

8-20 mm in circumference, tubular in basal 2-6 mm, with one keel, translucent white

with pink tinge at apex, sparsely shortly hairy. Calyx 15—19 mm long, translucent

white with slight pink tinge, glabrous but sparsely hairy at apex (margins of teeth), 3

teeth, unilateral slit 8-10 mm. Floral tube 18—21 mm, pinkish at base, light yellow

towards apex, glabrous externally. Dorsal corolla lobe ovate, 16—23 * 6—7 mm, semi-

translucent yellow with red tinge at apex, glabrous, apex mucronate, mucro 2.5 mm,

shortly hairy. Lateral corolla lobes ovate, 16-20 x 6-8 mm, semi-translucent yellow

with red tinge at apex, apex slightly concave, glabrous. Labellum broadly spathulate,

bilobed at apex, 21—23 mm long, 18—21 mm broad at apex, c. 5 mm broad at base, lobes

more or less rounded, split c. 7-8 mm, yellow with red-orange central patch, with short

glandular hair. Lateral staminodes obovate, 20-24 x 7-10 mm, yellow, covered with

glandular hairs. Stamen 7 mm long, filament 2.5—3 mm long, pinkish at base, light

yellow towards apex, glabrous, anther 5 mm long, anther thecae dehiscing along their

entire length, connective tissue deep yellow, covered with glandular hairs, anther crest

c. | mm long. Epigynous glands two, 4-6mm, cream white sometimes with slight

pink tinge at base, ochraceous towards apex, sometimes connate. Ovary trilocular with

axile placentation, 2-3 mm long, 2 mm in diam., cream white with slight pink tinge

at apex, densely hairy. Style white, glabrous, stigma white, apex ciliate, quadrangular

with transverse ciliate ostiole. Fruits unknown.

Habitat and phenology. This species occurs in evergreen broad-leaved mixed forest,

on limestone, at elevations about 100—200 m. It flowers in March to April.

1ys benenica, anew Vietnamese ginger

Fig. 1. Distichochlamys benenica Q.B.Nguyen & Skorni¢k.: A & B. Habit. C. Close-up of

flowers. Photos: Q.B.Nguyen (A); Jana Leong-Skorni¢kova (B, C).

198 Gard. Bull. Singapore 64(1) 2012

Fig. 2. Distichochlamys benenica Q.B. Nguyen & Skorniék.: A. Rhizome. B. Root tubers.

C. Flower in bract (far left) and floral dissection (from left to right): bract, bracteole, corolla

lobes, staminodes and labellum, floral tube with stamen, calyx, ovary with epigynous glands.

D. Detail of ovary, calyx, floral tube and stamen. Photos: Jana Leong-Skorniékova.

Distribution and IUCN assessment. We have examined all Distichochlamys specimens

available at AAU, E, HN, SING and VNMN, but discrimination of Distichochlamys

species from dried material, if not accompanied by spirit collection, is challenging.

From all data available it appears that all Distichochlamys species, including the

newly described D. benenica, are rather restricted in their distribution and therefore

susceptible to any habitat changes.

Distichochlamys benenica, a new Vietnamese ginger 199

Distichochlamys benenica is so far known only from the type locality in Ben

En National Park, which has about 85 km of primary vegetation in a total area of about

166 km”. The primary habitat outside the park has been destroyed. We estimate that the

area of occupancy of this species within Ben En National Park is less than 20 km? and

therefore propose to treat this species provisionally under category Vulnerable (VU):

D2.

Etymology. The specific epithet is derived from the type locality, Ben En National

Park.

Key to Distichochlamys species

la. Inflorescence bracts spreading, loosely imbricate: labellum deeply cleft to c. half

OS EEE 2 tect etc ee D. citrea

b. Inflorescence bracts appressed to floral axis, densely imbricate: labellum divided

mibeieinexcnuine less than half its length ...............2.......s0c.-csssccseccssecescsesceneeee 2

2a. Lateral staminodes yellow with two red linear patches at the base .....................0...

0 RSE, SECT ES 7S | 3

3a. Labellum with red patch at base and two round lobes at apex ............ D. benenica

b. Labellum yellow with purple veins, dark yellow medium band and two emarginate

DD ES cece: pt on D. orlowii

ACKNOWLEDGEMENTS. We thank the curators of AAU, E, HN, P, SING and VNMN for

letting us examine the specimens in their care, the Asian Zingiberaceae Information Centre

at the Singapore Botanic Gardens and the Zingiberaceae Resource Centre at the Royal

Botanic Garden Edinburgh (<http://elmer.rbge.org.uk/ZRC/>) for providing protologues and

related references. The first author thanks the authorities, in particular Mr. Le Duc Thuan, for

permission to carry out research in Ben En National Park and Prof. Luu Dam Cu (VNMN) for

providing facilities for research. The research of the second author is funded by the National

Parks Board, Singapore and the Czech Science Foundation, GACR (grant nos. 521/09/0202

and P506/10/0623). We thank Dr J. F. Veldkamp (L) for translating the diagnosis into Latin,

and the two reviewers, Dr. Mark Newman (E) and Tran Hiru Dang, for comments on this

manuscript.

References

Kress, W.J., Prince, L.M. & Williams, K.J. (2002) The phylogeny and a new classification of

the gingers (Zingiberaceae): evidence from molecular and morphological data. Amer: J.

Bot. 89: 1682-1696.

200 Gard. Bull. Singapore 64(1) 2012

Larsen, K. & Newman, M. (2001) A new species of Distichochlamys from Vietnam and some

observations on generic limits in Hedychiae (Zingiberaceae). Nat. Hist. Bull. Siam Soc.

49:77.

Newman, M.F. (1995) Distichochlamys, a new genus from Vietnam. Edinburgh J. Bot. 52(1):

65-69.

Ngambriabsakul, C. (2001) The systematic of the Hedychieae (Zingiberaceae), with emphasis

on Roscoea Sm. Unpublished Ph.D. thesis. University of Edinburgh.

Ngamriabsakul, C., Newman, M.F. & Cronk, Q.B. (2004) The phylogeny of tribe Zingibereae

(Zingiberaceae) based on ITS (nrDNA) and t#rnL-F (cpDNA) sequences. Edinburgh J.

Bot. 60(3): 483-507.

Rehse, T. & Kress, W.J. (2003) Distichochlamys rubrostriata (Zingiberaceae), a new species

from Northern Vietnam. Brittonia 55(3): 205—208.

Searle, R.J. & Henderson, T.A. (2000) A preliminary phylogeny of the Hedychieae tribe

(Zingiberaceae) based on ITS sequences of the nuclear rRNA cistron. In: Wilson, K.L.

& Morrison, D.A. (eds) Monocots, Systematics and Evolution. Australia, Cillongwood:

CSIRO.

Gardens’ Bulletin Singapore 64(1): 201-209. 2012 201

Four new species of Aspidistra Ker Gawl. (Asparagaceae)

from China and Vietnam with a comment on A. longifolia

Hook.f. and A. hainanensis W.Y.Chun & F.C.How

Hans-Juergen Tillich' and Leonid V. Averyanov’

‘Ludwig-Maximilians-University, Systematic Botany,

Menzinger Str. 67, D-80638 Munich, Germany

hjtillich@gmx.de

*Komarov Botanical Institute, Russian Academy of Sciences,

Prof. Popov Str. 2, St. Petersburg 197376, Russian Federation

ABSTRACT. Four new species of Aspidistra Ker Gawl. (Asparagaceae) are described and

illustrated: A. basalis Tillich, A. columellaris Tillich, A. gracilis Tillich from China, and A.

coccigera L.V.Averyanov & Tillich from Vietnam. The application of the name 4. longifolia

Hook.f. to plants from SE Asia and the intraspecific variability of A. hainanensis W.Y.Chun &

F.C.How across its range from peninsular Malaysia to SE China is also discussed.

Keywords. Asparagaceae, Aspidistra, China, Malay Peninsula, Thailand, Vietnam

Introduction

The genus Aspidistra Ker Gawl. (Asparagaceae: Nolinoideae) is distributed from

Assam (India) in the west to southern Japan in the east, and from Central China

southwards to the Malay Peninsula but the centre of diversity is SE China (Guangxi

Province) and adjacent northern Vietnam. During the past three decades the number of

known species has increased considerably from 11 in 1980 to more than 100.

The recognition of new Aspidistra species is ongoing and stems largely from

more widespread collecting, especially in remote areas of SE Asia combined with

more detailed study of living plants. Since the publication of a comprehensive key to

the genus (Tillich 2008), additional new species have been published (Hou et al. 2009;

Lin et al. 2009, 2010; Lin & Liu 2011; Xu et al. 2010).

As a result of this ongoing work, a further four new species can now be added.

One of them was collected by the second author in Vietnam, while three more species

have been in cultivation for several years at the Juniper Level Botanical Gardens,

Raleigh, North Carolina, USA. This Garden obtained the plants from several collectors,

mostly with incomplete collection location details. Cuttings of these plants were sent

to the Botanical Garden Munich and added to the rich Aspidistra collection where they

grew vigorously and came to flower during the summer and autumn of 2009 and 2010.

Once they flowered, it was apparent that these plants represented four clearly different

and undescribed species (see Tillich 2008) and this paper accordingly names them

formally.

202 Gard. Bull. Singapore 64(1) 2012

Aspidistra basalis Tillich, sp. nov. (Fig. 1D & 3H, I.)

Species Aspidistra lurida Ker-Gawl. similis, sed differt pedicello 0.5 cm longo, floribus

ad solum prostratis, stigmate convexo, laeve, 8 maculis albis periphericis proviso,

lamina foliorum ca. 40 x 4-5 cm. TYPE: Cultivated plant in the Munich Botanical

Garden, originally from the Nanjing Botanical Garden, Jiangsu Province (?), China,

Tillich 5720 (M, including flowers in the liquid collection).

Rhizome epigeous, diameter 8—10 mm, richly branched, covered with deltoid scales.

Cataphylls up to 6 cm long. Leaves solitary, 2—5 cm apart. Petiole 3—5 cm, stout, stiff,

ventrally with a deep v-shaped furrow. Blade narrowly lanceolate, 35-45 x 4-5 cm,

without clear limitation, tapering gradually to petiole, sharply folded at base, dark

green, shiny, margin finely serrate and revolute, midrib sharply protruding abaxially.

Peduncle up to 0.5 cm, or flower subsessile. Perigone bowl-shaped, blackish purple

inside and out. Lobes 8, in two whorls of 4, outer lobes deltoid, 6-7 mm long and

wide, inner lobes 6-7 x 5 mm, more or less romboid distally. All lobes with 4 low

basal keels, the submarginal keels connected to those of the adjoining lobe, the two

median keels disappearing in the middle of the tube. Stamens 8, inserted at tube base,

subsessile, anthers broadly ovate, ca. 1.5 mm long. Pistil mushroom shaped, 5—6 mm

high. Stigma circular, c. 10 mm in diameter, smooth, convex, light red, the margin

with 8 soft strongly reflexed lobes and 8 submarginal, white-edged, radially elongated,

weak indentations (Fig. 3H). Fruit unknown.

Etymology. The species name relates to the flower position adjacent to the soil or litter

surface.

Notes. The species is similar to 4. /urida Ker Gawl. but differs clearly by its shorter

pedicels (0.5 cm), placing the flowers 1n horizontal position at ground level, the pink

(not white) stigma with its marginal lobes bent downwards nearly to the base of the

perigone tube, and the v-shaped leaf blades in cross-section, c. 40 x 4—5 cm (not flat,

15—20' x 3-5 cm):

The original plant from the Nanjing Botanical Garden was found in cultivation

along with A. minutiflora Stapf and A. retusa K.Y.Lang & S.Z.Huang, so it was

surprising that this remained undescribed.

Aspidistra coccigera L.V.Averyanov & Tillich, sp. nov. (Fig. 2C—G)

Species Aspidistra marasmioides H.-J.Tillich similis, sed rhizomate breve nec longe

repente, petiolo 17-20 cm longo, lamina 15—18 * 6—7.5 cm, perigonio late cupiforme

20 mm diametro. TYPE: Vietnam, Quang Binh Province, Minh Hoa District, Dan Hoa

Municipality, Chuong Lon Mountain, Vietnam-Laotian border, about 6 km N of Cha

Lo border station, around point 17°44’04” N, 105°46°53” E; L.V.Averyanov, P.K.Loc,

N.T. Vinh & N.S.Khang HAL 11693, , 19 April 2008 (holo HN; iso LE, M).

19>)

Four new species of Aspidistra

10 mm 2mm

Fig. 1. A-B. Aspidistra columellaris. A. Flower longitudinally dissected. B. Stigma in upper

view. C—E. Rhizomes and typical leaves. C. A. columellaris. D. A. basalis. E. A. gracilis.

204 Gard. Bull. Singapore 64(1) 2012

Rhizome with very short internodes, leaves crowded, cataphylls up to 5 cm long. Petiole

17-25 cm long, stiff, blade ovate-acuminate, base rounded, 15—18 = 6.0—7.5 cm, light

green, kept obliquely upright. Peduncle 2.5—6 cm long, erect or ascendant, purple,

with 5 purple scales, 2 subtending the flower and 3 along the scape. Perigone widely

cupuliform. Tube diameter 15-17 mm, white externally, internally blackish purple

in upper third, white below. Lobes 6, (blackish-) purple, reflexed, smooth. Stamens

inserted at mid-tube, filaments 1.5 mm long, protruding horizontally, anthers ovoid,

1.5 mm long. Pisti! mushroom-like, c. 13 mm long, slightly exceeding the perigone,

ovary depressed-conical, basal diameter c. 4 mm, style delicate, cylindrical, white,

stigma circular, prominently convex, smooth, bright pink. Fruit depressed-globose,

diameter c. 1.5 cm, deep purple, softly echinate.

Ecology. The species grows in primary broad-leaved forest on very steep slopes and

rocky cliffs of remnant mountains of solid marble-like highly eroded limestone at

elevations of 400-750 m az.s.l. The species is locally abundant.

Etymology. The species name relates to its purplish-red fruits.

Notes. The flowers of this species are similar to 4. marasmioides Tillich from Haiphong

Province, Vietnam. The latter species is distinguished by its richly branched, long

creeping rhizomes, numerous leaves forming a dense carpet of horizontally oriented

blades, and the flowers completely hidden below the leaves. In 4. coccigera, there are

clusters of fewer, much larger leaves, the blades are erect to semi-erect and the flowers

and their bright pink stigmas are clearly displayed.

Aspidistra columellaris Tillich, sp. nov. (Fig. 1A—C; 2A, B)

Species Aspidistra leyeensis Y.Wan & C.C.Huang similis, sed differt perigonio 12

mm diametro, lobis erectis 4—5 mm longis, lamina foliorum scortea, ovalis, 1S—20 x

5—7 cm. TYPE: Cultivated plant in the Munich Botanical Garden, collected in China

(Province and collector unknown), Ji/lich 5719 (M, including flowers in the liquid

collection).

Rhizome epigeous, diameter 5 mm, with soon decaying scales. Cataphylls up to 7

cm long. Leaves solitary, 1-1.5 cm apart. Petiole slender, 20-35 cm. Blade ovate-

acuminate, coriaceous, 18-20 x S—7 cm, with numerous white spots. Peduncle 0.5—2.5

cm, with 4-5 purple spotted scales. Perigone cupulate, thick-walled, changing from

green to beige externally, lobes abaxially spotted brownish-red. Tube 7-8 mm high,

12-15 mm in diameter, finely verrucose, purple-black internally. Lobes 8, deltoid,

c. 4 mm long and wide, fleshy, erect to somewhat recurved, adaxially purple with

beige tips, finely verrucose, with two low verrucose keels, each keel approaching the

neighboring keel of the adjacent lobe and extending to the base of the tube. Stamens 8,

inserted in the lower third of the tube, filaments short, directed obliquely downwards,

anthers 2.5-3.0 mm, hooked around filaments, oriented downwards. Pistil 5 mm high,

Four new species of Aspidistra

Fig. 2. A, B. Aspidistra columellaris. A. Flower. B. Flower longitudinally dissected, and stigma

in upper view. C-G. Aspidistra coccigera. C. Plant at natural habitat. D. Plants with flowers and

fruits prepared for herbarisation. E. Fruits. F. Flowers in upper view. G. Flower longitudinally

dissected. Bar scale: A, B, E-G: 1 cm; C, D: 10 cm.

206 Gard. Bull. Singapore 64(1) 2012

flat-topped. Style a stout column with 4 longitudinal groves alternating with the stigma

lobes, dark purple. Stigma 7—8 mm in diameter, delicate, thin, upper surface purple,

finely verrucose,cruciform with 4 arms bifurcating to 8 flattened, pale lobes (Fig. 1B

& 2B).

Etymology. The name relates to the column-like style.

Notes. The species is similar to A. /eyeensis Y.Wan & C.C.Huang with the stigma

divided into separate lobes, but is clearly distinguished mainly by the characters in

Table 1.

Table 1. Characteristics distinguishing A columellaris and A. leyeensis.

A. columellaris A. leyeensis

Leaf blade ovate, c. 20 x 5-7 cm lanceolate, c. 40 x 4-6 cm

Perigone diameter 12 mm 20-25 mm

Perigone lobes erect reflexed

stout, with 4 longitudinal

Style slender, smooth

grooves

: divided into 4 flat, slightly divided into 8 narrowly-ovate

Stigma as

bifurcate lobes lobes

Aspidistra gracilis Tillich, sp. nov. (Fig. 1E & 3F, G)

Species A. oblanceifolia FT. Wang & K.Y. Lang similis, sed differt foliis anguste

lanceolatis, lamina 25-30 * 2—2.5 cm, pedunculo erecto, 3—4.5 cm longo, ovario

albo, stigmate convexo, laeve. TYPE: Cultivated plant in the Munich Botanical

Garden, collected in China, Hong Kong S.A.R., Lantau Island, Sunset Peak (collector

unknown), Tillich 5718 (M, including flowers in the liquid collection).

Rhizome thin, with very short internodes, numerous /eaves and dry petioles of decayed

leaves densely clustered together, petiole 15—20 cm, semi-circular in cross section,

blade narrowly lanceolate, 25—30 x 2.0-2.5 cm, light green with scattered paler green

spots. Peduncle erect, 3.0-4.5 cm, with 3 scales along the scape and 2 subtending the

flower. Flower obliquely erect to horizontal. Perigone campanulate, 14-16 mm long,

12-14 mm in diameter, grey-green or reddish-purple mottled externally, smooth and

purple internally. Lobes 8, deltoid, straight or slightly spreading, thick-fleshy, 3.04.0

mm long and wide, finely verruculose adaxially. Stamens 8, inserted near tube base.

Pistil mushroom shaped, ca 10 mm long, white, stigma circular, conspicuously convex,

smooth. Fruit depressed-globose, pale green, mottled with red, c. 15 mm in diameter.

Etymology. The name relates to the dainty flowers, borne on an erect peduncle in

horizontal position.

Four new species of Aspidistra 207

Fig. 3. A-E. Aspidistra hainanensis W.Y.Chun & F.C.How s.1. A—-C. Plant from Thailand. A.

Habit, note the many white spots on leaves. B. Flower longitudinally dissected. C. Flower in

upper view, note the stigma composed of irregularly folded lobes. D, E. Plant from Laos. D.

Flower in longisection. E. Flower in upper view, note the fine line pattern on the smooth stigma

surface. F, G. Aspidistra gracilis. F. Flower longitudinally dissected. G. Flower in natural

position. H, I. Aspidistra basalis. H. Flower in upper view. I. Flower longitudinally dissected.

Bar scale: A: 20 cm; B—I: | cm.

208 Gard. Bull. Singapore 64(1) 2012

Notes. The species is similar to A. oblanceifolia F.T.Wang & K.Y. Lang and A. insularis

Tillich, but is clearly distinguished as shown in Table 2.

Table 2. Characteristics distinguishing A gracilis from two related species.

A. gracilis A. oblanceifolia A. insularis

Leaf blade 25-30 x 22.5 em 35-50 x 2.5—-4 cm 40-50 x 7-8 cm

Peduncle 34.5 cm 0.3—2 cm 0.2—0.5 cm

Flower position horizontal vertical vertical

Perigone shape narrow-campanulate campanulate cupuliform

Stigma colour white purple purple

’ convex, circular conical, 8-lobed, square, with 4

Siemasshape smooth smooth prominent radial ribs

Notes on Aspidistra hainanensis W.Y.Chun & F.C.How in SE Asia

In SE Asia there is a group of Aspidistra species that ranges from peninsular Malaysia

through Thailand and Southwest-China (Yunnan) eastwards to Laos, Hainan, and

Southeast-China (Guangxi) and 1s characterised by tufted, oblanceolate to linear

leaves. The flowers of plants from different sites across that region are similar at first

sight, bearing a campanulate to slightly urceolate perigone and mushroom-shaped

pistils. However, they are distinguishable by several minor characters, such as the

number and course of the adaxial perigone ribs, details of the stigma surface, and in

colour patterns. The leaves also vary from variously spotted to homogeneously green.

As an example, two floral variants are shown in Fig. 3. The plant in Fig. 3A—C

was purchased from a Thai flower market and is now in cultivation at the Botanical

Garden Munich. The plant in Fig. 3D—E was collected by H. Billensteiner in Laos,

Bokeo Province, and is now in cultivation at the Palmengarten Frankfurt/M. During the

past several decades, some of these local variants have been described as new species:

A. hainanensis W.Y.Chun & H.W.How, A. vingjiangensis Peng, A. larutensis deWilde

& Vogel (Chun & How 1977, Peng 1989, de Wilde & Vogel 2005) and Phonsena &

de Wilde (2010) recently discussed the taxonomy of this group. We agree with the

conclusion of Phonsena & de Wilde (2010) that excessive splitting is unwarranted, and

that presumably “local Aspidistra populations represent clones of uniform plants, each

clone conserving its characteristics against those of remote populations”.

Experience with Aspidistra has shown that taxonomically meaningful

solutions can be obtained only by studying plants from a great number of well-

documented collection sites side by side in cultivation and this is one of the best

ways to circumscribe taxa. However, there is a problem in that Phonsena & de Wilde

(2010) placed all these SE Asian plants into 4. longifolia Hook.f., a species decribed

from Assam, India, and based on two specimens from the Griffiths Herbarium at Kew

Four new species of Aspidistra 209

(Hooker 1892). One of these specimens (Griffith 5887, barcode 000099915) was

designated twice as Type, initially by Tillich (2008) and independently by Phonsena

& de Wilde (2010). Unfortunately, neither of these herbarium specimens are suitably

preserved for detailed flower analysis but Hooker’s description leaves no doubt that

A. longifolia is clearly different from the SE Asian plants as he states “perianth 1/2 in.

diam., fleshy, subglobose contracted at the mouth; lobes very small”. This character

combination clearly differs from the campanulate to slightly urceolate flowers with

lobes (nearly) as long as or longer than the tube, which is the condition seen in all the

SE Asian plants so far examined.

We therefore propose to regard the SE Asian Aspidistra specimens with

oblanceolate to lineate, tufted leaves as part of the 4. hainanensis W.Y.Chun &

H.W.How complex, until more comprehensive cultivation experiments can provide

clarity regarding their variability and taxonomic status.

ACKNOWLEDGEMENTS. We are grateful to Prof. D. Podlech (Munich) for the Latin

diagnoses, and Todd Rounsaville (Raleigh, NC, USA) and Clemens Bayer (Palmengarten

Frankfurt) for providing plant material.

References

Chun, W.Y. & How, F.C. (1977) Aspidistra Ker Gawl. Flora Hainanica 4: 114-115, 533.

Hou, M.F., Liu, Y., Kono, Y. & Peng, C.-I. (2009) Aspidistra daxinensis (Ruscaceae), a new

species from limestone areas in Guangxi, China. Bot. Stud. (Taipei) 50: 371-378.

Lin, C.-R., Liang, Y.-Y. & Liu, Y. (2009) Aspidistra bamaensis (Ruscaceae), a new species

from Guangxi, China. Ann. Bot. Fenn. 46: 416-418.

Lin, C.-R., Peng, C.-I., Kono, Y. & Liu, Y. (2010) Aspidistra obconica, Asparagaceae

(Ruscaceae), a new species from limestone areas in Guangxi, China. Bort. Stud. (Taipei)

51(2): 263-268.

Lin, C.-R. & Liu, Y. (2011) Aspidistra punctatoides sp. nov. (Ruscaceae) from limestone areas

in Guangxi, China. Nord. J. Bot. 29: 189-193.

Peng, L. (1989) One new species of the genus Aspidistra from Yunnan. Acta Bot. Yunnan.

11(2): 173-174.

Phonsena, P. & de Wilde, W.J.J.O. (2010) The genus Aspidistra Ker Gawl. (Asparagaceae/

Ruscaceae) in Thailand. Thai Forest Bull., Bot. 38: 48-58.

Tillich. H.-J. (2008) An updated and improved determination key for Aspidistra Ker Gawl.

(Ruscaceae, Monocotyledons). Feddes Repert. 119: 449-462.

de Wilde, W.J.J.O. & Vogel, A. (2005) A new species of Aspidistra (Convallariaceae) from

Perak, Peninsular Malaysia. Folia Malaysiana 6(3-4): 125-130.

Xu, W.F., He, H.Z. & Yang. L. (2010) Aspidistra chishuiensis (Ruscaceae), a new species from

Guizhou, China. Ann. Bot. Fenn. 47: 118-120.

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Gardens’ Bulletin Singapore 64(1): 211-219. 2012 211

Phylogenetic study of the Schismatoglottis Nervosa

Complex (Araceae: Schismatoglottideae)

A.P.J. Ting'’, S.Y. Wong’, J. Jamliah! and P.C. Boyce®

‘Department of Plant Science & Environmental Ecology,

Faculty of Resource Science & Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan, Sarawak, Malaysia

*april_day15@yahoo.com

*Pusat Pengajian Sains Kajihayat (School of Biological Sciences),

Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Malaysia

ABSTRACT. The Schismatoglottis Nervosa Complex (Araceae: Schismatoglottideae)

currently comprises 10 species: Schismatoglottis adoceta S.Y.Wong, S. elegans A.Hay, S. liniae

S.Y.Wong, S. tessellata S.Y.Wong, S. ulusarikeiensis S.Y.Wong, S. matangensis S.Y.Wong, S.

simonii S.Y.Wong, S. turbata S.Y.Wong, and S. nervosa Ridl., occurring in Borneo, with each

endemic to a specific locality and most to a particular geology; and one species (S. brevicuspis

Hook.f.) widespread in Peninsular Malaysia and Sumatera, where it is restricted to granites.

Based on analysis of the matK region, a preliminary biogeographical hypothesis for the origins

and subsequent taxagenesis of the Nervosa Complex is presented. This study also provides

insight into possible evolution of localised mesophytic endemics in everwet, humid, and

perhumid megathermal Sundaic forests. Two clades are resolved: one north of, and another

south of, the Lupar Divide.

Keywords. Araceae, biogeography, Borneo, matK, Schismatoglottis Nervosa Complex,

vicariance

Introduction

Geology and tectonics of Borneo

Borneo is the third largest island in the world and the second largest tropical island

after New Guinea. Borneo is situated in a tectonically intricate region between three

marginal basins: the South China, Sulu and Celebes Seas (Hall et al. 2008), the latter

two on the eastern edge of the Sunda Shelf. Borneo has a complex geological history

having been formed by Mesozoic accretion of oceanic crustal material (ophiolite),

island arcs and microcontinental fragments accreted to the Palaeozoic continental core

of the now Schwaner Mountains (Hutchison 1989; Moss & Wilson 1998; Hall et al.

2008). Despite this rather active formation, Borneo is now a stable area with little or

no seismic activity, and has no active volcanoes (Hall 2002; Hall et al. 2008).

Plant endemism in Borneo

Borneo is frequently cited—rightly although nowadays somewhat repetitively—as

one of the world’s areas of ‘mega’-biodiversity. With a flora comprising at least 3000

212 Gard. Bull. Singapore 64(1) 2012

tree species alone (MacKinnon et al. 1996; Slik et al. 2003), Borneo’s floristic patterns

are significantly correlated with topography and geography (Slik et al. 2003). Present-

day floristic patterns are probably influenced by Pleistocene glaciations and are

suggested to be of recent emergence after the Pleistocene glacial period. The Southeast

Asian mainland and Sumatera were formerly connected to Borneo by land bridges

during the Pleistocene glacial (Morley 2000) and the emergence (more accurately,

re-emergence) of lowland humid forest is almost certainly owing to rapid radiation of

the relictual fragmentary floras from southeast Borneo (Meratus), and/or expansion of

forest remnants from the Riau Pocket (see Corner 1960; Ashton 2005). It has also been

suggested that Pleistocene refugia extended to north and northwest Borneo (Ashton

1972; Wong 1998), providing yet other points of taxon radiation.

It is estimated that c. 37% of Borneo’s 15,000 vascular plant species are

endemic (Roos et al. 2004; Welzen et al. 2005), although these figures are likely both

underestimates given that studies are consistently revealing that a high percentage of

the mesophytic flora appears to be endemic, and includes a great many novelties, e.g.,

the findings in Zingiberales alone: Theilade & Mood (1997, 1999), Nagamasu & Sakai

(1999) and Takano & Nagamasu (2007).

Sabah and Sarawak together are frequently cited to have the most endemic

plant species in Borneo, notably in mountainous areas (Moss & Wilson 1998).

Frequently it is stated that much of the plant endemism of Borneo occurs in areas

of the north, west Sarawak (notably), Sabah’s Crocker Range (including Kinabalu)

and Trusmadi Range, the central Bornean mountain chain (the Schwaner—Muller—

Iran Range), and also in the southeastern Meratus range (Mackinnon et al. 1996).

However, it is important to remember that the collecting density of much of highland

Borneo is almost infinitesimally small, and thus the taxonomic wealth of the numerous

smaller and isolated upland areas (e.g., the Kalimantan Kapuas Hilir, and flanking

(Sarawak) Klingkang range, the Kapuas Hulu, Sambiliung, etc.) has yet to be sampled

in any scientifically meaningful way. Furthermore, increasingly it is becoming

apparent that the /ow/and areas have, despite their frequently degraded condition, still

an extraordinary untapped reserve of novel taxa, many with highly restricted, often

geologically obligate, ranges.

The Schismatoglottis Nervosa Complex

Since 2000, the genus Schismatoglottis has been the focus of attention aimed at

resolving both its taxonomy and phylogeny (Hay & Yuzammi 2000). Wong (2010)

delimitated the Nervosa complex by the presence of aromatic vegetative tissues

(terpenoids), longitudinally ribbed petioles, and leaf blades with tessellate tertiary

venation. Seven novel species additional to the two (Schismatoglottis nervosa and

S. elegans) recognised by Hay & Yuzammi (2000) were proposed. In addition, S.

brevicuspis was recognised as belonging to the Nervosa complex.

Thus, as now defined the Nervosa Complex comprises 10 described species.

A further four species await description. Schismatoglottis nervosa and S. elegans are

Schismatoglottis Nervosa Complex yA)

restricted to Karst limestones in the southwest and northeast of Sarawak, respectively;

Schismatoglottis matangensis and S. turbata are species of sandstones, with S.

matangensis occurring on soft sandstones under perhumid moist forest, while S. turbata

occurs in humid forest where it is restricted to the tops of small Bornhardts comprised

of very hard sandstone. Schismatoglottis adoceta, S. tessellata and S. ulusarikeiensis

are restricted to shales, S. simonii is unusual in that it occurs on both limestones (the

type) and sandstones, while S. /iniae and S. brevicuspis are granite obligates. Of the

four species awaiting formal description, one is endemic to basalt in East Sabah, and

one each to limestone at Mulu, shale in west Kapit, and limestone in southern central

Bintulu.

This study investigates taxonomic relatedness within the Nervosa Complex

and possible correlation between phytogeographical patterns and phylogeny of the

complex, and presents a hypothesis of taxagenesis of the localised endemics.

Materials and methods

Sample collection and outgroup selection

The ingroup for the analysis consisted of 13 species (each with one accession, except

for S. simonii, from two localities) from Sabah and Sarawak and one species from

West Malaysia. Of these, 13 accessions were collected from the living collection

at Semenggoh Botanical Research Centre, Kuching, Sarawak, and one accession

was collected from Perak, Peninsular Malaysia, supplied through Universiti Sains

Malaysia, Pulau Pinang (see Table 1). The outgroup taxon selected was Apoballis

acuminatissima S.Y.Wong & P.C.Boyce, established as the Asian sister species to

Schismatoglottis (Wong et al. 2010).

DNA extraction, PCR and sequencing

DNA was extracted from fresh samples using the protocol of Wong et al. (2010). The

primers used for the matK gene region, which partially includes the 5’ flanking region

of the #rnK intron were: matK 19F (Gravendeel et al. 2001), and matK 2R (Steele &

Vilgalys 1994).

Polymerase Chain Reaction (PCR) amplifications were performed ina Biometra

Tgradient thermal cycler. PCR conditions included an initial 2-min denaturation at

95°C, 35-40 cycles of 1 min at 95°C (denaturation), | min at 50—60°C (annealing),

and 2 min at 72°C (extension), followed by a final 10-min extension at 72°C. PCR

products were purified using a PureLink™ PCR Purification Kit (Invitrogen Corp.).

The purified products were then sent to a commercial company for sequencing by

including two internal primers: 390F and 1326R (Cuénoud et al. 2002).

Data analyses

Sequences were assembled and manually aligned using BioEdit Sequence Alignment

Editor v7.0.5 (Hall 1999). Parsimony analysis was performed with PAUP* 4.0b10

(Swofford 2000) using PaupUp graphical interface (Calendini & Martin 2005) with

214 Gard. Bull. Singapore 64(1) 2012

Table 1. GenBank accessions, vouchers and locality information for species of the

Schismatoglottis Nervosa Complex (Araceae: Schismatoglottideae) in the present study.

GenBank Voucher

Species Bees fe Location Geology

Schismatoglottis ARI74 Ga aL aNer ane in havent

ane 1N570740 arawak, Serian, Gunung Ampungan _ limestone

ge SEED JN570742 AR944 Sarawak, Bau, Gunung Bidi limestone

nervosa

Schismatoglottis INS70748 AR1087 Sarawak, Kapit, Taman Rekreasi eaten

tessellata Sebabai

Se NUE JN570739 ARI408 Sarawak, Kapit, Belaga shales

adoceta

SEPSIS JNS70746 ARIS79 Sarawak, Sarikei, Ulu Sarikei shales

ulusarikeiensis

ee za fas tel JN570751 ARI686 Sarawak, Serian, Mongkos shales

Schismatogiottis 1NS70741 AR1864 Sarawak, Matang, Kubah National a rastene

matangensis Park

aie Los JN570747 ARI1877 — Sarawak, Miri, Niah National Park limestone

sorimatogions JN570744. AR1930.— Sarawak, Miri, Mulu limestone

sp. nov. aff nervosa

pea as JN570743 AR2062 ~~ Sarawak, Lundu, Gunung Gading granite

SECS RTS JN570749 = AR2078 Sarawak, Bintulu, Bukit Sarang limestone

sp. nov. aff nervosa

see eee JN570750 AR2143 Sarawak, Lundu, Sempadi sandstone

UIC

Sehismatesiots JNS70745. AR2482 Sabah, Tawau basalt

sp. nov. aff nervosa

Selisinatos Ons GQ220910 AR2677 Peninsular Malaysia, Perak granite

brevicuspis

all characters treated as “unord” and with equal weight. Gaps were treated as missing

values. Random addition sequence was used as starting point and 10000 replicates

retained. Branch-swapping was undertaken using the tree-bisection-reconnection

(TBR) algorithm. No more than 3 trees of score (length) >/= to 1 were saved for each

replicate. Steepest descent option was not in effect and branches were collapsed if

maximum branch length was zero. The “MulTrees” option was in effect and topological

constraints not enforced. 10000 bootstrap replicates were completed. The consistency

index (CI), homoplasy index (HI), retention index (RI) and rescaled consistency index

(RCI) were calculated with one of the most-parsimonious trees (MPTs).

Results

Unaligned sequences of the matK region of the ingroup taxa ranged from 1448 to

1740 bp. The final length for the aligned sequences comprised 1436 characters which

Schismatoglottis Nervosa Complex 215

included two additions in S. simonii AR1686 (from 626 to 631 bp) and A. acuminatissima

(from 1505 to 1510bp). To align the sequences with the outgroup, 11 gaps were added.

3 variable sites were found from 200bp to 1400bp. From the 1436 bps, 1413 (98%)

characters were constant, 14 (1%) of them were parsimony-uninformative and 9 (1%)

of them were parsimony-informative characters.

One of 28 most parsimonious trees was selected (Fig. 1). There are two

apparent clusters within the Nervosa Complex with weakly supported bootstrap

values: S. simonii AR1686, S. adoceta AR1408, S. tessellata AR1087, S. sp. nov. aff.

nervosa AR2078, S. brevicuspis AR2677, S. ulusarikeiensis AR1579 and S. elegans

AR1877 (bootstrap 43%); and S. nervosa AR944, S. simonii AR174, S. matangensis

AR1864, S. liniae AR2062, S. turbata AR2143, S. sp. nov. aff. nervosa AR1930

and AR2482 (bootstrap: 50%). Within these, however, Schismatoglottis adoceta,

S. brevicuspis, S. tessellata and S. sp. nov. aff. nervosa AR2078 form a group with

strong bootstrap support (100%), and this is weakly grouped (bootstrap: 36%) with

S. simonii AR1686. Schismatoglottis ulusarikeiensis and S. elegans are sister to this

clade (bootstrap: 64%). Schismatoglottis nervosa, S. simonii AR174, S. matangensis,

S. liniae and S. turbata form a strongly supported clade (bootstrap: 100%). The S. sp.

nov. aff. nervosa AR1930 and S. sp. nov. aff. nervosa AR2482 clade 1s also strongly

supported (bootstrap: 100%).

: > Apoballis acuminatissima AR1197 . Outgroup

Schismatoglottis simonii AR1686 fice Line

clade

43 Schismatoglottis sp. nov. aff. nervosa AR2078 | ctaaoe

Lupar Line

clade

Schismatoglottis elegans AR1877

Schismatoglottis nervosa AR944

Lower

Lupar Line

clade

Schismatoglottis sp. nov. aff. nervosa AR1930

Fig. 1. One of 28 most parsimonious trees for the Schismatoglottis Nervosa complex based on

the matK region. Tree length 26. Consistency index (CI) 0.885. Retention index (RI) 0.870.

Rescaled consistency index (RC) 0.769. Homoplasy index (HI) 0.115. Although some clusters

are weakly supported, potential clade relationships that may correspond to either side of the

Lupar Line are indicated as hypotheses for continuing work.

216 Gard. Bull. Singapore 64(1) 2012

Discussion

The Nervosa Complex comprises two general clusters: (((Rejang-Peninsular

Malaysia), West Borneo) Rejang-NE Borneo) and (W Borneo, NE Sarawak-NE

Borneo). Apparently, the cluster radiations are separated to below and above the Lupar

Divide. The upper Lupar Divide clade comprises S. adoceta and S. tessellata (both

Kapit, on shales), S. sp. nov. aff. nervosa AR2078 (Bintulu), and S. elegans in Miri.

The Lower Lupar Line clade comprises Schismatoglottis simonii AR1686, S. nervosa,

S. simonii AR174, S. matangensis, S. liniae, and S. turbata. The Lupar Line was an

active convergent plate margin between the Late Cretaceous (65—99mya) to Palaeocene

(54.8-65mya), with the margins extending through Engkilili and Lubok Antu (Tan

1979), approximately in line with the saddle that currently separates the Kapuas Hilir

and Kapuas Hulu mountains. Hutchison (1996) stated that the sediments of the Lupar

Line are the result of fluvial system deposition from the interior of a proto-Sundaland.

The Lupar Formation comprises turbidites (deposits comprising rhythmic alternations

of fine-grained, graded sandstones and shale beds) and igneous rocks (Honza et al.

2000). Active subduction of the oceanic crust occurred during the Cretaceous (65—

144 mya) or early Tertiary (33.7—65 mya). The Rejang-Peninsular Malaysia clade (S.

adceta, S. tessellata and S. sp. nov. aff. nervosa AR2078, S. brevicuspis), 1s estimated

to have evolved from the last common ancestor during the Early Tertiary (33.7—65

mya). Schismatoglottis adoceta, S. tessellata and S. sp. nov. aff. nervosa AR2078 are

in the Belaga and Lupar formations in Sarawak and Embaluh Group and Selangkai

Formations in Kalimantan, circumscribed in the South by the Lupar Line ophiolite

and in the North by the Bukit Mersing Line (Hutchison 1989), The Rajang Group

comprises turbidite sedimentation dating from the Late Cretaceous (65-99 mya) to

Late Eocene (33.7-41.3mya), formed by accretion at a subduction trench (Honza et

al. 2000). Bedding dips are generally southward, but become younger northward.

Our preliminary results also suggest that the Belaga might act as geographical barrier

for the distribution of S. simonii, S. tessellata, S. sp. nov. aff. nervosa AR2078, and

S. adoceta.. Schismatoglottis adoceta and S. tessellata are found on shales, whereas

S. sp. nov. aff. nervosa AR2078 is locally restricted to limestones. The inclusion of

S. brevicuspis (granite, Perak, Peninsular Malaysia) may be explicable as a relict of

the former Riau Pocket flora (Corner 1960), running from present-day Terengganu

to north-east Borneo, and might in former times have provided a corridor of everwet

“stepping stones”. Indeed, morphologically, S. simonii is morphologically proximate

to S. sp. nov. aff. nervosa AR2078.

Schismatoglottis ulusarikeiensis is grouped with S. elegans to form the

Rejang-NE Sarawak cluster/clade. Schismatoglottis ulusarikeiensis occurs on shales

at Ulu Sarikei, while S. e/egans is restricted to the Niah limestones at Gunung Subis,

which are composed of limestone from the Tangap Formation (Hazebroek & Abang

Kashim 2000). The southern part of the Niah system consists of sandstone from the

Nyalau Formation. The Subis Limestone was formed by reefs, coralline algae and tiny

shellfish of about 23 mya (Hazebroek & Abang Kashim 2000).

Schismatoglottis Nervosa Complex DAT |

The West Borneo clade comprises S. nervosa, S. simonii, S. matangensis, S.

liniae and S. turbata, and is considered be perhaps Middle Eocene (33.7—54.8 mya)

in origin. The sediments on which two of the species (S. matangensis and S. turbata)

occur are of Eocene origin (Moss & Wilson 1998). However, S. nervosa is restricted

to the Bau Limestone, which are Upper Jurassic (180-206mya) (Wolfenden 1965),

although its origin is dependent on when the Bau limestones emerged. Schismatoglottis

matangensis, S. simonii AR174 and S. turbata are restricted to sandstones but on

separate localities and differing ecologies: S. matangensis is found on soft sandstones

under perhumid moist forest in Matang, Kuching, S. simonii AR174 on sandstone

in Gunung Ampungan, Serian, while S. turbata is restricted to micro-Bornhardts

comprised of very hard sandstone in Lundu. Schismatoglottis liniae is restricted to

granite beneath perhumid forest at Gunung Gading, Lundu.

Schismatoglottis sp. nov. aff. nervosa AR1930 is restricted to the Mulu

limestones. The Mulu Formation comprises Palaeocene (54.8-6Smya) to Eocene

(33.7-54.8mya) miogeoclinal (non-volcanic) formations of limestone, sandstone,

shale and slate (Hutchison 1989), suggesting the as-yet undescribed species might

have evolved during the Eocene. Schismatoglottis sp. nov. aff. nervosa AR2482, is

restricted to basalt outcrops at Bukit Quoin, Tawau (E Sabah), predominantly the result

of young volcanism: the Tawau-Gunung Wullersdorf area is predominatly composed

of basaltic-andesite which resembles the Pliocene (1.8—5.3mya) eruptions of the

Mostyn Estate of Kunak (Hutchison 2005). Bukit Quoin is further considered a young

volcanic edifice as the volcanic rocks have weathered only to a depth of 6m (Kirk

1962), suggesting that the volcanism of the Tawau area ceased during the Quaternary.

Although Schismatoglottis sp. nov. aff. nervosa AR1930 and S. sp. nov. aff. nervosa

AR2482 are sister taxa; S. sp. nov. aff. nervosa AR2482 occurs on the young basalts,

suggesting an adaptation that may have occurred recently (probably during Pliocene

or Pleistocene).

Conclusion

The Schismatoglottis Nervosa complex diversified in Borneo with one widespread

species in West Malaysia to Sumatera. The Bornean local endemism of the complex

is unique with each species being adapted to their geological localities. This seems to

be correlated with speciation radiations and numerous vicariance events. Analyses of

matK data resolves the Schismatoglottis Nervosa complex into two apparent clusters,

one above and one below the Lupar Divide: a Rejang-Peninsular Malaysia clade and

a W Borneo clade, with the exception of two species from NE Sarawak-NE Borneo

that grouped with species from W Borneo. Recent and rapid radiation of the Nervosa

Complex in Borneo is perhaps owing to vicariance events from spatial and temporal

isolations, combined with geological and ecological factors. On-going analyses

involving further gene regions and mapping of morphological and ecological data are

being undertaken to better resolve incongruences.

218 Gard. Bull. Singapore 64(1) 2012

ACKNOWLEDGEMENTS. The first author is grateful to the award of a bursary by the

Singapore Botanic Gardens to enable her to travel to present her paper at the 8th Flora

Malesiana Symposium, August 2010. The paper is part of the first author’s on-going MSc

study on the Araceae of Borneo and is partially funded by the Ministry of Higher Education,

Malaysia by fundamental research grant scheme vote: FRGS/01(12)/709/2009(25). Research

and fieldwork in Borneo is, most recently, under Sarawak Forestry Department Research Permit

No. NPW.907.4.4(V)-77 & Park Permit No. 34/2009. The continuing collaboration and support

of the Forest Department Sarawak are gratefully acknowledged.

References

Ashton, P.S. (1972) The Quaternary geomorphological history of western Malesia and lowland

forest phytogeography. In: Ashton, P.S. & Ashton, M. (eds) The Quaternary era in Malesia

miscellaneous ser. no. 13, pp. 35—49. Hull: University of Hull, Department of Geography.

Ashton, P.S. (2005) Lambir s Forest: The World's Most Diverse Known Tree Assemblage? In:

Roubik, D.W., Sakai, S. & Abang A. Hamid Karim (eds) Pollination Ecology and the

Rain Forest: Sarawak Studies, pp. 191-216. New York: Springer.

Calendini, F. & Martin, J.-F. (2005) PaupUP v1.0.3.1 A free graphical frontend for Paup* DOS

software.

Corner, E.J.H. (1960) The Malayan flora. In: R.D. Purchon (ed) Proceedings of the Centenary

and Bicentenary Congress of Biology, pp. 21-24. Singapore: University of Malaya Press.

Cuénoud, P., Savolainen, V., Chatrou, L.W., Powell, M., Grayer, R.J. & Chase, M.W. (2002)

Molecular phylogenetics of Caryophyllales based on nuclear 18S rDNA and plastid rbcL,

atpB, and matK DNA sequences. Amer. J. Bot. 89: 132-144.

Gravendeel, B., Chase, M.W., Vogel, E.F., Roos, M.C., Mes, T.H.M. & Bachmann, K. (2001)

Molecular phylogeny of Coelogyne (Epidendroideae; Orchidaceae) based on plastid

RFLPS, matkK, and nuclear ribosomal ITS sequences: Evidence for polyphyly. Amer. J.

Bot. 88: 1915-1927.

Hall, T.A. (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis

program for Windows 95/98/NT. Nucleic Acids Symp. Ser. 41: 95-98.

Hall, R. (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific:

computer based reconstructions, model and animation. J. Asian Earth Sci. 20: 353-431.

Hall, R., Hattum, M.W.A. van & Spakman, W. (2008) Impact of India-Asia collision on SE

Asia: The record in Borneo. Tectonophysics 451: 366-389.

Hay, A. & Yuzammi (2000) Schismatoglottideae (Araceae) in Malesia I — Schismatoglottis.

Telopea 9(1): 1-177.

Hazebroek, H.P. & Abang Kashim bin Abang Moshidi (2000) National Parks of Sarawak. Kota

Kinabalu: Natural History Publications (Borneo).

Honza, E., John, J. & Banda, R.M. (2000) An imbrications model for the Rajang Accretionary

Complex in Sarawak, Borneo. J. Asian Earth Sci.18: 751-759.

Hutchison, C.S. (1989) Geological Evolution of South-East Asia. New York: Oxford University

Press.

Hutchison, C.S. (1996) The ‘Rajang accretionary prism’ and ‘Lupar Line’ problem of Borneo.

In: Hall, R. & Blundell, D. (eds) Tectonic Evolution of Southeast Asia, Geological Society

Special Publication 106, pp. 247-261. London: Geological Society.

Hutchison, C.S. (2005) Geology of north-west Borneo: Sarawak, Brunei and Sabah. The

Netherlands: Elsevier.

Schismatoglottis Nervosa Complex 219

Kirk, H.J.C. (1962) The Geology and Mineral Resources of the Semporna Peninsula North

Borneo. British Borneo Geological Survey Memoir 14.

Mackinnon, K., Hatta, G., Halim, H. & Mangalik, A. (1996) The ecology of Kalimantan, the

ecology of Indonesia series II]. Singapore: Periplus Editions (HK) Ltd.

Morley, R.J. (2000) Origin and evolution of tropical rain forests. UK: John Wiley & Sons.

Moss, S.J. & Wilson, M.E.J. (1998) Biogeographic implications of the tertiary paleogeographic

evolution of Sulawesi and Borneo. In: Hall, R. & Holloway, J.D. (eds) Biogeography

and Geological Evolution of Southeast Asia, pp. 133-163. The Netherlands: Backbuys

Publishers.

Nagamasu, H. & Sakai, S. (1999) Orchidantha inouei (Lowiaceae), a new species from Borneo.

Nord. J. Bot. 19: 149-152.

Roos, M.C., Kessler, P.J.A., Gradstein, S.R. & Baas, P. (2004) Species diversity and endemism

of five major Malesian islands: diversity-area relationships. J. Biogeogr. 31: 1893-1908.

Slik, J.W.F., Poulsen, A.D., Ashton, P.S., Cannon, C.H., Kartawinata, K., Lanniari, I.

Nagamasu, H., Nakagawa, M., Nieuwstadt, M.G.L. van, Payne, J., Purwaningsih, Saridan,

A., Sidiyasa, K., Verburg, R.W., Webb, C.O. & Wilkie, P. (2003) A floristic analysis of the

lowland dipterocarp forests of Borneo. J. Biogeogr. 30: 1517-1531.

Steele K.P. & Vilgalys, R. (1994) Phylogenetic analysis of Polemoniaceae using nucleotide

sequences of plastid gene matK. Syst. Bot. 19: 126-142.

Swofford, D.L. (2000) PAUP*; Phylogenetic Analysis Using Parsimony (*and other methods),

version 4. Massachusetts, Sunderland: Sinauer Associates.

Takano, A & Nagamasu, H. (2007) Myxochlamys (Zingiberaceae), a new genus from Borneo.

Acta Phytotax. Geobot. 58(1): 19-32.

Tan, D.N.K. (1979) Lupar Valley, West Sarawak, Malaysia. Geological Survey of Malaysia

Report 13.

Theilade, I. & Mood, J. (1997) Five new species of Zingiber (Zingiberaceae) from Borneo.

Nord. J. Bot. 17: 337-347.

Theilade, I. & Mood, J. (1999) Six new species of Zingiber (Zingiberaceae) from Borneo.

Nord. J. Bot. 19: 513-524.

Welzen, P.C. van, Slik, J.W.F. & Alahuhta, J. (2005) Plant distribution patterns and plate

tectonics in Malesia. Biol. Skr. 55: 199-217.

Wolfenden, E.B. (1965) Bau mining district, West Sarawak, Malaysia (Part 1: Bau). Geological

survey Borneo region, Malaysia Bulletin 7:\—147.

Wong, K.M. (1998) Patterns of plant endemism and rarity in Borneo and the Malay Peninsula.

In: Peng, C.-I. & Lowry II, P.P. (eds) Rare, threatened and endangered floras of Asia

and the Pacific Rim, Monograph Series No. 16, pp. 139-169. Tatwan: Academia Sinica,

Institute of Botany.

Wong, S.Y. (2010) Studies on Schismatoglottideae (Araceae) of Borneo XIII: A revision of the

Schismatoglottis nervosa complex. Gard. Bull. Singapore 62(1):177—209.

Wong S.Y., Boyce, P.C., Ahmad Sofiman, O. & Leaw, C.P. (2010) Molecular phylogeny of

tribe Schismatoglottideae based on two plastid markers and recognition of a new tribe,

Philonotieae, from the neotropics. Taxon 59(1): 117-124.

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Gardens’ Bulletin Singapore 64(1): 221-256. 2012 |

The plant taxa of H.N. Ridley, 4.

The primitive angiosperms (Austrobaileyales, Canellales,

Chloranthales, Laurales, Magnoliales, Nymphaeales and

Piperales)

I.M. Turner

Fairfield, Pett Level Road, Winchelsea Beach,

East Sussex TN36 4ND, U.K.

turner! 87@pbtinternet.com

ABSTRACT. The names of plant taxa authored by H.N. Ridley from the orders of

primitive angiosperms are enumerated. A total of 157 taxa across 11 families (Annonaceae,

Aristolochiaceae, Chloranthaceae, Illiciaceae, Lauraceae, Magnoliaceae, Monimiaceae,

Nymphaeaceae, Piperaceae, Trimeniaceae and Winteraceae) and seven orders (Austrobaileyales,

Canellales, Chloranthales, Laurales, Magnoliales, Nymphaeales and Piperales) are listed with

synonyms and accepted names. The types are listed for those taxa that Ridley described.

Lectotypes are designated for 37 taxa. Melodorum breviflorum Ridl. (Annonaceae) is transferred

to Fissistigma, and two Ridley species in Piperaceae that are later homonyms are provided with

new names: Peperomia kerinciensis 1.M.Turner for Peperomia villosa Ridl. (1917, nom. illegit.

non P. villosa C.DC. (1866)) and Piper angsiense 1.M.Turner for Piper venosum Ridl. (1925,

nom. illegit. non P. venosum (Miq.) C.DC. (1869)).

Keywords. Austrobaileyales, Canellales, Chloranthales, Laurales, Magnoliales, Nymphaeales,

Piperales, primitive angiosperms, Ridley

Introduction

This paper continues an intermittent series on the plant taxa named by Henry Nicholas

Ridley (1855-1956). The three parts published to date (Turner & Chin 1998a, b; Turner

2000), dealt with the pteridophytes, gymnosperms and Zingiberales, respectively. The

focus shifts to the primitive angiosperms in the current paper.

Ridley described many plant species. Among the primitive angiosperm orders

there are numerous examples which are enumerated in this paper. Most of these taxa

were described in the course of Ridley’s many papers documenting the diversity of

South-East Asian plants often in relation to the research expeditions undertaken by

Ridley and others.

The three families best represented from among the primitive angiosperms are

Annonaceae, Lauraceae and Piperaceae. They are here ordered in terms of declining

attention from taxonomists since Ridley’s day. The Annonaceae are a diverse family in

tropical Asia represented by many genera. There has been and remains a considerable

activity in Annonaceae systematics. The Lauraceae are more poorly served despite

Gard. Bull. Singapore 64(1) 2012

the efforts of the late A.J.G.H. Kostermans. The Piperaceae in Malesia have been

particularly neglected. Ridley’s account in his Flora of the Malay Peninsula is still the

only treatment available for the area, though it relies very heavily on the work of C. de

Candolle, particularly his paper describing many new species for the Malay Peninsula

(Candolle 1912). Ina reversal of the general position, the Piperaceae of eastern Malesia

(New Guinea) have received more recent attention, largely thanks to Drs. W.L. Chew

and R.O. Gardner, than the far west of the region. One surprise from the enumeration

is that Ridley seems never to have named any members of the Myristicaceae, despite

the nutmegs being both diverse and common in the lowland forests of South-East Asia.

Warburg’s monographic work on the family (Warburg 1897) may be one reason for

this. Another possible explanation 1s that the Singapore Botanic Gardens did not have

professional tree climbers, so canopy trees were probably little collected by Ridley—

there was a vast number of novelties to be found in the lower layers of the forest.

An enumeration of Ridley’s primitive angiosperm taxa

The taxa Ridley described as new are listed against Arabic numbers, and purely

nomenclatural novelties are itemised with Roman numerals. Ridley’s combination

starts each entry and is given in bold. Synonyms, including basionym where relevant,

are given. Where possible a currently accepted name is included in bold small

capitals, either included in the list of synonyms if homotypic to Ridley’s taxon or

below, preceded by ‘=’, if heterotypic. In some cases the identity of Ridley’s taxon

is uncertain (e.g., sheets are determined but the information seems not to have been

published), in which instances a question mark is employed, or unknown in which case

no accepted name 1s listed. The place of publication of all names is given and reference

to important revisions and monographs are included.

For each of the taxa Ridley described, the types are listed including as many

syntypes as have been located, and the herbaria in which they are found. The author

has seen most of the specimens, but some records come from published sources,

information from correspondents or on-line databases. There are doubtless duplicates

in other herbaria and there remain a few instances where no type material has yet been

found.

A note on typification

A high proportion of Ridley’s taxa requires lectotypification because he rarely

designated types from among the various specimens he cited when publishing new

taxa. Specimen citation by Ridley can frequently be careless—he often omitted

collection numbers, rarely noted collection dates, and generally failed to state in which

herbaria material was deposited. He was also inconsistent in annotating the specimens

he saw. Typification is therefore often problematic. Ridley’s career can be divided into

two main periods—the Singapore years (1888-1912) and the retirement to Kew (1913

onwards). This information is important in making decisions relating to typification of

Ridley taxa as it helps identify what is likely to be original material, but experience

oS)

Ridley’s taxa among primitive angiosperms

has shown that many types one might expect, from the date of publication, to be in

Singapore are found at Kew. In order to reduce the number of repetitive explanatory

notes in the list, if choice of lectotype was simply based on selecting the specimen of

highest quality from among candidate syntypes, no note is included.

AUSTROBAILEYALES

Illiciaceae

L. Illicitum cambodianum Hance var. tenuifolium Ridl., Fl. Malay

Penins. 1 (1921) 19, as ‘tenuifolia’. ILLICIUM TENUIFOLIUM (RidI.)

A.C. Sm., Sargentia 7 (1947) 68; Saunders, Fl. Males. 13 (1997)

182; Phoon, Fl. Pen. Malaysia, ser. II, 1 (2010) 142. SYNTYPES:

Peninsular Malaysia, Pahang, Telom Cascade, H.N. Ridley 13534,

November 1908 (lectotype, designated by Smith (1947), K (barcode

no. K000442171); isolectotype, BM); Gunong Tahan; Selangor,

Menuang Gasing, C.B. Kloss s.n., February 1912 (BM, K). Perak,

Thaiping Hills, H.H. Kunstler. Kedah, Kedah Peak, L.M. Bell & Md

Haniff s.n., March 1911 (K[x2]).

2 Illicium cambodianum var. crassifolium Ridl., Fl. Malay Penins. | (1921)

19, as ‘crassifolia’. SYNTYPES: Peninsular Malaysia, Selangor,

Gunong Menkuang, 5000 feet, H.C. Robinson s.n., 18 January

1913 (lectotype, designated here, K (barcode no. K000442171);

isolectotype, BM (barcode no. BM000017468)); Bukit Kutu (flowers

fascicled on stems), H.N. Ridley 7218, May 1895 (K); Sempang

Mines, H.N. Ridley 15691, April 1911 (K); Semangkok Pass.

Perak, Gunong Kerbau, Md Haniff 4017, May 1909 (K); Gunong

Batu Putih, L. Wray 251, s. dat. (K). Malacca, Mt Ophir, Pahang,

K’luang Terbang, WD. Barnes 10876, 1900 (K); Gunong Berembun

near Telom, H.N. Ridley 13563, November 1908, (BM, K); Gunong

Tahan, H.N. Ridley 16268, July 1911 (K); H.N. Ridley 16009, July

1911 (K).

= ILLICIUM RIDLEYANUM A.C. Sm., Sargentia 7 (1947) 66; Saunders, FI.

Males. 13 (1997) 181; Phoon, Fl. Pen. Malaysia, ser. II, 1 (2010) 140.

The lectotype selected for Ridley’s var. crassifolium is the holotype of

Illicium ridleyanum A.C. Sm., which seems appropriate given Smith’s

choice of epithet for the species. Making these taxa homotypic

synonyms may simplify future rearrangements in //licium.

Gard. Bull. Singapore 64(1) 2012

Trimeniaceae

ee TRIMENIA PAPUANA RidI., Trans. Linn. Soc., Bot. 9 (1916) 144; Rodenburg,

Blumea 19 (1971) 9; Philipson, Fl. Males. 10 (1986) 331. TYPE:

New Guinea, Mt Carstensz, Camp VIb, 3900 ft, C.B. Kloss s.n., 17

February 1913 (holotype, BM (barcode: BM000950880)).

CANELLALES

Winteraceae

4. Drimys buxifolia Ridl., Trans. Linn. Soc., Bot. 9 (1916) 12. Tasmannia

buxifolia (Ridl.) A.C. Sm., Taxon 18 (1969) 288. TYPE: New Guinea,

Mt Carstensz, Camp XIII to XVI (13-11 on sheet), 8300 to 10500 ft,

C.B. Kloss s.n., 1912-1913 (holotype, BM (Barcode: BM00574841);

isotype, K).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns et al., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

5 Drimys densifolia Ridl., Trans. Linn. Soc., Bot. 9 (1916) 12. Tasmannia

densifolia(Ridl.) A.C. Sm., Taxon 18 (1969) 289. TYPE: New Guinea,

Mt Carstensz, Camp XII to XIII, 10500 to 11000 ft, C.B. Kloss s.n.,

29 January 1913 (holotype, BM (barcode no. BM 000574843)).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns et a/., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

6. Drimys elongata Ridl., Hook. Icon. Pl. 31 (June 1916) t. 3051; Trans.

Linn. Soc., Bot. 9 (August 1916) 12. Zasmannia elongata (Ridl.) A.C.

Sm., Taxon 18 (1969) 288. SYNTYPES: New Guinea, Mt Carstensz,

Camp III to VHI, 2500 to 4900 ft, CB. Kloss s.n., December 1912 (BM

(barcode no. BM00574844)); Camps VIb and VIc, 3900 to 5500 ft,

C.B. Kloss s.n., 17 February 1912 (BM (barcode no. BM00574845));

Camp VIb, s.dat., C.B. Kloss s.n. (lectotype, designated by Vink

(1970), K (barcode no. K0006816245)).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns et a/., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

Ridley’s taxa among primitive angiosperms

11.

bho

Drimys grandiflora Ridl., Trans. Linn. Soc., Bot. 9 (1916) 11. Tasmannia

grandiflora (Ridl.) A.C. Sm., Taxon 18 (1969) 288. TYPE: New

Guinea, Mt Carstensz, Camp XIII, 10500 ft, C.B. Kloss s.n., 29

January 1913 (holotype, BM (barcode no. BM000574847): isotype,

K).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns et a/., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

Drimys parviflora Ridl., Trans. Linn. Soc., Bot. 9 (1916) 12. Bubbia

parviflora (Ridl.) Burtt, Hook. Icon. Pl. 34 (1936) t. 3315, p. 3.

Tasmannia parviflora (Ridl.) A.C. Sm., Taxon 18 (1969) 288.

TYPE: New Guinea, Mt Carstensz, Camps XI to XII, 8300 to 11900

ft, C.B. Kloss s.n., 28 January 1913 (holotype, BM (barcode no.

BM000574842); isotype, K).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns et a/., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

Drimys rosea Ridl., Trans. Linn. Soc., Bot. 9 (1916) 11. Zasmannia rosea

(Ridl.) A.C. Sm., Taxon 18 (1969) 288. TYPE: New Guinea, Mt

Carstensz, Camp XII to XIII, 10,000 ft, C.B. Kloss s.n., 29 January

1913 (holotype BM (barcode no. BM 00057446); isotype, K).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110; Hopkins in Johns ef a/., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

Drimys umbellata Ridl., Trans. Linn. Soc., Bot. 9 (1916) 11. Bubbia

umbellata (Ridl.) Dandy, J. Bot. 72 (1934) 41. ZyGoGyNuM

UMBELLATUM (Ridl.) W. Vink, Blumea 31 (1985) 54. TYPE: New

Guinea, Mt Carstensz, Camp VIb, 3900 ft, C.B. Kloss s.n., January

1913 (holotype, BM (barcode no. BM000574861)).

Drimys vaccinioides Ridl., Trans. Linn. Soc., Bot. 9 (1916) 13. Tasmannia

vacciniodes (Ridl.) A.C. Sm., Taxon 18 (1969) 288. TYPE: New

Guinea, Mt Carstensz, Camp XIII, 10500 ft, C.B. Kloss s.n. (holotype,

BM, ? lost).

= TASMANNIA PIPERATA (Hook.f.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2

(1858) 110. Hopkins in Johns et al., Guide Alp. & Subalp. Fl. Mt.

Jaya (2006) 496.

I have not been able to find the type material of Drimys vaccinioides. Vink

(1970) also reported it missing.

Gard. Bull. Singapore 64(1) 2012

CHLORANTHALES

Chloranthaceae

Chloranthus brachystachys var. melanocarpa Ridl., J. Straits Branch

12. Roy. Asiat. Soc. 87 (1923) 89. SARCANDRA GLABRA (Thunb.)

Nakai var. MELANOCARPA (Ridl.) Verdc., Kew Bull. 40 (1985) 217;

Verdcourt, Fl. Males. 10 (1986) 136. TYPE: Sumatra, Berastagi, H.N.

Ridley s.n., February 1921 (holotype, K (barcode no. K000802684);

isotype, SING).

LAURALES

Lauraceae

(3. ACTINODAPHNE CONCINNA Ridl., J. Fed. Malay States Mus. 5 (1914) 44.

TYPE: Peninsular Malaysia, Selangor, Gunong Menkuang, 5000

feet, H.C. Robinson s.n., 18 January 1913 (holotype, K (barcode no.

K000793048); isotypes, BM, SING (barcode no. SING 0051764)).

14. Actinodaphne gelonioides Ridl., Fl. Malay Penins. 3 (1924) 111. TYPE:

Peninsular Malaysia, Perak, Gunong Kerbau, H.C. Robinson s.n.,

June 1913 (holotype, K (barcode no. K000793057); isotype, BM).

= ACTINODAPHNE OLEIFOLIA Gamble, Bull. Misc. Inform. Kew 1910 (1910)

313; Kochummen, Tree FI. Malaya 4 (1989) 106.

15: Actinodaphne oleifolia Gamble var. glabra Ridl., Fl. Malay Penins. 3

(1924) 111. SYNTYPES: Peninsular Malaysia, Pahang, Gunung

Tahan, H.N. Ridley 16169, July 1911 (lectotype, designated here, K

(barcode no. K000793056)); H.N. Ridley 16109, July 1911 (K).

= ACTINODAPHNE OLEIFOLIA Gamble, Bull. Misc. Inform. Kew 1910 (1910)

313; Kochummen, Tree Fl. Malaya 4 (1989) 106.

I do not believe the collection numbers of the syntypes are given in error.

Both sheets have original SING labels which have different initial

determinations.

16. BEILSCHMIEDIA BREVIPES Ridl., Fl. Malay Penins. 3 (1924) 86; Kochummen,

Tree Fl. Malaya 4 (1989) 118. TYPE: Peninsular Malaysia, Pahang,

Ulu Rompin, F-W. Foxworthy Forest Department FMS 3181, 21

April 1919 (holotype, K (barcode no. K000768680)).

Ridley’s taxa among primitive angiosperms 27

(vs

20.

Beilschmiedia longipedicellata Ridl.. Bull. Misc. Inform. Kew 1926

(1926) 475; DEHAASIA LONGIPEDICELLATA (Ridl.) Kosterm., New

& Crit. Mal. Pl. IV. (For. Serv. Indones., Div. Plann.) (1955) 37:

Kostermans, Bot. Jahrb. Syst. 93 (1973) 462. TYPE: Peninsular

Malaysia, Selangor, Semeniyih, H.L. Hume 8432, 13 August 1921

(holotype, SING (barcode no. SING 0055156): isotype, BO).

Ridley did not cite the Hume collection number or state in which herbaria

the type material was deposited. Given the data and place of the

original publication, Kew would seem the likely institution, but I

have failed to locate any relevant specimens. Hume worked for the

FMS Museums and his collections were deposited in the Museum’s

herbarium from where they were indefinitely loaned to the Singapore

Herbarium (van Steenis-Kruseman 1950). In SING there is a sheet

of Hume 8432 which is labelled Beilschmiedia longipedicellata in

Ridley’s hand. I assume this to be the holotype which Ridley must

either have seen during a visit to Kuala Lumpur or was sent to

Kew for him to name and later returned to Kuala Lumpur and then

transferred to Singapore.

Beilschmiedia sumatrensis Ridl., J. Malayan Branch Roy. Asiat. Soc.

1 (1923) 89, as “Beilschimedia’. TYPE: Sumatra, Berastagi woods,

HN. Ridley s.n., February 1921 (lectotype, designated here, K

(barcode no. K000778438)).

There is another sheet at K (barcode no. K000778439) which may be a

duplicate, or another gathering by Ridley from the same locality.

The lectotype has a collection label in Ridley’s hand; that of the

‘duplicate’ seems not to be written by him.

BEILSCHMIEDIA TONKINENSIS Ridl., J. Straits Branch Roy. Asiat. Soc. 82

(1920) 190, as ‘Beilschimedia’; Kochummen, Tree Fl. Malaya 3

(1989) 122. TYPE: Thailand, Satun, Setul, H.N. Ridley 15465.

Kostermans (1964) and Kochummen (1989) treat this as anew combination

based on B. glomerata var. tonkinensis Lecomte. However, Ridley

refers to B. tonkinensis as ‘sp. nov.’ in the publication. I have not

managed to trace any type material.

Cinnamomun iners var. angustifolium Ridl., Fl. Malay Penins. 3 (1924)

93. TYPE: Peninsular Malaysia, Perak, Temengoh, Kelantan River,

HN. Ridley 14516, July 1909 (lectotype, designated by Soh (2011),

SING (barcode no. SING 0055564): isolectotype, K).

= CINNAMOMUM INERS (Nees & T.Nees) Reinw. ex Blume, Bijdr. (1826)

570; Kochummen, Tree FI. Malaya 4 (1989) 126.

Gard. Bull. Singapore 64(1) 2012

CINNAMOMUM KUNSTLERI Rid1., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

191; Kochummen, Tree Fl. Malaya 3 (1989) 127. TYPE: Peninsular

Malaysia, Perak, Larut, Dr Kings Collector [H.H. Kunstler]

5568, Febuary 1884 (lectotype, designated here, K(barcode no.

K000778640); isolectotypes, BM (barcode no. 000950947), K[x2],

SING (barcode no. 0051701), L (barcode no. L0035840)).

Of the three sheets of King’s Collector 5568 at K, one came from the

personal herbarium of J.S. Gamble and was only donated to Kew in

1925 so is unlikely to be original material of Cinnamomum kunstleri

Ridl. The other two sheets are annotated by Ridley with the species

name. All three sheets have red ‘type’ tickets with one sheet indicated

as holotype. I designate this sheet as the lectotype.

Cinnamomum parvifolium Ridl., J. Fed. Malay States Mus. 6 (1915) 54,

non C. parvifolium Lecomte (1913); CINNAMOMUM MICROPHYLLUM

Ridl., Fl. Malay Penins. 3 (1924) 92, nom. nov.; Kochummen, Tree

Fl. Malaya 4 (1989) 127. TYPE: Peninsular Malaysia, Perak, Gunong

Kerbau, H.C. Robinson s.n., 20 March 1913 (holotype, K (barcode

no. K000778644)).

CINNAMOMUM PUBERULUM Ridl., Fl. Malay Penins. 3 (1924) 96;

Kochummen, Tree Fl. Malaya 4 (1989) 128. SYNTYPES: Peninsular

Malaysia, Pahang, Bentong, Ulu Raub, Ahmad [F.W. Foxworthy’s

collector] 5096, 5 January 1920 (lectotype, designated here, K

(barcode no. K000778651)); Ahmad 5063, 29 December 1919 (K,

SING (barcode no. SING 0055569); Ahmad 5095, 5 January 1920

(SING (barcode no. SING 0055568)).

CINNAMOMUM SCORTECHINI Gamble var. SELANGORENSE Ridl., Fl. Malay

Penins. 3 (1924) 95; Kochummen, Tree Fl. Malaya 4 (1989) 131.

TYPE: Peninsular Malaysia, Selangor, Ulu Kali, 4. Burn-Murdoch

339 [A. Hashim leg.] (lectotype, designated by Kostermans (1970),

SING (barcode no. SING 0051711); isolectotypes, K[*2]).

Cinnamomum velutinum Ridl., J. Straits Branch Roy. Asiat. Soc. 82

(1920) 190. TYPE: Peninsular Malaysia, Pahang, Gunong Tahan,

Teku woods, H.N. Ridley 16110, 17 July 1911, (holotype, K (barcode

no. K000778660); isotype, SING (barcode no. SING 0051702).

= C. scorrecuinu' Gamble, Bull. Misc. Inform. Kew 1910 (1910) 219;

Kochummen, Tree Fl. Malaya 4 (1989) 130.

Ridley’s taxa among primitive angiosperms

26.

DS:

Jas

30.

Cryptocarya tenuifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

190. TYPE: Peninsular Malaysia, Selangor, Ulu Gombak, H.N.

Ridley s.n., 9 March 1915 (holotype, K).

= CRYPTOCARYA ZOLLINGERIANA Miq., Fl. Ind. Bat. 1(1) (1858) 924;

Kochummen, Tree Fl. Malaya 4 (1989) 137.

Dehaasia elliptica Ridl., Fl. Malay Penins. 5 (1925) 329. SYNTYPES:

Peninsular Malaysia, Selangor, Klang, Bukit Cheraka, [FW.

Foxworthy| 3462 (lectotype, designated here, K (barcode no.

K000350888)); Kuala Lumpur, Public Gardens, F}W. Foxworthy

4706, 5 July 1919 (K (barcode no. K000350887));.

= DEHAASIA POLYNEURA (Miq.) Kosterm., Bot. Jahrb. Syst. 93 (1973) 466.

Kostermans cited Foxworthy 4760 as the type of Dehaasia elliptica but

while this specimen was annotated by Ridley it was not cited in the

protologue.

DEHAASIA LANCIFOLIA Ridl., J. Linn. Soc., Bot. 38 (1908) 320; Kostermans,

Bot. Jahrb. Syst. 93 (1973) 461. SYNTYPES: Peninsular Malaysia,

Gunung Tahan 5000-6000 ft, L. Wray & H.C. Robinson 5468, 5 July

1905 (lectotype, designated here, BM; isolectotypes, K, SING[*2]);

H.C. Robinson 5525, 12 July 1905 (BM).

Endiandra crassifolia Ridl., Bull. Misc. Inform. Kew 1926 (1926) 79.

NOTHAPHOEBE CRASSIFOLIA (Ridl.) Kosterm., Reinwardtia 7 (1968)

354. TYPE: Sumatra, Mentawai Islands, Siberut, C.B. Kloss 14593,

27 September 1924 (holotype, K (barcode no. K000778906)).

Lindera caudifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

192. SYNTYPES: Peninsular Malaysia, Penang, Penang Hill, Road

to Penara Bukit, 2000 ft, C. Curtis 2447, June 1890 (lectotype,

designated here, K (barcode no. K000815584); isolectotype, BM);

2000 ft. Tiger Hill, H.N. Ridley s.n., March 1915 (K, BM).

= LINDERA SUBUMBELLIFLORA (Blume) Kosterm., J. Sci. Res., Indonesia

1 (1952) 127; Kochummen, Tree Fl. Malaya 4 (1989) 148, as

‘subumbellifera .

The Curtis collection may be numbered 2442 - the last figure is not clear.

230 Gard. Bull. Singapore 64(1) 2012

Suh Lindera cinnamomea Ridl., J. Fed. Malay States Mus. 6 (1915) 173.

NEOLITSEA CINNAMOMEA (Ridl.) Kosterm., Reinwardtia 6 (1962)

287. TYPE: Peninsular Malaysia, Pahang, Gunong Tahan, H.N.

Ridley 16124, July 1911 (holotype, K (barcode nos. K000815497,

K000815498)).

There are two sheets of Ridley 16/24 at K. One has the blue collection

label typically used in the Singapore Herbarium the other has details

written on a Herb. Hort. Bot. Reg. Kew slip. The two sheets were

received ar Kew on the same date — 9 November 1911. The sheet

with the Singapore label is annotated ‘Lindera cinnamomea Ridl.’ in

Ridley’s hand. The other sheet has ‘cinnamomea’ in Ridley’s hand

on the Kew slip. Above the collection label on each sheet someone

has written ‘bis’. My interpretation is that these two sheets represent

a single specimen — there was too much material to mount on one

sheet so the excess was mounted on a second sheet and the label

details copied on to a Kew slip. A type must consist of a single

specimen [ICBN (McNeill et al. 2006) Art. 8.1]. A specimen may be

mounted as more than one preparation as long as the parts are clearly

labelled as being part of the same specimen [ICBN Art. 8.3]. While

‘bis’ (Latin = twice), seems to have several meanings when put on

herbarium sheets, in this case I infer that it means that the specimen

was mounted on two sheets and the holotype consists therefore of

two sheets.

i). LINDERA CONCINNA Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 192;

Kochummen, Tree Fl. Malaya 4 (1989) 146. SYNTYPES: Peninsular

Malaysia, Pahang, Gunong Tahan, L. Wray & H.C. Robinson 5455, 2

July 1905 (lectotype, designated here, K (barcode no. K000815604);

isolectotype, BM (barcode no. BM000951093)); H.N. Ridley 16108,

July 1911 (K[<2]).

33: LINDERA MONTANA Ridl., J. Fed. Malay States Mus. 6 (1915) 172;

Kochummen, Tree Fl. Malaya 4 (1989) 146. SYNTYPES: Peninsular

Malaysia, Pahang, Gunong Tahan, Wray’s camp, 3300 ft, HN.

Ridley 16161, July 1911 (lectotype, designated here, K (barcode

no. K000815596) (excluding fruiting material in capsule)); Gunung

Tahan 5600 ft, H.N. Ridley 16161, July 1911 (K, fruiting material in

capsule only (barcode no. K000815597)).

Ridley 16161 consists of a herbarium sheet with an attached capsule and

two collection labels. The mounted specimen is a leafy twig with

flowers that the label indicates was collected at 3300 ft on Gunung

Tahan. The capsule contains fruits that were collected at 5600 ft.

Though Ridley included these under the same collection number

they clearly represent separate gatherings and so lectotypification is

required.

Ridley’s taxa among primitive ansiosperms

35:

36.

37.

38.

—

Lindera selangorensis Ridl.. J. Fed. Malay States Mus. 5 (1914) 44.

SYNTYPES: Peninsular Malaysia, Selangor, Gunong Menkuang

Lebah, H.C. Robinson s.n., 6 February 1913 (lectotype, designated

here. K (barcode no. K000815585): isolectotypes. BM. SING

(barcode no. SING 0046618)): Sempang Mines. H.N. Ridley 15589,

Apmil 1894. (K[<2], SING (barcode no. SING 0046596)).

= Linpera Lucipa (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 147.

The species was described in a paper on plants collected from Gunong

Mengkuang Lebah. so the Robinson specimen is chosen as lectotype.

Lindera stricta Ridl., J. Fed. Malay States Mus. 6 (1915) 172. TYPE:

Peninsular Malaysia, Pahang, Gunong Tahan, HN. Ridley 16128,

July 1911 (holotype, K (barcode no. K000815595); isotype. BM).

= LinpeRA MonTANA Ridl.. J. Fed. Malay States Mus. 6 (1915) 172:

Kochummen, Tree Fl. Malaya 4 (1989) 146.

LiTsEA ACRANTHA Ridl.. J. Fed. Malay States Mus. 10 (1920) 152:

Kochummen, Tree Fl. Malaya 4 (1989) 151. TYPE: Peninsular

Malaysia, Kedah-Perak boundary, Gunong Bintang. nat. coll. F MS.

Mus. s.n., June 1917 (holotype, K (barcode no. K000797079)).

LITSEA CINERASCENS Rid. J. Linn. Soc., Bot. 41 (1913) 296: Kochummen,

Tree Fl. Malaya 4 (1989) 153. SYNTYPES: Peninsular Malaysia.

Selangor, Gunong Menuang Gasing, C.B. Kloss. Pahang, Telom.

HN. Ridley 13781, November 1908 (K; SING (barcode no. SING

0055903)).

I have not been able to trace the Kloss specimen. As the species was

described in a paper conceming plants collected on Gunong Menuang

Gasing, it seems prudent to defer lectotypification in this case.

Litsea glabrifolia Ridl., J. Fed. Malay States Mus. 10 (1920) 152. TYPE:

Peninsular Malaysia, Pahang, Gunong Senyum, 1 H_N. Evans s.n.,

June 1917 (holotype. K: isotype. BM).

= Litsea INSIGNIS (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 142:

Kochummen, Tree Fl. Malaya 4 (1989) 159.

ios)

39)

40.

4].

Gard. Bull. Singapore 64(1) 2012

Litsea machilifolia Gamble var. montana Ridl., Fl. Malay Penins. 3

(1924) 127. TYPE: Peninsular Malaysia, Pahang, Bukit Fraser

(Sempang), /.H. Burkill & R.E. Holttum Forest Dept., F. M.S. 7784,

16-30 September 1922 (holotype, K (barcode no. K000797064)).

= LITSEA MACHILIFOLIA Gamble, Bull. Misc. Inform. Kew 1910 (1910)

320; Kochummen, Tree Fl. Malaya 4 (1989) 160.

There are further Burkill & Holttum collections from Fraser’s Hill in K

made on the same visit but collected under SF numbers. None of these

is annotated by Ridley as var. montana so I assume the annotated

sheet to be the holotype.

Litsea persella Ridl., Agric. Bull. Straits Fed. Malay States 5 (1906)

6. Cylicodaphne persella (Ridl.) Nakai, Bull. Tokyo Sci. Mus.

No. 22 (1948) 32. TYPE: Borneo, Sarawak, Kuching, H.N. Ridley

12429, September 1905 (lectotype, designated here, K (barcode no.

K000815214)).

?=Litsea garciae S. Vidal, Revis. Pl. Vasc. Filip. (1886) 228.

In the protologue Ridley wrote ‘I found it beautifully in flower in Sarawak

in September 1905, in the town of Kuching, and later Mrs Ellis and

Mr Hewitt sent me seeds and fruits of it.” I have not been able to

locate any fruit or seed material, but there is a flowering specimen as

Kew which I designate the lectotype of Litsea persella.

Litsea tenuipes Ridl., Bull. Misc. Inform. Kew 1926 (1926) 79. TYPE:

Sumatra, Mentawai Islands, Sipora, C.B. Kloss 1475], 22 October

1924 (holotype, K (barcode no. K000815203)).

?=Litsea insignis (Blume) Boerl., Handl. Fl. Ned. Ind. 3 (1900) 142.

Litsea terminalis Ridl., Fl. Malay Penins. 5 (1925) 330. TYPE: Peninsular

Malaysia, Perak, Gunong Inas, R.H. Yapp 494, 19 December 1899

(holotype, K (barcode no. K000797080)).

= LITSEA ACRANTHA Ridl., J. Fed. Malay States Mus. 10 (1920) 152;

Kochummen, Tree Fl. Malaya 4 (1989) 151.

Nothaphoebe angustifolia Ridl., J. Fed. Malay States Mus. 7 (1916)

50, as ‘Nothophoebe’. TYPE: Peninsular Malaysia, Kedah, Kedah

Peak, 3800 ft, H.C. Robinson & C.B. Kloss 5996, December 1915

(holotype, K (barcode no. K000778892); isotype, SING (barcode no.

SING 0056072)).

= PERSEA FRUTICOSA (Kurz) Kosterm., Reinwardtia 7 (1968) 355;

Kochummen, Tree Fl. Malaya 4 (1989) 173.

Ridley’s taxa among primitive angiosperms

44.

45.

233

NOTHAPHOEBE CONDENSA Ridl., J. Straits Branch Roy. Asiat. Soc. 82

(1920) 191; Kochummen, Tree Fl. Malaya 4 (1989) 170. TYPE:

Peninsular Malaysia, Pulau Langkawi, Kerap [Kesap], Md. Aniff

15473, February 1911 (holotype, K (barcode no. K000575991):

isotype, SING (barcode no. SING 0056073).

Phoebe tavoyana var. villosa Ridl., Fl. Malay Penins. 3 (1924) 105.

SYNTYPES: Peninsular Malaysia, Pulau Langkawi, Gunong

Raya, Md Aniff 15469, February 1911 (lectotype, designated here,

K); Kedah, Bukit Jenun, Md Haniff SFN 1210, 19 November 1915

(K[<2]). Burma, Mergui.

= PHOEBE TAVOYANA (Meissn.) Hook.f., Fl. Brit. India 5 (1886) 143;

Kochummen, Tree Fl. Malaya 4 (1989) 178.

CINNAMOMUM MICROPHYLLUM Ridl., Fl. Malay Penins. 3 (1924) 92, nom.

nov. pro Cinnamomum parvifolium Ridl.; Kochummen, Tree FI.

Malaya 4 (1989) 127.

Cinnamomum rhynchophyllum var. lampongum (Miq.) Ridl., FI.

Malay Penins. 3 (1924) 93. Basionym: Cinnamomum lampongum

Migq., Fl. Ned. Ind., Eerste Bijv. 3 (1861) 358.

= CINNAMOMUM RHYNCHOPHYLLUM Miq., Fl. Ned. Ind. 1 (1858) 895. Soh,

Blumea 56 (2011) 256.

Monimiaceae

46.

Palmeria paniculata Ridl., Trans. Linn. Soc., Bot. 9 (1916) 144. NAR B:

New Guinea, Mt Carstensz, Camp VIc, 5500 ft, C.B. Kloss s.n.,

1912-1913 (?BM).

= PALMERIA GRACILIS Perkins, Bot. Jahrb. 31 (1902) 745: Philipson, Fl.

Males. 10 (1986) 269.

There is a specimen in BM, labelled Camp Dundok Padang, which may

be the type.

Gard. Bull. Singapore 64(1) 2012

MAGNOLIALES

Annonaceae

47.

48.

50.

49.

Mezzettiopsis Ridl., Bull. Misc. Inform. Kew 1912 (1912) 389. TYPE:

Mezzettiopsis creaghii Ridl.

= OrorpuEa Blume, Bijdr. (1825) 18.

Uvariella Ridl., Fl. Malay Penins. 1 (1922) 35. TYPE: Uvariella leptopoda

(King) Ridl.

= Uvaria L., Sp. Pl. (1753) 536.

Alphonsea keithii Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911) 66.

TYPE: Thailand, Phangnga, Bangtaphan, 4. Keith 3/1, 16 April 1890

(holotype, SING (barcode: SING0048653)).

Artabotrys havilandii Ridl., Bull. Misc. Inform. Kew 1912 (1912) 382.

SYNTYPES: Borneo, Sarawak, Kuching, G.D. Haviland 1629

(lectotype, designated by Turner (2009a), K; isolectotype, SAR);

near Kuching, G.D. Haviland & C. Hose 3340, 24 November 1894

(BM, K, SAR); s. loc., O. Beccari P.B. 381 (K); O. Beccari P.B. 786;

O. Beccari P.B. 713 (K), O. Beccari P.B. 554 (FI-B, K),,.

= ARTABOTRYS MAINGAYI Hook.f. & Thomson, Fl. Brit. India 1 (1872) 55;

Turner, Folia Malaysiana 10 (2009) 62.

Beccari’s PB. 554 1s the type of Artabotrys pleianthus Diels, a synonym

of Artabotrys roseus Boerl.

ARTABOTRYS HIRTIPES Ridl., Bull. Misc. Inform. Kew 1912 (1912) 383;

Turner, Folia malaysiana 10 (2009) 86. SYNTYPES: Borneo,

Sarawak, Rejang Kapit, G.D. Haviland 2326, March 1893 (lectotype,

designated by Turner (2009a), K); Kuching, G.D. Haviland 2106,

19 January 1893 (K, SING); G.D. Haviland & C. Hose 2106, 29

November 1894 (K).

Disepalum grandiflorum Ridl., Bull. Misc. Inform. Kew 1912 (1912)

384. SYNTYPES: Borneo, Sarawak, Baram, C. Hose 142 (lectotype,

designated by Johnson (1989), K; isolectotypes, B, BM); Marudi, C.

Hose 214 (K).

= DISEPALUM ANOMALUM Hook.f., Trans. Linn. Soc. 23 (1860) 156;

Johnson, Brittonia 41 (1989) 375.

Ridley’s taxa among primitive angiosperms

aS.

aD:

56.

58.

iv)

DREPANANTHUS CARINATUS Ridl., Fl. Malay Penins. 1 (1922) 38.

Cyathocalyx carinatus (Ridl.) J.Sinclair, Gard. Bull. Singapore 14

(1955) 241. TYPE: Peninsular Malaysia, Selangor, Sungei Buluh,

F-W. Foxworthy 2263, December 1923 (holotype, K (barcode no.

K000691221); isotype. SING).

Goniothalamus caudifolius Ridl., Bull. Misc. Inform. Kew 1914 (1914)

324. TYPE: Peninsular Malaysia, Perak, Pondok Tanjong, 4.M.

Burn-Murdoch 180, 1909 (holotype, K (barcode no. K000691878);

isotype, K).

= GONIOTHALAMUS TENUIFOLIUS King, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 61 (1892) 71; Saunders, Bot. J. Linn. Soc. 142 (2003) 331.

GONIOTHALAMUS PARALLELIVENIUS Ridl., Bull. Misc. Inform. Kew 1912

(1912) 385, as ‘parallelovenius’; Saunders, Bot. J. Linn. Soc. 139

(2002) 244. TYPE: Borneo, Sarawak, O. Beccari P.B. 3772, s. dat.

(holotype, K (barcode no. K000691904): isotypes, FI-B. P).

Goniothalamus pendulifolius Ridl., Fl. Malay Penins. 5 (1925) 287.

TYPE: Peninsular Malaysia, Pahang. 6 miles north of Bentong, /.H.

Burkill & Md. Haniff SFN 16501, 5 November 1924 (lectotype,

designated by Saunders (2003), K (barcode no. K000691852):

isolectotypes, K (barcode no. K000691853), SING[*2]).

= GONIOTHALAMUS UVARIOIDES King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.

61 (1892) 78, Saunders, Bot. J. Linn. Soc. 142 (2003) 326.

GONIOTHALAMUS UNDULATUS Ridl., J. Fed. Malay States Mus. 10 (1920)

81. TYPE: Thailand, Tasan, C.B. Kloss 6836, s. dat. (holotype, K

(barcode no. K000691832): isotype, SING).

Melodorum breviflorum Ridl., J. Malayan Branch Roy. Asiat. Soc. 1

(1923) 51. FissisTIGMA BREVIFLORUM (Ridl.) I.M. Turner, comb.

nov. TYPE: Sumatra, Berastagi, H.N. Ridley s.n., 8 February

1921 (lectotype, designated here, K (barcode no. K000691997);

isolectotypes, K[*2]).

The generic name Me/odorum Lour. has a complex nomenclatural history

that is not entirely resolved (Ueda 1986). For a long period it was

wrongly applied to Fissistigma Griff. Merrill (1919) corrected

this mistake and transferred many species from Melodorum to

Fissistigma. However Ridley did not follow Merrill when publishing

this new species from Sumatra (or in his Flora of the Malay

Gard. Bull. Singapore 64(1) 2012

Peninsula). Melodorum breviflorum has the features typical of

Fissistigma including its climbing habit, scalariform leaf venation,

branched inflorescences, inner petals slightly smaller then outer

petals and excavated at the base. | am not able to match the specimens

to any other described Fissistigma species so I transfer Melodorum

breviflorum to Fissistigma.

Melodorum fagifolium Ridl., Bull. Misc. Inform. Kew 1912 (1912) 386.

Fissistigma fagifolium (Ridl.) Merr., Philipp. J. Sci., C. 15 (1919)

131. Fissistigma kingii (Boerl.) Burkill var. fagifolium (Ridl.) Airy

Shaw, Bull. Misc. Inform. Kew 1939 (1939) 238. TYPE: Borneo,

Sarawak, Entagut River, C. Hose 397, December 1894 (lectotype,

designated here, K (barcode no. K000574676); isolectotypes, A, B,

BMAK EYP):

= FISSISTIGMA KINGH (Boerl.) Burkill, Bull. Misc. Inform. 1935 (1935)

317. Turner, Nordic J. Bot. 27 (2009) 368.

There are two herbarium sheets at K with the relevant Hose collection

number. The better specimen 1s chosen as lectotype.

Melodorum longipetalum Ridl., Bull. Misc. Inform. Kew 1912 (1912)

387. FISSISTIGMA LONGIPETALUM (Ridl.) Merr., Philipp. J. Sci., C.

15 (1919) 133. TYPE: Borneo, Sarawak, Kuching, G.D. Haviland

2102 [Ngaing leg.], 11 January 1893 (holotype, K (barcode no.

K000574672); isotypes, BM, SAR, SING).

Melodorum ovalifolium Ridl., Bull. Misc. Inform. Kew 1912 (1912)

387. Fissistigma ovalifolium (Ridl.) Merr., Philipp. J. Sci., C. 15

(1919) 134. Oxymitra ovalifolia (Ridl.) J. Sinclair, Sar. Mus. J. 5

(1951) 607. Richella ovalifolia (Ridl.) Steenis, Blumea 12 (1964) 357.

FRIESODIELSIA OVALIFOLIA (RidlI.) I1.M. Turner, Blumea 55 (2010) 118.

SYNTYPES: Borneo, Sarawak, Kuching, G.D. Haviland & C. Hose

3151, 17 May 1894 (lectotype, designated by Turner (2010), K); G.D.

Haviland & C. Hose 3141, 17 May 1894 (K, SAR[*2], SING).

Melodorum paniculatum Ridl., Bull. Misc. Inform. Kew 1912 (1912)

386. FISSISTIGMA PANICULATUM (Ridl.) Merr., Philipp. J. Sci., C. 15

(1919) 134. TYPE: Borneo, Sarawak, near Kuching, G.D. Haviland

1845, October 1892 (holotype, K (barcode no. K000574667);

isotype, K).

One of the two sheets of Haviland 1845 at K has ‘Type’ written in Ridley’s

hand on it. I take this to be the holotype.

Ridley’s taxa among primitive angiosperms 23T

63. Melodorum rigidum Ridl., Bull. Misc. Inform. Kew 1912 (1912) 386.

Fissistigma rigidum (Ridl.) Mert., Philipp. J. Sci.. C. 15 (1919)

135. SYNTYPES: Bomeo, Sarawak, Kuching, G.D. Haviland 421

[Garai leg.] [d.k.m.a.] (lectotype. designated here, K (barcode no.

K000574656)): s. loc., O. Beccari PB. 393 (FI-B. K, NY(fragment)):.

= PYRAMIDANTHE PRISMATICA (Hook.f. & Thomson) Merr.. J. Straits

Branch Roy. Asiat. Soc., Spec. No. (1921) 262.

Of the syntypes at K, the Haviland specimen is annotated as ‘Type’ in

Ridley’s hand and therefore chosen as lectotype. When Haviland

began collecting in Sarawak he used a complex four-letter code for

specimens that combined elements of the date and specimen number

(Stapf 1907). He was advised to desist from this confusing practice

and use a simple running number sequence. Many of the earlier coded

collections were renumbered, but unfortunately he tended to give

different gatherings of the same species the same number. Therefore

it is important to use other information, such as date of collection,

when dealing with typification of species based on Haviland

collections. In this case, other herbaria (BO, SAR, SING) have G_D.

Haviland 42] specimens but these are of different collecting dates

compared to the K specimen, so are not isolectotypes.

64. Mezzettia pauciflora Ridl.. Bull. Misc. Inform. Kew 1912 (1912) 389.

TYPE: Borneo, Sarawak, Kuching, G.D. Haviland 1952, 1 December

1892 (lectotype, designated here, K (barcode no. K000574873:

isolectotypes, K, SAR, SING).

= MEZZETTIA UMBELLATA Becc., Nuovo Giorn. Bot. Ital. 3 (1871) 187: van

der Heijden & KeBler, Blumea 35 (1990) 220.

65. Mezzettiopsis creaghii Ridl., Bull. Misc. Inform. Kew 1912 (1912) 389.

OROPHEA CREAGHI (Ridl.) Leonardia & P.J.A. Kessler. Blumea 46

(2001) 149. TYPE: Borneo, British North Borneo [Sabah], Tinkango,

C.V. Creagh s.n., April 1895 (holotype, K (barcode no K000574862):

isotype, BM).

66. MILIvSA AMPLEXICAULIS Ridl., J. Straits Branch Roy. Asiat. Soc. 54 (1910)

13: Mols & KeBler, Blumea 48 (2003) 436. TYPE: Peninsular

Malaysia, Pulau Langkawi, Coah [Kwah], C. Curtis 3205, April 1896

(lectotype, designated here, SING (barcode nos. SING 0045995);

isolectotypes, K, SING (barcode no. SING 0045994), UC).

238

67.

68.

- 69.

70.

Gard. Bull. Singapore 64(1) 2012

Miliusa concinna Ridl., J. Fed. Malay States Mus. 10 (1920) 127.

SYNTYPES: Thailand, Phangnga, Hat Sunuk [Hat Sanuk], Koh

Lak, Hamid FMS Field No. 3802, 3 April 1919 (lectotype, designated

here, K (barcode no. K000691547); isolectotype, KEP); near Koh

Lak, Hamid FMS Field No. 3820, 13 April 1919 (K, KEP).

= POLYALTHIA EVECTA (Pierre) Finet & Gagnep., Bull. Soc. Bot. Fr. 53,

Mémoire 4 (1906) 91.

Miliusa filipes Ridl., J. Fed. Malay States Mus. 10 (1920) 81. TYPE:

Thailand, Tasan, C.B. Kloss 6968 (holotype, K).

MILIUSA PARVIFLORA Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911)

65; Mols & KeBler, Blumea 48 (2003) 450. SYNTYPES: Peninsular

Malaysia, Perlis, Tebing Tinggi, H.N. Ridley 15340, March 1910

(lectotype, designated here, SING; isolectotypes, BM, K); limestone

rocks, Bukit Lagi, H.N. Ridley 15239, March 1910 (SING);.

Mols & Kefler (2003) cited Ridley 15340 as the holotype of M. parviflora

but the original protologue cites both 15239 and 15340. Though

misused terms relating to typification can be corrected [ICBN

(McNeill et al. 2006) Art. 9.8], Mols and KeBler’s statement is not a

valid lectotypification for a publication after 1 January 2001 because

‘designated here’ or its equivalent was not employed [ICBN Art. 7.11].

MITREPHORA ALBA Ridl., J. Fed. Malay States Mus. 5 (1915) 158,

Weerasooriya & Saunders, Syst. Bot. Monogr. 90 (2010) 39. TYPE:

Thailand, Koh Samui western side, H.C. Robinson 5717, 13 May

1913 (lectotype, designated here, K (barcode no. K000691973);

isolectotypes, K[ 2] (barcode no. K000691974)).

Of the three sheets of Robinson 5717 at K, two have original field tickets

labelled with the date and Koh Samui. The chosen lectotype also

has Robinson’s collection label giving more details of the plant and

collecting location. Both sheets are labelled *Mitrephora alba Ridl.’ in

Ridley’s hand. The third sheet does not have any of Robinson’s labels.

Mitrephora crassipetala Ridl., J. Straits Branch Roy. Asiat. Soc. 54

(1912) 13. TYPE: Peninsular Malaysia, Pahang, Tahan River,

Mat s.n., 23 September 1893 (holotype: SING (barcode no. SING

0119273)).

= ANAXAGOREA JAVANICA [Blume, FI. Javae Anon. (1830) 66, t. 32] var.

JAVANICA, Sinclair, Gard. Bull. Singapore 14 (1955) 347, 466.

Ridley’s taxa among primitive angiosperms

I Ex

73:

74.

hos

76.

239

MITREPHORA KEITHII Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911)

66; Weerasooriya & Saunders, Syst. Bot. Monogr. 90 (2010) 70.

TYPE: Thailand, Phangnga, Bangtaphan, 4. Keith 373, 13 May 1890

(holotype, SING (barcode no. SING 0045997): isotype, K).

Mitrephora obtusa Blume var. glabra Ridl., Sarawak Mus. J. 1 (1913)

86. TYPE: Borneo, Sarawak, Mt Koum limestone, G.D. Haviland

1714 [Kalong leg.], 22 September 1892 (lectotype, designated here,

K (barcode no. K000574547); isolectotypes, BM, L[<2], SAR[*2],

SING[*2]).

= MITREPHORA UNIFLORA Weeras. & R.M.K. Saunders, Syst. Bot. Monogr.

90 (2010) 122.

MITREPHORA RUFESCENS Ridl., Bull. Misc. Inform. Kew 1912 (1912)

385. SYNTYPES: Sumatra, Sungei Buluk, Padang, O. Beccari PS.

968 (lectotype, designated by Weerasooriya & Saunders (2010),

BM; isolectotypes, FI-B, K), September 1878; O. Beccari P.S. 916,

September 1878 (BM, FI-B, K[ <2], L). Borneo, Sarawak, O. Beccari

PB. 1616 (FI-B, K); Matang, G.D. Haviland 1035 [leg. Garai]

[e.c.q.b.] (K, SAR, SING); near Bau limestone G.D. Haviland 2249,

6 December 1893 (K, SAR).

The Sarawak syntypes are Mitrephora longipetala Miq. (Weerasooriya &

Saunders 2010).

Orophea setigera Ridl., Trans. Linn. Soc., Bot. 9 (1916) 16. TYPE: New

Guinea, Mt Carstensz, Camp III to VIII, 2500-4900 ft, C.B. Kloss

s.n. (holotype, BM).

Oxymitra linderifolia Ridl., Bull. Misc. Inform. Kew 1912 (1912) 385.

Friesodielsia linderifolia (Ridl.) Steen., Blumea 12 (1964) 360.

SYNTYPES: Borneo, Sarawak, Kuching, G.D. Haviland 2333, 19

April 1893 (lectotype, designated here, K (barcode no. K000691760);

isolectotypes, BM, K, SAR, SING); near Kuching, G.D. Haviland &

C. Hose 3335, 11 October 1894 (K, BM, GH, SAR).

= FRIESODIELSIA GLAUCA (Hook.f. & Thomson) Steenis, Blumea 12 (1964)

359:

The Haviland collection was erroneously referred to as 3333 rather than

2333 in the original publication.

240

78.

IGE

80.

81.

Gard. Bull. Singapore 64(1) 2012

POLYALTHIA ANGUSTISSIMA Ridl., J. Straits Branch Roy. Asiat. Soc. 54(1910)

11; Sinclair, Gard. Bull. Singapore 14 (1955) 286. SYNTYPES:

Singapore: Bukit Timah, H.N. Ridley 8050 (lectotype, designated by

Bunchalee & Chantaranothai (2006), SING; isolectotype, K); sandy

woods, Changi, H.N. Ridley 5917 (SING, CAL); Gardens’ Jungle,

H.N. Ridley 4813 (K, SING). Peninsular Malaysia, Johore: Kwala

Sembrong, H.J/. Kelsall 4047 (CAL, SING).

POLYALTHIA ASTERIELLA Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

169; Sinclair, Gard. Bull. Singapore 14 (1955) 309. TYPE: Perak,

Taiping Hills, Maxwell’s Hill, H.N. Ridley 2986, February 1891

(holotype, K (barcode no. K000691443); isotypes, CAL, SING).

Polyalthia castanea Ridl., J. Fed. Malay States Mus. 8 (1917) 14. TYPE:

Sumatra, Korinchi (Kerinci), Sandaran Agong at 2,450 feet. Small

tree, flowers orange-brown.

I have failed to locate any (type) material of Polyalthia castanea.

POLYALTHIA CHRYSOTRICHA Ridl., Fl. Malay Penins. | (1922) 57; Sinclair,

Gard. Bull. Singapore 14 (1955) 302. TYPE: Peninsular Malaysia,

Selangor, Menuang Gasing, C.B. Kloss s.n., February 1912 (holotype,

K (barcode no. K000691577)).

Polyalthia coriacea Ridl., Bull. Misc. Inform. Kew 1912 (1912) 383.

ENICOSANTHUM CORIACEUM (Ridl.) Airy Shaw, Bull. Misc. Inform.

Kew 1939 (1939) 277. SYNTYPES: Borneo, Sarawak, Mt Buan

Limestone, G.D. Haviland 2002 [Garai leg.], 15 December

1892 (lectotype, designated here, K (barcode no. K000691411);

isolectotype, SING); Bidi, C.J. Brooks 1057, April 1909 (BM).

Polyalthia curtisii Ridl., J. Straits Branch Roy. Asiat. Soc. 54 (1910) 11.

TYPE: Peninsular Malaysia, Penang, Telok Bahang, C. Curtis 3644

(holotype, SING; isotype, K).

= POLYALTHIA GLABRA (Hook.f. & Thomson) J. Sinclair, Gard. Bull.

Singapore 14 (1955) 315.

Ridley’s taxa among primitive angiosperms 24]

83.

84.

85.

86.

87.

Polyalthia eriantha Ridl., Bull. Misc. Inform. Kew 1912 (1912) 384, as

‘erianthia’. Enicosanthum erianthum (Ridl.) Airy Shaw, Bull. Misc.

Inform. Kew 1939 (1939) 276. Meiogyne eriantha (Ridl.) J.Sinclair,

Sarawak Mus. J. 5 (1951) 604. TYPE: Borneo, Sarawak, Kuching,

G.D. Haviland 410 [d.e.q.b.] (holotype K (barcode no. K000691133,);

isotypes, K[ <2] (barcode nos. K000691134, K000691135), SAR).

= MEIOGYNE VIRGATA (Blume) Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865)

12; van Heusden, Blumea 38 (1994) 495.

The holotype sheet is annotated ‘Type’ in Ridley’s hand.

Polyalthia hirta Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 168,

non (Mig.) F. Muell. (1877); POoLYALTHIA HIRTIFOLIA J. Sinclair,

Gard. Bull. Singapore 14 (1955) 300, nom. nov. TYPE: Peninsular

Malaysia, Penang, Pulau Butong Reserve, C. Curtis 2745, March

1892 (lectotype, designated here, K (barcode no. K000691452);

isotypes, BM, K (barcode no. K000691453), SING).

Polyalthia montana Ridl., Journ. Linn. Soc., Bot. 41 (1913) 288. TYPE:

Peninsular Malaysia, Selangor, Ulu Langat, Menuang Gasing, C.B.

Kloss s.n., February 1912 (lectotype, designated by Turner (2009b),

K; isolectotype, BM).

= TRIVALVARIA CosTATA (Hook.f. & Thomson) I.M. Turner, Kew Bull. 64

(2009) 577.

POLYALTHIA PARVIFLORA Ridl., J. Straits Branch Roy. Asiat. Soc. 61

(1912) 49; Gard. Bull. Singapore 14 (1955) 299. SYNTYPES:

Peninsular Malaysia, Pulau Langkawi, Kuala Malacca, C. Curtis

2533, September 1890 (lectotype, designated here, SING (barcode

no. SING 0059296); isolectotype, CAL); Pulau Tengah, H.N. Ridley

15893, April 1911 (K, BM, SING (barcode no. SING 0059295)).

POLYALTHIA PUMILA Ridl., J. Straits Branch Roy. Asiat. Soc. 54 (1910)

12; Gard. Bull. Singapore 14 (1955) 289. SYNTYPES: Peninsular

Malaysia, Perak, Dindings, Sera Woods, H.N. Ridley 7996, March

1896 (lectotype, designated here, SING (barcode: SING 0059298);

isolectotype, K); Johore, Gunong Janing, H.W. Lake & H.J. Kelsall

s.n., 20 October 1892 (SING (barcode: SING 0059299)).

88.

89.

90.

Gard. Bull. Singapore 64(1) 2012

Polyalthia purpurea Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

168. TYPE: Singapore, cultivated in Singapore Botanic Gardens,

H.N. Ridley 12014, June 1904 (lectotype, designated here, K (barcode

no. K000691573); isolectotype, SING).

= POLYALTHIA SCLEROPHYLLA Hook.f. & Thomson, FI. Brit. Ind. 1 (1872)

65; Sinclair, Gard. Bull. Singapore 14 (1955) 311.

Polyalthia velutinosa Ridl., J. Straits Branch Roy. Asiat. Soc. 59 (1911)

64. TYPE: Peninsular Malaysia, Pulau Lankawi, Gunong Raya at

1500 feet, [Aniff] Haniff 15552, February 1911 (holotype, SING

(barcode no. SING 0058808); isotypes, BM, K).

= POLYALTHIA CINNAMOMEA Hook.f. & Thomson, FI. Ind. 1 (1855) 138;

Sinclair, Gard. Bull. Singapore 14 (1955) 286.

Popowia rufescens Ridl., Bull. Misc. Inform. Kew 1926 (1926) 59. TYPE:

Sumatra, Mentawai Islands, Siberut, C.B. Kloss 14607 (holotype, K

(barcode no. K000691711); isotype, BO).

= PopowlA PISOCARPA (Blume) Endl. ex Walp., Repert. Bot. Syst. 1 (1842)

74; Sinclair, Gard. Bull. Singapore 14 (1955) 468.

Unona conchyliata Ridl., Bull. Misc. Inform. Kew 1912 (1912) 384.

Desmos conchyliatus (Ridl.) Merr., J. Straits Branch Roy. Asiat.

Soc., Spec. No., (1921) 255. TYPE: Borneo, Sarawak, Kuching,

G.D. Haviland 771 [1779], 13 October 1892 (holotype, K (barcode

no. K000691930)).

=MEIOGYNE MONOSPERMA (Hook.f. & Thomson) Heusden, Blumea 38

(1994) 502.

Ridley referred to Haviland 1779 in describing the species. There are

three sheets at Kew bearing this number, but all differ in collection

date. One, which has the original collection number 771, has been

annotated by Ridley as type. Other herbaria (B, BO, L, SAR, SING)

have Haviland specimens with the number 1779 but none appears to

be a duplicate of the holotype.

Unona filipes Ridl., J. Linn. Soc. Bot. 41 (1913) 287. Desmos filipes (Ridl.)

Ridl., J. Straits Branch Roy. Asiat. Soc. 75 (1917) 6. DASYMASCHALON

FILIPES (Ridl.) Ban, Bot. Zhurn. 60(2): 229; Wang et al., Syst. Bot. 34

(2009) 259. SYNTYPES: Peninsular Malaysia, Selangor, Menuang

Gasing, C.B. Kloss s.n., February 1912 (lectotype, designated here,

K (barcode no., K000198741); isolectotypes, K[<2] (barcode nos.

K000198739, K00019840); Perak, Fr B. Scortechini 342, September

Ridley’s taxa among primitive angiosperms 243

93.

94.

95.

1892 (BO, CAL, K, SING); Larut, L. Wray 609, May 1886 (K), Larut,

Dr King’s Collector [H.H. Kunstler] 529/ (K).

Nurmawati (2003) referred to the Herbarium Bogoriense duplicate of

the Scortechini collection as the lectotype of Unona filipes but

this is invalid as a lectotypification as the term ‘designated here’

or equivalent was not used [ICBN Art. 7.11]. I choose one of the

Kloss specimens as the lectotype because, not only are they good

specimens, but the protologue was published in the account of

Kloss’s expedition to Gunung Menuang Gasing. The chosen sheet

has the species name in Ridley’s hand.

Unona jambosifolia Ridl., Bull. Misc. Inform. Kew 1912 (1912) 384.

Desmos jambosifolius Merr., J. Straits Branch Roy. Asiat. Soc. Spec.

No. (1921) 255. Polyalthia jambosifolia (Ridl.) D.M. Johnson,

Contrib. Univ. Mich. Herb. 22 (1999) 99. SYNTYPES: Borneo,

Sabah, Kudat, M. Fraser 174, July 1885, (lectotype, designated here,

K (barcode no. K000691580); Batu Putik, CV. Creagh s.n. (BM, K

(barcode no. K000691581);.

= POLYALTHIA MicROTUS Miq., Ann. Mus. Bot. Lugd. Bat. 2 (1865) 13.

The Creagh specimen is Polyalthia igniflora D.M. Johnson.

Unona purpurata Ridl., Sar. Mus. J. 1(3) (1913) 79, nom. illegit. Desmos

purpuratus (Ridl.) Merr., J. Straits Branch Roy. Asiat. Soc., Spec.

No., (1921) 256, nom. illegit. TYPE: Same as Unona conchyliata

Ridl.

= MEIOGYNE MONOSPERMA (Hook.f. & Thoms.) Heusden, Blumea 38

(1994) 502.

UVARIA CAULIFLORA Ridl., Bull. Misc. Inform. Kew 1912 (1912) 382.

SYNTYPES: Borneo, Sarawak, O. Beccari PB. 1120 (FI-B, K):

Tegora, G.D. Haviland 417 [c.k.q.a.] [Garai leg.] (lectotype,

designated here, K (barcode no. K000380300); isolectotype,

K(barcode no. K000380298)); near Kuching, G.D. Haviland 417

[Ulak leg.], 6 October 1892 (K); foot of Bembang, G.D. Haviland

409 (K, SAR).

The lectotype sheet selected is annotated ‘Type’ in Ridley’s hand.

244

96.

Die

98.

100.

Gard. Bull. Singapore 64(1) 2012

UvarRIA LANUGINOSA Ridl., Bull. Misc. Inform. Kew 1912 (1912) 382.

TYPE: Borneo, Sarawak, near Kuching, G.D. Haviland & C. Hose

3334, 13 November 1894 (holotype, K (barcode no. K000380301);

isotypes, BM, SAR).

In the protologue, Ridley cited the type as 334, but the correct number is

3334.

Xylopia congesta Ridl., Bull. Misc. Inform. Kew 1912 (1912) 387.

POLYALTHIA CONGESTA (Ridl.) J. Sinclair, Sarawak Mus. Journ. 5

(1951) 607. TYPE: Borneo, Sarawak, O. Beccari P.B. 2654 (holotype,

K (barcode no. K000691591); isotypes, BM, FI-B).

XYLOPIA CORHFOLIA Ridl., Bull. Misc. Inform. Kew 1912 (1912) 388.

SYNTYPES: Borneo, Sarawak, near Kuching, G.D. Haviland &

C. Hose 3337, 6 November 1894 (lectotype, designated here, K

(barcode no. K000574692); isolectotype, SAR); near Kuching, G.D.

Haviland 1906 [leg. Garai], 26 October 1892 (K); s. loc., O. Beccari

PB. 2652 (FI-B, K), O. Beccari PB. 3333.(F1-B, );.

Xylopia havilandii Ridl., Bull. Misc. Inform. Kew 1912 (1912) 388.

SYNTYPES: Borneo, Sarawak, near Kuching, G.D. Haviland

2334, 17 May 1893 (lectotype, designated here, K (barcode no.

K000574708); isolectotypes, SAR, SING); G.D. Haviland & C.

Hose 3352, 29 December 1894 (K).

= XYLopIA FUSCA Maingay ex Hook.f. & Thomson, FI. Brit. India 1 (1872)

83.

Xylopia lanceola Ridl., Bull. Misc. Inform. Kew 1912 (1912) 389.

SYNTYPES: Borneo, Sarawak, O. Beccari P.B. 3368 (K); O. Beccari

PB. 1968 (lectotype, designated by Sinclair (1951), K (barcode no

K000574695):; isolectotypes, A, B, FI-B).

= XyLopia cAuDATA Hook.f. & Thomson var. RETICULATA J. Sinclair,

Sarawak Mus. J. 5 (1951) 608.

Ridley misreported PB. 1968 as 1908 in the protologue — an excusable

error as the labels are hand-written and the down stroke on the 6 is

very feint. In K the specimen G.D. Haviland 2066 is annotated in

Ridley’s hand as type of Xy/lopia lanceola although the specimen

was not cited in the original publication.

Ridley’s taxa among primitive angiosperms

101.

102.

ill.

245

XYLOPIA PULCHELLA Ridl., Bull. Misc. Inform. Kew 1912 (1912) 388.

TYPE: Borneo, Sarawak, near Kuching, G.D. Haviland 2101, 16

January 1893 (holotype, K (barcode no. K000574761): isotype,

SAR[*2]).

Xylopia rotundata Ridl., Bull. Misc. Inform. Kew 1926 (1926) 469.

TYPE: Borneo, Sarawak, Kuching, Siol, 7 January 1924, Jantan

bin Omar [Foxworthy’s collector] 00357 (holotype, SING; isotype,

NY).

= DREPANANTHUS BIOVULATUS (Boerl.) Surves. & R.M.K. Saunders, Taxon

59 (2010) 1730.

Sphaerocoryne (Scheff. ex Boerl.) Scheff. ex Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 8. Basionym: Polyalthia section Sphaerocoryne

Scheff. ex Boerl., [Cat. Pl. Phan. 1 (1899) 26, nomen] Icon. Bogor. 1

(1899) 196. TYPE: Sphaerocoryne aberrans (Maingay ex Hook.f. &

Thomson) Ridl. (designated by Fries (1959)).

Although Ridley made no direct reference to Boerlage’s sectional name,

ICBN rules apply such that the genus must be considered to be based

on section Sphaerocoryne [ICBN Art. 33.3, viz. Ex. 9].

Canangium latifolium {[Hook.f. & Thomson] Pierre ex Ridl., Fl. Malay

Pen. 1 (1922) 44, nom. superfi. Unona latifolia Hook.f. & Thomson,

FI. Brit. India 1 (1872) 60, non U. latifolia Dunal.

Ridley’s intention was to transfer Unona latifolia Hook.f. & Thomson

to Canangium, but as this is a later homonym the transfer is

invalid (hence square brackets in citation) and as a new name the

combination is superfiuous as Pierre had provided the replacement

Unona brandisiana for Hooker and Thomson’s name, which Safford

had already transferred to Canangium (Turner & Veldkamp 2009).

Canangium monospermum (Hook.f. & Thomson) Ridl., J. Straits Branch

Roy. Asiat. Soc. 75 (1917) 7. MEIOGYNE MONOSPERMA (Hook.f. &

Thomson) Heusden, Blumea 38 (1994) 502. Basionym: Cananga

monosperma Hook.f. & Thomson, FI. Brit. Ind. 1 (1872) 57.

246

Vi.

Vil.

Vill.

Gard. Bull. Singapore 64(1) 2012

Desmos dasymaschalus (Blume) Safford var. wallichii (Hook.f. &

Thomson) Ridl., Fl. Malay Penins. 1 (1922) 47. Dasymaschalon

blumei Finet & Gagnep. var. wallichii (Hook.f. & Thomson) Ban,

Bot. Zhurn. 60 (1975) 228. Dasymaschalon dasymaschalum (Blume)

I.M. Turner var. wallichii (Hook.f. & Thomson) I.M. Turner, Gard.

Bull. Singapore 58 (2007) 277. DASYMASCHALON WALLICHII (Hook.f.

& Thomson) Jing Wang & R.M.K. Saunders, Syst. Bot. 34 (2009)

263. Basionym: Unona dasymaschala var. wallichii Hook.f. &

Thomson, FI. Ind. | (1855) 135.

Desmos filipes (Ridl.) Ridl., J. Straits Branch Roy. Asiat. Soc. 75 (1917) 6.

DASYMASCHALON FILIPES (Ridl.) Ban, Bot. Zhurn. 60(2): 229; Wang

et al., Syst. Bot. 34 (2009) 259. Basionym: Unona filipes Ridl., J.

Linn. Soc., Bot. 41 (1913) 287.

MEZZETTIA HAVILANDI (Boerl.) Ridl., Sarawak Mus. Journ. 1(3) (1913)

96; van der Heijden & Kessler, Blumea 35 (1990) 221. Basionym:

Mezzettia parviflora Becc. var. havilandii Boerl., Icon. Bogor. |

(1899) 111.

Mitrephora chrysocarpa (Miq.) Boerl. var. sakumbangensis (Miq.)

Ridl., Sar. Mus. J. 1 (1913) 87. Basionym: Orophea chrysocarpa

Miq. var. sakumbangensis Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865)

24.

= PSEUDUVARIA RETICULATA (Blume) Miq., Fl. Ned. Ind. 1(2) (1858) 33; Su

& Saunders, Syst. Bot. Monogr. 79 (2006) 150.

Polyalthia crinita (Hook.f. & Thomson) Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 6. Basionym: Unona crinita Hook.f. &

Thomson, FI. Brit. Ind. 1 (1872) 61.

= POLYALTHIA STENOPETALA (Hook.f. & Thomson) Finet & Gagnep., Bull.

Soc. Bot. France Mem. 4 (1906) 96; Sinclair, Gard. Bull. Singapore

14 (1955) 285. (Note that this combination is often attributed to

Ridley (J. Straits Branch Roy. Asiat. Soc. 75 (1917) 6) but Finet and

Gagnepain have priority on the name by more than a decade).

Ridley’s taxa among primitive angiosperms 247

Xl.

Ml,

Xlll.

XIV.

XV.

XVI.

Polyalthia desmantha (Hook.f. & Thomson) Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 6. Desmos desmanthus (Hook.f. & Thomson)

Saff., Bull. Torrey Bot. Club 39 (1912) 508. PoLYALTHIA CAULIFLORA

Hook.f. & Thomson var. DESMANTHA (Hook.f. & Thomson) J.

Sinclair, Gard. Bull. Singapore 14 (1955) 295. Basionym: Unona

desmantha Hook.f. & Thomson, FI. Brit. Ind. 1 (1872) 61.

Polyalthia submitrata (Miq.) Ridl., Sar. Mus. J. 1 (1913) 82. Basionym:

Monoon submitratum Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 16.

= SPHAEROCORYNE AFFINIS (Teijsm. & Binn.) Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 8.

Polyalthia wrayi (Hemsl.) Ridl., J. Straits Branch Roy. Asiat. Soc. 75

(1917) 6. Desmos wrayi (Hemsl.) Saff., Bull. Torrey Bot. Club 39

(1912) 507. POLYALTHIA CAULIFLORA Hook.f. & Thomson var. WRAYI

(Hemsl.) J. Sinclair, Gard. Bull. Singapore 14 (1955) 296. Basionym:

Unona wrayi Hemsl., Hook. Icon. Pl. 16 (1887) t. 1553.

Popowia nervosa (Hook.f.) Ridl., Fl. Malay Penins. 1 (1922) 78.

TRIVALVARIA NERVOSA (Hook.f. & Thomson) J. Sinclair, Gard. Bull.

Singapore 14 (1955) 197. Basionym: Ellipeia nervosa Hook.f. &

Thomson, FI. Brit. Ind. 1 (1872) 52.

Popowia pumila (King) Ridl., Fl. Malay Penins. | (1922) 78. Trivalvaria

pumila (King) J. Sinclair, Gard. Bull. Singapore 14 (1955) 198.

Basionym: Ellipeia pumila King, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 61 (1892) 27.

= TRIVALVARIA CosTATA (Hook.f. & Thomson) I.M. Turner, Kew Bull. 64

(2009) 577.

Sphaerocoryne aberrans (Maingay ex Hook.f. & Thomson) Ridl.,

J. Straits Branch Roy. Asiat. Soc. 75 (1917) 8. Popowia aberrans

(Maingay ex Hook.f. & Thomson) Pierre ex Finet & Gagnep., Bull.

Soc. Bot. Fr. Mem. 4(2) (1906) 109. Melodorum aberrans (Maingay

ex Hook.f. & Thomson) J.Sinclair, Gard. Bull. Singapore 14 (1953)

41. Mitrella aberrans (Maing. ex Hook.f. & Thomson) Ban, Bot.

Zhurn. 59(2) (1974) 244. Basionym: Polyalthia aberrans Maingay

ex Hook.f. & Thomson, FI. Brit. Ind. 1 (1872) 67.

= SPHAEROCORYNE AFFINIS (Teijsm. & Binn.) Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 8.

248

XVil.

XVIII.

XIX.

Magnoliaceae

103.

Gard. Bull. Singapore 64(1) 2012

SPHAEROCORYNE AFFINIS (Teijsm. & Binn. ex Boerl.) Ridl., J. Straits

Branch Roy. Asiat. Soc. 75 (1917) 8. Basionym: Polyalthia affinis

Teijsm. & Binn. ex Boerl., Ic. Bogor | (1899) t. 63.

Sphaerocoryne siamensis (Boerl.) Scheff. ex Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 8. Basionym: Polyalthia siamensis Boerl., Ic.

Bogor | (1899) 195.

=SPHAEROCORYNE AFFINIS (Te1jsm. & Binn.) Ridl., J. Straits Branch Roy.

Asiat. Soc. 75 (1917) 8.

Ridley cited three synonyms when making this combination: Unona

mesnyi Pierre (Fl. Forest. Cochinch. (1881) t. 17), Popowia mesnyi

Craib (Bull. Misc. Inform. Kew 1914 (1914) 5) and the apparently

unpublished Popowia edulis Pierre. The first two, predating Boerlage’s

Polyalthia siamensis, at first sight makes Ridley’s new combination

superfluous (though this does not invalidate the combination [ICBN

Art. 52.3]); but Pierre cited Polyalthia aberrans Maingay ex Hook.f.

& Thomson (FI. Brit. Ind. 1 (1872) 67) and Melodorum clavipes

Hance (J. Bot. 15 (1877) 328) as synonyms when describing Unona

mesnyi, rendering the name illegitimate. Craib, while explicitly

excluding Polyalthia aberrans from Unona mesnyi, did not refer to

Melodorum clavipes, so failed to validate Pierre’s taxon as Popowia

mesnyi. Therefore Sphaerocoryne siamensis was not superfluous

when Ridley published it.

Uvariella leptopoda (King) Ridl., Fl. Malay Penins. | (1922) 35. Uvarta

LEPTOPODA (King) R.E. Fr., Ark. Bot. 3 (1953) 40. Basionym: Ellipeia

leptopoda King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 61 (1892) 25.

Talauma beccarii Ridl., Bull. Misc. Inform. Kew 1912 (1912) 381.

Magnolia candollei (Blume) Noot. var. beccarii (Ridl.) Noot.,

Blumea 32 (1987) 375. MAGNOLIA LILIIFERA Baill. var. BECCARII

(Ridl.) Govaerts, World Checklist & Bibliogr. Magnoliac. (1996)

71. TYPE: Borneo, Sarawak, Mt Sadong, O. Beccari P.B. 3959,

November 1867 (holotype, K (barcode no. K000681543); isotype,

FI-B).

Ridley’s taxa among primitive angiosperms

104.

105.

XX.

249

Talauma oblanceolata Ridl., Fl. Malay Penins. 5 (1925) 286. TYPE:

Peninsular Malaysia, Pahang/Selangor, Sempang, H.N. Ridley

15590, April 1911 (holotype, K (barcode no. K000395248); isotype,

SING).

= MAGNOLIA LILIIFERA Var. OBOVATA (Korth.) Govaerts, World Checklist

& Bibliogr. Magnoliac. (1996) 71.

Talauma singapurensis Ridl., Bull. Misc. Inform. Kew 1914 (1914)

323. Magnolia candollei (Blume) Noot. var. singapurensis (Ridl.)

Noot., Blumea 32 (1987) 375. MAGNOLIA LILIFERA Baill. var.

SINGAPURENSIS (Ridl.) Govaerts, World Checklist & Bibliogr.

Magnoliac. (1996) 71. SYNTYPES: Singapore, Chan Chu Kang,

HN. Ridley 5091 (lectotype, designated by Dandy (1928), BM;

isolectotypes, B (?destroyed), K, SING); Bukit Mandai Road, H.N.

Ridley 3656 (BM (carpological coll.), K).

K has a sheet of H.N. Ridley 2110, Bukit Mandai, January 1891, on which

are mounted capsules containing material from the two syntypes.

Presumably Ridley used all these collections in describing the

species, but failed to cite 2110 in the protologue. Dandy (1928)

effectively lectotypified the species name to a BM collection nearly

half a century before Nooteboom (1987) proposed the duplicate in

SING as lectotype.

Talauma longifolia (Blume) Ridl., J. Fed. Malay States Mus. 7 (1916)

38. Basionym: TZalauma mutabilis Blume var. longifolia Blume, FI.

Jav. Magnol. (1829) 37.

= MAGNOLIA LILUFERA Baill. var. LILIIFERA, Govaerts, World Checklist &

Bibliogr. Magnoliac. (1996) 33.

NYMPHAEALES

Nymphaeaceae

XX1.

Barclaya kunstleri (King) Ridl., Fl. Malay Penins. | (1922) 117.

HybROSTEMMA KUNSTLERI (King) B.C. Stone, Gard. Bull. Singapore

35 (1982) 71. Basionym. Barclaya motleyi var. kunstleri King, J.

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 58 (1887) 390.

250 Gard. Bull. Singapore 64(1) 2012

PIPERALES

Aristolochiaceae

106. ARISTOLOCHIA GLAUCIFOLIA Ridl., Bull. Misc. Inform. Kew 1925 (1925)

88; Ding Hou, Fl. Males. 10 (1984) 93. TYPE: Sumatra, Bencoolen,

C.J. Brooks s.n. (holotype, K (barcode no. K000820375)).

107. ARISTOLOCHIA KLOSSII Ridl., Bull. Misc. Inform. Kew 1926 (1926) 78,

Ding Hou, FI. Males. 10 (1984) 93. TYPE: Sumatra, Mentawai

Islands, Sipora, C.B. Kloss SEN 14767, 24 October 1924 (holotype,

K (barcode no. K000820374); isotypes, BO, SING).

108. ARISTOLOCHIA MINUTIFLORA Ridl. ex Gamble, Bull. Misc. Inform. Kew

1910 (1910) 79; Ding Hou, FI. Males. 10 (1984) 93. SYNTYPES:

Peninsular Malaysia, Perak, H.N. Ridley 8022, March 1896

(lectotype, designated here, SING; isolectotype, BM); H.N. Ridley

10259 (SING), Kunstler [King’s Collector] 1964.

109. Aristolochia pithecurus Ridl., J. Bot. 52 (1914) 296. PARARISTOLOCHIA

PITHECURUS (Ridl.) Mich. J. Parsons, Bot. J. Linn. Soc. 120 (1996)

207. TYPE: New Guinea, Central Province, Sogeri Region, Mt

Korkoko 2500 ft, H.O. Forbes 621 (holotype, BM; isotypes, L,

WRSL).

110. Thottea hirsuta Ridl., Journ. As. Soc. Mal. 1 (1923) 87. TYPE: Sumatra,

Sibolangit, Bukit Pasang, Mohamed Nur SFN 7223, 11 August

1921 (holotype, K; isotypes, BM, BO, SING (barcode no. SING

0097461)).

= THOTTEA MACRANTHA (Boerl.) Ding Hou, Fl. Males. 10 (1984) 81.

NL THOTTEA PARVIFLORA Ridl., J. Straits Branch Roy. Asiat. Soc. 57 (1911)

89; Ding Hou, FI. Males. 10 (1984) 67. TYPE: Peninsular Malaysia, |

Perak, Upper Perak, Temengoh hill woods, H.N. Ridley 14580, :

August 1904 (holotype, SING (barcode no. SING 0097415)). |

Ridley’s taxa among primitive angiosperms

Piperaceae

112:

114.

15.

116.

AE?

Peperomia hedyotidea Ridl., Trans. Linn. Soc., Bot. 9 (1916) 142.

SYNTYPES: New Guinea, Mt Carstensz, Camp VIc, C.B. Kloss

s.n., 22 February 1913 (lectotype, designated here, BM (barcode

BM000950712); isolectotype, K); Camp III, 2500 ft, C.B. Kloss s.n.,

Febuay 1913 (BM (barcode no. BM000950711)); Camp VIb, C-.B.

Kloss s.n., 2 February 1913, (BM (barcode no. BM000950710), K).

At both BM and K all the syntypes are mounted on one herbarium sheet.

Peperomia malaccensis Ridl., Bull. Misc. Inform. Kew 1895 (1895) 185.

TYPE: Peninsular Malaysia, Malacca, R. Derry s.n. (holotype, K

(barcode no. KO00820265)).

? = Peperomia gemella Mig.

Peperomia muscicola Ridl., Trans. Linn. Soc., Bot. 9 (1916) 141.

TYPE: New Guinea, Mt Carstensz, Camp I, 500 ft, C.B. Kloss

s.n., November—December 1913 (holotype, BM (barcode no.

BM009507 14): isotype, K).

Peperomia obliqua Ridl., Trans. Linn. Soc., Bot. 9 (1916) 142, non P.

obliqua Ruiz & Pav. (1798). PEPEROMIA RIDLEYI C. DC., Candollea

1 (1923) 407. TYPE: New Guinea, Camp IX to XI, 5500 to 8300

ft, C.B. Kloss s.n., February 1912 (holotype, BM (barcode no.

BM000950715).

Peperomia purpurea Ridl., Trans. Linn. Soc., Bot. 9 (1916) 141; non

P. purpurea Ruiz & Pavon. (1798). PEPEROMIA COLOSSINA & Dex.

Candollea 1 (1923) 383. TYPE: New Guinea, Camps I to III, 500 to

2500 ft, C_B. Kloss s.n., 22 November 1912 (holotype, BM (barcode

no. BM000950716); isotype, K).

Peperomia villosa Ridl., J. Fed. Malay States Mus. 8 (1917) 80, non

P. villosa C. DC. (1866). PEPEROMIA KERINCIENSIS I.M. Turner,

nom. nov. TYPE: Sumatra, Korinchi (Kerinci) Peak, Siolak, Daras

3000 ft, H.C. Robinson & C.B. Kloss s.n., 19 March 1914, annot.

‘Peperomia hispidula Ridl.’ (lectotype, designated here, K (barcode

no. K000820274)).

I have not traced any material labelled Peperomia villosa Ridl. In the Kew

herbarium there is a Robinson & Kloss specimen that agrees with

the details given in the protologue and which is labelled “Peperomia

hispidula Ridl.’, a name apparently never published by Ridley. One

118.

2 119.

Gard. Bull. Singapore 64(1) 2012

supposes Ridley, realizing his proposed name was already used for a

Caribbean species (Peperomia hispidula (Sw.) Dietr.), changed to P.

villosa without correcting the specimen. Unfortunately P. villosa is

also a later homonym, so I here provide an avowed substitute for it.

PIPER ARGYRITES Ridl. ex C.DC., Rec. Bot. Surv. India 6 (1912)

25; Suwanphakdee & Chantaranothai, Blumea 56 (2011) 239.

SYNTYPES: Peninsular Malaysia, Selangor, Gua Batu, H.N. Ridley

8176 (lectotype, designated by Suwanphakdee & Chantaranothai

(2011), SING; isolectotype, G); Ginding Bidai, H.N. Ridley 76/1.

Piper burkillii Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 189.

TYPE: Peninsular Malaysia, Negeri Sembilan, Tampin Hill,

LH. Burkill SFN 1177, 10 July 1916 (holotype, K (barcode no.

K000794927); isotype, SING (barcode no. SING 0068757)).

?= Piper curtisii C. DC.

Piper coactile Ridl., Journ. As. Soc. Mal. 1 (1923) 88. TYPE: Sumatra,

Berastagi, Karoo Plateau, 5000 ft, H.N. Ridley s.n., 15 February

1921 (holotype, K (barcode no. K000820249).

Piper cyrtostachys Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920)

189. TYPE: Peninsular Malaysia, Pahang, Gunong Berumbun near

Telom, H.N. Ridley 14013, November 1908 (holotype, K (barcode

no. K000794912)).

The Ridley collection number is the same as that for the type of Piper

subgrande Ridl., but the location and date are different.

Piper grande Ridl., Trans. Linn. Soc., Bot. 9 (1916) 112; non P. grande

Vahl (1798). Piper pergrande C. DC., Candollea 1 (1923) 268,

nom. nov. SYNTYPES: New Guinea, Mt Carstensz, Camp 1, C.B.

Kloss s.n., October-November 1912 (lectotype, designated by Chew

(1992), BM; isolectotype, K); Canoe Camp, 150 ft, C_B. Kloss s.n.,

October-November 1912 (BM, K).

= PIPER SUBBULLATUM K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee

(1900) 262; Chew, Blumea 37 (1992) 161.

Piper klossii Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 189.

TYPE: Peninsular Malaysia, Selangor, Gunong Menuang Gasing,

Ulu Langat C.B. Kloss s.n., February 1912 (holotype, K (barcode no.

K000794928)).

Ridley’s taxa among primitive angiosperms

Piper kraense Ridl., J. Fed. Malay States Mus. 10 (1920) 112, as

‘kraensis’. TYPE: Thailand, Tasan, C.B. Kloss 7045, Febrary—

March 1919 (holotype, K (barcode no. K000794895): isotype, SING

(barcode no. SING 0068765).

?= Piper majusculum Blume

124.

125. Piper kurzii Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 188.

SYNTYPES: Burma, Pegu, WS. Kurz 2220, 21 February 1871

(lectotype, designated here, K (barcode no. K000794423) (excluding

fruiting material in attached capsule (barcode no. K000794424)):

isotype, CAL). Perak, Temengoh, H.N. Ridley 14575, August 1909

(K); Selangor, Batu Caves, H.N. Ridley s.n.,4 March 1915 (K).

Given Ridley’s choice of epithet it seems appropriate to select the Kurz

specimen as lectotype. In the protologue, Ridley confused his Ks

and mistakenly referred to Kunstler as the collector of the lectotype

sheet. The Kew duplicate of this sheet has an attached capsule

containing one infructescence. This is labelled as Kurz 49/ and is

therefore specifically excluded from the lectotype as it represents a

different gathering. Piper kurzii C.DC. (Candollea | (1923) 198) is a

later homonym of Ridley’s species and probably represents the same

taxon as Kurz 2220 was cited (though pro parte — possibly explained

by inclusion of material of Kurz 49/) in the expanded description of

the species (Candollea 2 (1925) 205).

126. Piper melanocarpum Ridl., Journ. As. Soc. Mal. 1 (1923) 88. TYPE:

Sumatra, Berastagi, H.N. Ridley s.n., 11 February 1921 (holotype, K

(barcode no. K000820051)).

?= Piper nigrescens Blume

7. Piper montivagum Ridl., Trans. Linn. Soc., Bot. 9 (1916) 143. TYPE:

New Guinea, Mt Carstensz, Camps IX to XIII, 5500 to 10500 ft,

C.B. Kloss s.n. (holotype, BM (barcode no. BM000624276)).

= PIPER GiBBILIMBUM C. DC., Nova Guinea 8 (1910) 416; Johns in Johns

et al., Guide Alp. & Subalp. Fl. Mt. Jaya (2006) 401

128. Piper oreophilum Ridl., J. Straits Branch Roy. Asiat. Soc. 82 (1920) 188.

TYPE: Peninsular Malaysia, Pahang, Gunong Tahan in the gulley

on the way to the plateau about 4000 ft, H. N. Ridley 16106, August

1911 (holotype, K (barcode no. K000794911)).

254

13.0:

Sills

133:

134.

Gard. Bull. Singapore 64(1) 2012

Piper philodendron Ridl., J. Malayan Branch Roy. Asiat. Soc. 1 (1923)

87. TYPE: Sumatra, Berastagi woods, H.N. Ridley s.n., 14 February

1921 (holotype, K (barcode no. K000820057)).

Piper polycarpum Ridl., J. Fed. Malay States Mus. 10 (1920) 112, as

‘polycarpa’. TYPE: Thailand, Tasan, C.B. Kloss 6888 (holotype, K

(barcode no. K000794894)).

Piper puncticulatum Ridl., Trans. Linn. Soc., Bot. 9 (1916) 143. TYPE:

New Guinea, Mt Carstensz, Camp Vla, 3100 ft, C.B. Kloss s.n., 14

January 1913 (holotype, BM (barcode no. BM00062487); isotype,

K).

= PIPER GIBBILIMBUM C. DC., Nova Guinea 8 (1910) 416. Johns in Johns

et al., Guide Alp. & Subalp. Fl. Mt. Jaya (2006) 401.

Piper salticola Ridl., J. Malayan Branch Roy. Asiat. Soc. 1 (1923) 88.

SYNTYPES: Sumatra, Berastagi, H.N. Ridley s.n., 11 February 1921

(lectotype, designated here, K (barcode no. K00057307); Berastagi

forests, H.N. Ridley s.n., 13 February 1921 (K); Berastagi woods,

H.N. Ridley s.n., February 1921 (K).

? = Piper korthalsii Miq.

Piper subgrande Ridl., Fl. Malay Penins. 5 (1925) 329. TYPE: Peninsular

Malaysia, Perak, Ulu Batang Padang, H.N. Ridley 14013, November

1908 (holotype, K (barcode no. K000794913)).

?= Piper majusculum Blume

Note that the Ridley collection number is the same as that for the type of

Piper cyrtostachys Ridl., but the location and date are different.

Piper uncinulatum Ridl., J. Bot. 62 (1924) 300. TYPE: Peninsular

Malaysia, Pahang, Fraser’s Hill, 3000 ft, R.E. Holttum SFN 11343,

13 September 1923 (holotype, K (barcode no. K000794906)).

Piper venosum Ridl., Fl. Malay Penins. 5 (1925) 327, non P. venosum

(Miq.) C.DC. (1869). PipER ANGSIENSE I.M. Turner, nom. nov. TYPE:

Peninsular Malaysia, Negeri Sembilan, Gunong Angsi west side,

LH. Burkill 11510 (Md Nur leg.], 20 November 1923 (holotype, K).

Ridley’s name is a later homonym. In order to supply a legitimate name

for the species I propose an avowed substitute for Piper venosum

Ridl.

Ridley’s taxa among primitive angiosperms D5

XXIl. Piper subpenninerve (C. DC.) Ridl., Fl. Malay Penins. 3 (1924) 47.

Basionym: Piper nigrum var. subpenninerve C. DC., J. Asiat. Soc.

Bengal, Pt. 2, Nat. Hist. 75: 314.

ACKNOWLEDGEMENTS. The project was initiated with a small grant from the Singapore

Institute of Biology which is gratefully acknowledged. Support during the period of preparing

this paper has come from the Arnold Arboretum, Singapore Botanic Gardens, Forest Research

Institute Malaysia and Royal Botanic Gardens Kew. Particular thanks to Serena Lee and Paul

Leong (SING) for providing information and photographs. Dr. L. Gautier (G) kindly checked

for material of Piper argyrites in Geneva.

References

Bunchalee, P. & Chantharanothai, P. (2006) Notes on Polyalthia (Annonaceae). Thai Forest

Bull., Bot. 34: 1-3.

Candolle, C. de (1912) Piperaceae novae e peninsula Malayana. Rec. Bot. Surv. India 6: 1-27.

Chew, W.L. (1992) Studies in Malesian Piperaceae II. Blumea 37: 159-164.

Dandy, J.E. (1928) Malayan Magnolieae. Bull. Misc. Inform. Kew 1928(5): 183-193.

Fries, R.E. (1959) Annonaceae. Die natiirlichen Pflanzenfamilien, 2nd edition. Band 17a II:

1-171.

Johnson, D.M. (1989) Revision of Disepalum (Annonaceae). Brittonia 41: 356-378.

Kostermans, A.J.G.H. (1964) Bibliographia lauracearum. Bogor: Archipel.

Kostermans, A.J.G.H. (1970) Materials for a revision of Lauraceae III. Reinwardtia 8: 21-196.

MeNeill, J., Barrie, F. R., Burder, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K.,

Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J. (2006)

International Code of Botanical Nomenclature (Vienna code). Regnum Vegetabile 146:

1—S68.

Merrill, E.D. (1919) On the application of the generic name Melodorum of Loureiro. Philipp.

J ser., CG. 15: 125-137.

Mols, J.B. & KeBler, PJ.A. (2003) The genus Miliusa (Annonaceae) in the Austro-Malesia

Area. Blumea 48: 421-462.

Nooteboom, H.P. (1987) Notes on Magnoliaceae II Revision of Magnolia sections Maingola

(Malesian species), Aromadendron, and Blumiana. Blumea 32: 343-382.

Nurmawati, S. (2003) Malesian species of Dasymaschalon (Annonaceae). Floribunda 2(3):

67-81.

Saunders, R.M.K. (2003) A synopsis of Goniothalamus species (Annonaceae) in Peninsular

Malaysia, with a description of a new species. Bot. J. Linn. Soc. 142: 321-339.

Sinclair, J. (1951) Notes on Bornean Annonaceae. Sarawak Mus. J. 5: 597-609.

Smith, A.C. (1947) The families Illiciaceae and Schisandraceae. Sargentia 7: 1-224.

Soh, W.-K. (2011) Taxonomic revision of Cinnamomum (Lauraceae) in Borneo. Blumea 56:

241-261.

Stapf, O. (1907) Miscellaneous notes. Bull. Misc. Inform. Kew 1917: 197-198.

256 Gard. Bull. Singapore 64(1) 2012

Steenis-Kruseman, M.J. van (1950) Cyclopaedia of collectors. F/. Males., ser. 1, 1: 1-639.

Suwanphakdee, C. & Chantaranothai, P. (2011) A new species and three taxonomic changes in

Piper (Piperaceae) from Thailand. Blumea 56: 235-239.

Turner, I.M. (2000) The plant taxa of H.N. Ridley, 3. The Zingiberales. Asian J. Trop. Biol. 4:

1-47.

Turner, I.M. (2009a) Artabotrys (Annonaceae) in Borneo: new species and new synonyms.

Folia Malaysiana 10: 59-88.

Turner, I.M. (2009b) On the identity of Guatteria costata Hook.f. & Thomson (Annonaceae).

Kew Bull. 64: 577-578.

Turner, I.M. (2010) Richella in Malesia re-visited. Blumea 55: 118-119.

Turner, I.M. & Chin, S.C. (1998a) The plant taxa of H.N. Ridley, 1. The pteridophytes. Asian

J. Trop. Biol. 2: 31-40.

Turner, I.M. & Chin, S.C. (1998b) The plant taxa of H.N. Ridley, 1. The gymnosperms. Asian

J. Trop. Biol. 3: 33-35.

Turner, ILM. & Veldkamp, J.F. (2009) A history of Cananga (Annonaceae). Gard. Bull.

Singapore 61: 189-204.

Ueda, K. (1986) Comments on the uncertain application of Me/odorum Lour. (Annonaceae).

Taxon 35: 347-349.

Vink, W. (1970) The Winteraceae of the Old World I. Pseudowintera and Drimys — morphology

and taxonomy. Blumea 18: 225-354.

Warburg, O. (1897) Monographie der Myristicaceen. Abhandlungen der Kaiserlichen

Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher 68: \—680.

Weerasooriya, A.D. & Saunders, R.M.K. (2010) Monograph of Mitrephora (Annonaceae).

Syst. Bot. Monogr. 90: 1-167.

Gardens’ Bulletin Singapore 64(1): 257-269. 2012 257

Studies on Schismatoglottideae (Araceae) of Borneo XVII:

The Schismatoglottis Hottae Complex,

a new informal taxon, and three new species

from Sarawak, Malaysian Borneo

S.Y. Wong'”, P.C. Boyce’ and S.L. Low!

‘Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Samarahan, Sarawak, Malaysia

*sywong@frst.unimas.my

*Pusat Pengajian Sains Kajihayat (School of Biological Sciences),

Universiti Sains Malaysia 11800 USM, Pulau Pinang, Malaysia

ABSTRACT. On the basis of a suite of shared morphological characters, the Schismatoglottis

Hottae Complex is defined as a Borneo-endemic informal taxon in the Schismatoglottis

Asperata Group. Four species, three novel, are assigned to the Hottae Complex: S. hottae

Bogner & Nicolson, S. dilecta S.Y.Wong, P.C.Boyce & S.L.Low, sp. nov., S. mira S.Y.Wong,

P.C.Boyce & S.L.Low, sp. nov., and S. thelephora S.Y.Wong, P.C.Boyce & S.L.Low, sp. nov.

A key to species of the Hottae Complex is proffered. Schismatoglottis hottae is illustrated from

the Holotype herbarium material, the three novelties from living plants.

Keywords. Araceae, Borneo, Sarawak, Schismattoglottis, Schismatoglottis Hottae Complex

Introduction

Earlier papers of this series (Boyce & Wong 2006; Wong 2010), and Hay & Yuzammi

(2000), highlight that the Asperata Group (sensu Hay & Yuzammi 2000) is certainly

heterogeneous. Notwithstanding the phylogenetic veracity or otherwise of the

Asperata Group, during extensive fieldwork over the past 6 years in Sarawak we have

come to recognise that within the Asperata Group there exists a number of “species

complexes’—subsets of morphotaxa—the application of an informal nomenclature to

which provides a convenient tool for discussing potential phylogenetically significant

units (PSUs). One such is the Hottae Complex, defined here by long-persistent

somewhat bicarinate ligular sheaths, petioles and sometimes leaf blades with deciduous

indumentum, leaf blades abaxially with conspicuous pellucid secretory canals, solitary

inflorescences with a very short peduncle completely concealed within the subtending

cataphylls, staminate flowers with a blunt, narrowly pyramidal connective extended

well beyond the thecae, and a spathe limb darkening rapidly after opening and thence

deliquescing without crumbling. All species show an ecological preference for vertical

or very steep slopes with surface running water in lowland moist or perhumid gallery

forest, with one (S. thelephora sp. nov.) at least a facultative rheophyte.

258 Gard. Bull. Singapore 64(1) 2012

gap:

So far four species are known that display the above set of morphological characters:

S. hottae Bogner & Nicolson, and three novelties, here described.

Key to the Schismatoglottis Hottae Complex

la. Leaf blade lanceolate, abaxially with conspicuous pellucid secretory canals.

Staminate flower zone much narrower than the pistillate zone. Interstice staminodes

nipple-like, greatly exceeding the pistils and staminate flowers, upper-most

staminodes furnished with a terminal yellow-brown rostrum (vestigial anther?).

Anthers: deep red) i2..c.z.c:...c08t semester ete ee cee 4. S. thelephora

b. Leaf blade ovate to broadly ovate or ovato-cordate, abaxially with or without

conspicuous pellucid secretory canals. Staminate flower zone width equalling

or much exceeding the pistillate zone. Interstice staminodes not nipple-like,

equalling pistils. Anthers pale salmon-orange, or bright pink ..............eeeeeeeee 2

2a. Leaf blade adaxially minutely and softly densely tuberculate, abaxially often with

adventitious plantlets. Emerging leaves green. Pistillate zone much narrower than

the staminate zone, pale green. Staminate flowers pale brownish pink. Spathe

limb opening green. Sarawak (Bintulu, Bukit Satiam), shales ................ 3. S. mira

b. Leaf blade adaxially minutely pubescent or smooth, abaxially without adventitious

plantlets. Emerging leaves pink. Pistillate zone spadix sub-equalling or slightly

exceeding the staminate zone. Spathe limb opening medium to bright pink.

Sarawak atid Brome ..s.0..:scedecccesagesnee emt ceuet cece terete ween ene ee ee 3

3a. Plant to 35 cm tall. Leaf blade up to 30 cm x 20 cm, adaxially smooth, semi-

glossy when wet, abaxially pale green with conspicuous pellucid secretory

canals. Petioles, and midrib and primary lateral veins of abaxial surface sub-

microscopically pale greyish pubescent. Spadix c. 9 cm long. Sarawak (Mulu),

S) 02) eke PP ee eR Re oe a tae ees i once eeseda noaccuceeness 2. S. dilecta

b. Plant to 20 cm tall, often less. Leaf blade up to 12 cm x 9 cm, adaxially densely

minutely hairy, matte even when wet, abaxially pale green without conspicuous

pellucid secretory canals. Petioles, and mid-rib and primary lateral veins of

abaxial surface densely pale brown pubescent. Spadix c. 2.5 cm long. Brunet

(Teraja and Batu: Patam);isandstonesiz: 2.20) ere. en eee 1. S. hottae

1. Schismatoglottis hottae Bogner & Nicolson, Aroideana 2: 120 (1979); Mayo,

Bogner & Boyce, The Genera of Araceae, 184, pl. 49(i1), D (1997); Hay & Yuzammi,

Telopea 9(1): 70-71 (2000) — Schismatoglottis cordifolia M. Hotta, Mem. Coll. Sci.

Univ. Kyoto, ser. B, 32: 229 (1966) (non S. cordifolia Ridl. (1911)). TYPE: Brunei,

‘Seria Distr.’ [Belait], Bukit Teraja, Teraja Forest Reserve, 21 Dec 1963, M. Hotta

12886 (holo KYO, K! photo). (Fig. 1)

Schismatoglottis Hottae Complex and new Sarawak species 259

Small lithophytic herb to 20 cm tall. Stem very condensed and shortly erect, c. 3 em

long x 1 cm diam., older portions clothed in old leaf remains, with oldest portions

naked and somewhat decumbent; modules pleionanthic. Leaves several together (4—

7) with roots emerging from among their bases; petiole 5-9 cm long, slender, about

equalling the blade, very densely clothed in minute straight pale brown hairs, sheathing

only at the extreme base with the sheath extended into a narrowly triangular, abaxially

pubescent, bicarinate ligular portion 24 cm long; blade ovate to broadly ovate,

matte dark green, slightly metallic and densely minutely hairy throughout adaxially,

matte pale green, densely minutely hairy on all the venation and minutely tuberculate

between the veins abaxially, drying dark brown, 9—12 cm long x 7—9 cm wide, the base

retuse to cordate with rounded posterior lobes to 2 cm long, the apex broadly acute to

obtuse and shortly apiculate; midrib not or hardly prominent abaxially and adaxially;

primary lateral veins crowded, c. 13 on each side of the midrib, diverging at 45—60°,

alternating with lesser interprimaries and very occasionally branching near the midrib,

impressed adaxially, prominent abaxially; secondary veins abaxially prominent (due

to hairs), arising from the midrib and the bases of the primary veins; tertiary venation

obscure. Inflorescence solitary; peduncle very short, completely concealed within

subtending leaf bases. Spathe c. 3.5 cm long, green to salmon pink and then veined

with deeper pink; lower spathe ovoid, c. 1.1 cm long, differentiated from the limb by

a slight constriction; limb 2.2—2.8 cm long, broadly lanceolate, apiculate for 2—3 mm,

?deciduous. Spadix 2.5 cm long, sessile; pistillate zone obliquely inserted, c. 3 mm

long; pistils densely packed; ovary sub-globose, c. | mm diam.; stigma sessile, thickly

discoid, almost as wide as the ovary; sterile interstice slightly wider than male and

female zones, 2 mm long, c. 3 whorls of flat-topped irregularly polygonal staminodes

c. 0.6 mm diam.; staminate flower zone cylindric, slightly attenuate corresponding to

spathe constriction, c. 3 mm long; stamens somewhat laxly arranged, mostly in adnate

pairs with the blunt and narrowly pyramidal connective extended beyond the thecae;

appendix cylindric-ellipsoid, more than half the length of the spadix, about twice the

width of the male zone, c. 2 cm long x 6 mm thick, composed of flat-topped irregularly

polygonal staminodes c. | mm diam. Infructescence unknown.

Other specimens examined: BRUNEI: Belait District: Melilas Subdistr., Ulu Ingei, Bukit Batu

Patam, 04°05’N 114°42’E, 9 June 1989, P.-C. Boyce et al. 279 (BRUN, K, L); Labi Subdistr.,

Bukit Teraja, south of summit, 04°20’N 114°27°E, 20 March 1991, R.J.Johns 6872 (BRUN,

K).

Distribution. Brunei, to date recorded only from Bukit Teraja and Bukit Batu Patam.

Ecology. Lithophytic on sandstone boulders and low damp cliffs in lowland kerangas

and humid hill dipterocarp forest, 180-400 m alt.

Notes. Hay & Yuzammi (2000) remarked that S. hottae appeared allied to S. puberulipes

Alderw. by sharing velvety hairs on the leaf, a solitary short-pedunculate inflorescence,

and an ‘inflated’ appendix. However, there is confusion with regard application of

260 Gard. Bull. Singapore 64(1) 2012

cael

Wu OOM

KYO 00020228

NATIONAL HERBARIUM OF N'S.W

Schismatogloitis hottae Bogner & Nicolson

DET. A. Hay Dee 1999

HOLOTYPE

Schismatoglottis hottae Bogner et Nicolson

rom. nov.

[ Schismatoglottis cordifolia M.Hotta 1366

non Ridley 4310 |

che. oe Biever. uae ris “TYP US!

HERBARIUM UN S KYOTOENSIS

PLANTS OF BRUNEI (UM UNIVERSITATIS KYOTOENSIS

Kyoto Univ. Borneo Expedition, 1963 4 = ; :

Were M Hella

rx BPUADA O& Le potas

Loc, Bt. Teraja, Teraja forest reserve,

Seria district, alt. 300-400 n.

Date Dee, 21, 1963.

Coll. Mitsuru Horra no. 72866

Fig. 1. Schismatoglottis hottae Bogner & Nicolson. Holotype: M.Hotta 12886 (KYO). Image

courtesy of Dr. Hidetoshi Nagamasu (KYO). Used with permission.

Schismatoglottis Hottae Complex and new Sarawak species 261

the name S. puberulipes. The plant illustrated in Hay & Yuzammi (2000:79, Fig. 9),

and the substantial part of the description given is Schismatoglottis hayi S.Y.Wong

& P.C.Boyce endemic to Gua Niah (NE Sarawak) and allied to Mulu-endemic S.

multinervia, sharing with that species aromatic (terpenoids) vegetative tissues, fully

adnate petiolar sheaths, and a spathe limb rapidly darkening on opening but then

breaking into strips. Together with several other novelties, these are regarded by us

as belonging to the Multinervia Complex, and are most probably not closely allied to

the Hottae Complex. The lectotype of S. puberulipes (BOKR! + BO spirit!) matches

exactly S. gamoandra M.Hotta, a very distinctive species of doubtful affinity (Wong

& Boyce 2011).

2. Schismatoglottis dilecta S.Y.Wong, P.C.Boyce & S.L.Low, sp. nov., Schismatoglottis

mirae accedens, innovationibus clare roseis, laminis adaxialiter seminitidis,

abaxialiter canalis secretoriis pellucidis conspicuis venis similibus, sine proliferis

adventiis, zona pistillata spadicis aliis zonis in latitudine aequali spatha rosea differt.

TYPE: Malaysia, Sarawak, Limbang, Nanga Medamit, Mulu N.P., Melinau Gorge, 3

Oct. 2007, P.-C. Boyce et al. AR- 2312 (holo SAR!). (Fig. 2)

Moderately robust usually lithophytic herb to 35 cm tall. Stem stout, condensed,

up c. 15 cm long x 2 cm diam., active portion erect, completely obscured by leaf

bases, oldest parts tending to become decumbent, naked and ringed with conspicuous

cataphyll/prophyll and petiole scars; shoot modules pleionanthic. Leaves several to

rather few together (4-8) with roots emerging from among their bases, innovations

pink; petiole 10-25 cm long, stout, about equalling the blade, sub-microscopically

pale greyish pubescent, appearing matte greyish-green when dry and minutely asperate

when wet, sheathing only at the extreme base, the majority of the sheath forming a

narrowly triangular, bicarinate, somewhat fleshy ligular portion 3—5 cm long; blade

ovate to ovate-elliptic, semi-glossy slightly metallic dark green adaxially, much paler

dull lime green abaxially, blades drying medium brown, 10-20 cm long = 6—15 cm

wide, the base cordate with rounded posterior lobes to 3 cm long, the apex broadly

acute shortly apiculate; midrib prominent abaxially, deeply impressed adaxially,

silvery when living; primary lateral veins crowded, c. 15 on each side of the midrib,

diverging at 45—60°, irregularly alternating with only slightly lesser interprimaries and

very occasionally branching near the midrib, deeply impressed adaxially, prominent

abaxially, sub-microscopically pale greyish pubescent; primary lateral veins sub-

microscopically pale greyish pubescent and raised abaxially; interprimary veins

interspersed with vein-like pellucid glands abaxially; secondary and tertiary almost

invisible. Inflorescence solitary; peduncle very short, together with the lowermost

part of spathe completely concealed within subtending cataphyll bases. Spathe c.

8.5 cm long, peduncle and lower part of bright pink, spathe limb much paler pink,

veined with deeper pink; lower spathe cylindrical, c. 3 cm long, differentiated from the

cm long, very broadly lanceolate, apiculate for c. 4 mm, opening clear pale pink but

DED ;

262 Gard. Bull. Singapore 64(1) 2012

Fig. 2. Schismatoglottis dilecta S.Y.Wong, P.C.Boyce & S.L.Low. A. Plants in habitat; note the

semi-glossy leaf blades. B. Leaf blade, abaxial view. C. Plant with emerging inflorescence. Note

the long, persistent, fleshy ligules. D. Spadix at pistillate anthesis, spathe artificially removed.

E. Detail of interstice staminodes (below) and staminate flowers (above). F. Detail of appendix

staminodes. G. Post anthesis inflorescence. Note that the spathe limb has deliquesced and that

the lower spathe persists. All from P.C. Boyce et al. AR- 2312. Photo credits: Peter C. Boyce.

Schismatoglottis Hottae Complex and new Sarawak species 263

swiftly darkening during staminate anthesis and then soon deliquescent. Spadix stout,

8 cm long, sessile: pistillate zone very slightly conic-cylindrical, weakly obliquely

inserted, c. 1.5 cm long; pistils densely packed, medium pink: ovary sub-globose, c.

1.5 mm diam.; stigma sessile, thickly discoid, as wide as the ovary: sterile interstice

very slightly wider than pistillate and staminate zones, 0.5 cm long. c. 7 whorls of flat-

topped irregularly polygonal glossy pale pink staminodes c. 1 mm diam.; staminate

flower zone cylindric, c. 1.5 cm long, often with a few scattered groups staminodes

of similar form to those in the interstice; stamens very densely arranged, mostly in

adnate pairs with the blunt and narrowly oblong connective extended beyond the

thecae, anthers medium pink; appendix stoutly clavate-cylindric, c. 2/3 the length of

the spadix, exceeding the width of the staminate zone, dull pale pink, c. 4.5 cm long

x 10 mm thick, composed of very densely arranged flat-topped irregularly polygonal

staminodes c. | mm diam. Infructescence unknown.

Other specimens examined: Malaysia, Sarawak, Miri, Marudi, Long Lama, Mulu N_P., trail

to Long Lansat, Sungai Licat, 04°00°03.5°N 114°48°49.87E, 9 Aug. 2006, P.-C. Boyce et al.,

AR- 1974 (SAR).

Distribution. Mulu N.P., found so far in the extreme SW and NE parts of the park.

Ecology. Lowland moist gallery forest, restricted to vertical walls of low shale cliffs

above forest streams, mostly on red clays with leaf litter accumulated around the bases,

more rarely on almost bare shale with surface seepage moisture: c. 30 m asl.

Notes. Schismatoglottis dilecta is superficially very similar to S. mira (see below):

from a distance plants of both are virtually indistinguishable. Aside from the floral

characteristics noted above S. dilecta differs from S. mira by several vegetative

features, including the leaf blade adaxial surface texture (smooth vs papillose), the

presence abaxially of vein-like pellucid secretory veins, and an absence of adventitious

plantlets (S. dilecta). The form and texture of the petiolar sheath ligules also differs

(compare Fig. 2C & 3C).

Etymology. From dilectos (Latin) “lovely, esteemed”, in allusion to the highly attractive

foliage, and striking colour of the inflorescences.

3. Schismatoglottis mira S.Y.Wong, P.C.Boyce & S.L.Low, sp. nov.. Schismatoglottis

dilectae similis, innovationibus viridibus (non roseis) lamina adaxialiter minute et

molliter dense tuberculata, abaxialiter proliferis adventitiis multis, zona pistillata

spadicis aliis zonis multo angustiore, et spatha viridis facile distinguenda. TYPE:

Malaysia, Sarawak, Bintulu, Bukit Satiam, 02°59°07.4°N 112°55°47.0°E, 15 July

2006, P.C._Boyce et al. AR-1906 (holo SAR!). (Fig. 3)

264 Gard. Bull. Singapore 64(1) 2012

Moderately robust lithophytic herb to 45 cm tall. Stem stout, condensed, up c. 10 cm

long x 2.5 cm diam., active portion erect, completely obscured by leaf bases, oldest

portions usually naked and tending to be decumbent; shoot modules pleionanthic.

Leaves several to rather few together (3—7) with roots emerging from among their bases,

innovations very pale green; petiole 10-25 cm long, stout, very weakly D-shaped in

cross-section, sub-equalling the blade, minutely asperate, matte pale-green, sheathing

only at the extreme base, the greater part of the sheath forming a narrowly triangular

very weakly bicarinate, membranous ligular portion 2—7 cm long; blade broadly ovate,

carried sub-perpendicular to petiole, adaxially minutely and softly densely tuberculate,

abaxially often with adventitious plantlets, adaxially deep green with the midrib marked

by a somewhat irregular broad pale yellow band, abaxially pale grey-green with the

epidermal cells somewhat refractive, blades drying pale brown, 15—25 cm long x 10—

21 cm wide, the base cordate with rounded posterior lobes to 4 cm long, the apex blunt,

shortly apiculate; midrib prominent abaxially, somewhat impressed adaxially, pale

yellow when living; primary lateral veins crowded, c. 15 on each side of the midrib,

diverging at 45—60°, irregularly alternating with only slightly lesser interprimaries and

very occasionally branching near the midrib, slightly impressed adaxially, prominent

abaxially; interprimary veins interspersed; secondary and tertiary almost invisible.

Inflorescence solitary; peduncle very short, completely concealed within subtending

cataphyll bases. Spathe c. 9 cm long, spathe exterior medium somewhat glaucous

green, longitudinally veined deeper green; lower spathe cylindrical, c. 3.5 cm long, not

clearly differentiated from the spathe limb; limb 6—6.5 cm long, broadly funnel-form

lanceolate at pistillate anthesis, the terminal portion tending to remain weakly furled,

and apiculate for c. 2 mm, opening pale green internally but swiftly darkening to glossy

tan during staminate anthesis, and then rapidly deliquescent starting from the spathe

limb margins. Spadix stout, sub-equalling the spathe, c. 8.5 cm long, sessile; pistillate

zone cylindrical, much narrower than rest of spadix, somewhat obliquely inserted, c.

1.5 cm long; pistils rather densely packed, pale green; ovary sub-globose, c. 1.5 mm

diam., interspersed with a few prominent, stoutly clavate white staminodes; stigma

sessile, thickly discoid, almost as wide as the ovary; sterile interstice very slightly

wider than pistillate and staminate zones, c. 5 mm long, with c. 6 whorls of somewhat

rounded-topped irregularly polygonal staminodes c. | mm diam., lowermost whorls

glossy very pale yellow, upper whorls pale salmon-orange; staminate flower zone

cylindric, c. | cm long; stamens moderately densely arranged, mostly in adnate pairs

with the bluntly and narrowly oblong connective barely extended beyond the thecae,

anthers pale salmon-orange; appendix stoutly cylindric, c. 2/3 the length of the spadix,

equalling the width of the staminate zone, dull pale salmon-orange, c. 6 cm long =

10 mm thick, composed of very densely arranged flat-topped irregularly polygonal

staminodes c. | mm diam. Infructescence unknown.

Other specimens examined: Malaysia, Sarawak, Bintulu, Bukit Satiam, 02°59°26.1°N

112°55’54.4”, 11 Aug. 2004, P-C.Boyce & Jeland ak Kisai AR- 597 (SAR), AR-603 (SAR),

AR-618 (SAR); Bukit Satiam, 02°59713.3"N 112°55°57.5”E, 14 July 2006, P.C.Boyce et al.

AR-1888 (SAR); 02°59’07.4”N 112°55’47.0”E; 15 July 2006 P.-C. Boyce et al. AR-1906 (SAR).

Fig. 3. Schismatoglottis mira S.Y.Wong, P.C.Boyce & S.L.Low. A. Plants in habitat; n

matte, tuberculate leaf blades. B. Leaf blade, adaxial view to show the tuberculate surfac

Details of long, persistent, membranous ligules. D. Inflorescence at pistillate anthesis. Note

that the spathe limb has already darkened and is starting to deliquesce along the margins. E.

Detail of fertile portions of spadix and lower half of appendix: spathe artificially removed. F.

Detail of appendix staminodes. All from P.-C. Boyce et al. AR-1906. Photo credits: Peter C

Boyce.

266 Gard. Bull. Singapore 64(1) 2012

Distribution. Known only from Bukit Satiam, Bintulu.

Ecology. Vertical shale cliffs and margins of waterfalls in lowland moist evergreen

forest; 100-120 m asl.

Notes. A strikingly beautiful species with an extraordinary leaf blade texture that is so

far unique in the genus, although somewhat approached by S. hottae. The superficially

similar appearance to S. dilecta is remarkable, although the species differ in numerous

critical morphologies.

Etymology. Latin, mirus (wonderful) in reference to the appearance of the plant, and

the unique texture of the adaxial surface of the leaf blade.

4. Schismatoglottis thelephora S.Y.Wong, P.C.Boyce & S.L.Low, sp. nov., 4b

omnibus speciebus complexi Hottae foliorum laminis lanceolatis, zona staminata

pistillata multo angustiore, staminodiis papilliformibus pistillis multo excedentibus,

floribus staminatis rubrissimis differt. TYPE: Malaysia, Sarawak, Bintulu, Tatau,

GT Plantations, Sungai Pandan Kecil, trail behind Camp C to intake point for water

supply-pipe, 02°42°40.1°N 113° 20’ 37.9"E, 5 Sept. 2010, PC Boyce, S.¥Y.Wong &

S.L.Low AR-3082 (holo SAR!). (Fig. 4)

Small facultative rheophytic herb to 25 cm tall, but usually somewhat less. Stem rather

stout, condensed, creeping and rooting strongly, up c. 25 cm long x 2.5 cm diam.,

active portion completely obscured by leaf bases; shoot modules pleionanthic. Leaves

several together (5—11), with roots emerging from among their bases, innovations

very pale green; petiole 7-17 cm long, relatively stout, very weakly D-shaped

in cross-section, sub-equalling to rarely exceeding the blade, minutely asperate,

matte medium-green, sheathing only at the extreme base, the majority of the sheath

forming a narrowly triangular, bicarinate, fleshy ligular portion 1.5—3 cm long; blade

oblanceolate to lanceolate or very narrowly ovate, thick to sub-succulent, 7-20 cm

long x 2-8 cm wide, the base acute to cuneate or weakly obtuse, tip acuminate for 1—3

cm and then conspicuously tubular-mucronate; adaxially semi-glossy medium green,

abaxially paler with the epidermal cells somewhat refractive and scattered vein-line

pellucid secretory canals, blades drying straw yellow; midrib and primary veins tough

and abaxially prominent, adaxially very weakly impressed; primary lateral veins 3—9

on each side of the midrib, irregularly alternating with interprimaries ill-differentiated

from the secondary venation, diverging at c. 30°; secondary venation mostly arising

from the midrib, some from near the bases of the primary veins in the lower part of the

leaf; tertiary venation obscure. Inflorescence solitary; peduncle very short, completely

concealed within subtending cataphyll bases. Spathe c. 3.5 cm long, spathe exterior

medium green, longitudinally veined deeper green; lower spathe ovoid, c. 1.5 cm long,

very pale greenish-white, with a shallow, broad constriction differentiating it from the

spathe limb; limb c. 2 cm long, broadly lanceolate, apiculate for c. 2 mm, opening pale

Schismatoglottis Hottae Complex and new Sarawak species 267

Fig. 4. Schismatoglottis thelephora S.Y.Wong, P.C.Boyce & S.L.Low. A. Plants in habitat.

B. Leaf blade, adaxial view to show vein-like pellucid secretary canals. C. Details of long,

persistent, membranous ligules. D. Leaf-tip tubule. E. Emerging inflorescence. F. Spadix at

staminate anthesis, spathe artificially removed. G. Detail of fertile portions of spadix and lower

half of appendix; spathe artificially removed. The nipple-like staminodes are prominent. Note,

too, the slender strings of pollen. All from P.-C. Boyce, S.¥. Wong & S.L. Low AR-3082. Photo

credits: Peter C. Boyce.

268 Gard. Bull. Singapore 64(1) 2012

green but swiftly darkening to dark glossy brown during staminate anthesis, and then

rapidly deliquescent starting from the spathe limb margins. Spadix somewhat stout,

sub-equalling spathe, c. 3 cm long, sessile; pistillate zone weakly conic-cylindrical,

somewhat obliquely inserted, c. 5 mm long; pistils rather densely packed, white; ovary

rhombic-subglobose, c. 0.5 mm diam.; stigma sessile, discoid, smaller than the ovary

diam.; sterile interstice very slightly narrower than pistillate zone, c. 5 mm long, c.

6 whorls of elongated nipple-like staminodes, these up to twice as long as pistils,

and with at least a few of these interspersed with the upper pistillate flowers, waxy

white with a yellow-brown tip perhaps associated with a vestigial connective, each c.

2 mm long and 0.3 mm diam.; staminate flower zone cylindric, c. 4 mm long; stamens

rather loosely arranged, mostly in adnate pairs with the blunt and narrowly triangular

connective extended beyond the thecae, deep yellow with the anthers (thecae) deep

red, lowermost staminate flowers transitioning to staminodes, these dull yellow barely

tinged red, these then transitioning to the elongated white staminodes present on the

interstice; appendix bullet-shaped, c. 2 the length of the spadix, much wider than the

staminate zone, dark yellow, c. 2 cm long x 6 mm thick, composed of rather loosely

arranged flat-topped irregularly polygonal staminodes c. 0.7 mm diam. Infructescence

subtended by the fleshy persistent lower spathe, rather loosely urceolate.

Other specimens examined: Malaysia, Sarawak, Bintulu, Tatau, GT Plantations, Sungai Likau,

02°44°37.6”"N 113°26°10.5”E, 6 Sept. 2010, PC. Boyce, S. Y. Wong & S.L.Low AR- 3092 (SAR).

Distribution. Known only from the neighbourhood of the type locality.

Ecology. Rheophytic along the lower fringes of lowland perhumid gallery forest on

sandstones and sandstone-derived clays; 50-70 m asl.

Notes. Schismatoglottis thelephora is remarkable, indeed unique, in the genus by the

greatly elongated nipple-like staminodes that clothe the interstice between the pistillate

and staminate flower zones, and also occur scattered among the uppermost pistillate

flowers. The deep red anthers are distinct from all other species in the Hottae Complex,

and furthermore this colour has not been previously recorded for the genus. Pollen is

released in very slender white strings.

Etymology. From Greek thele-, a nipple, and Latin -phora, to bear, in allusion to the

staminodes separating the pistillate and staminate flower zones.

ACKNOWLEDGEMENTS. Research and fieldwork in Borneo was conducted, most

recently, under Sarawak Forestry Department Research Permit Nos. NPW.907.4.4(V)-77

& NCCD.907.4.4(Jld.VI)-56 & Park Permit Nos. 34/20010 & 27/2011. The continuing

collaboration and support of the Forest Department Sarawak, and Sarawak Forestry Corporation,

are gratefully acknowledged. We would like to thank Mr. Wong Kee Heng, Mr. Wilfred Rogers

and Mr. Serum Ungau of GT Plantation Sdn. Bhd. for facilitating our field trip at Sg. Kakus,

Bintulu.

Schismatoglottis Hottae Complex and new Sarawak species 269

References

Boyce, P.C. & Wong, S.Y. (2006) Studies on Schismatoglottideae (Araceae) of Borneo I: A trio

of new Schismatoglottis from Sarawak. Gard. Bull. Singapore 58: 7-18.

Hay, A. & Yuzammi (2000) Schismatoglottideae (Araceae) in Malesia I — Schismatoglottis.

Telopea 9(1): 1-177.

Wong, S.Y. (2010) Studies on Schismatoglottideae (Araceae) of Borneo XIII: A Revision of the

Schismatoglottis nervosa species complex. Gard. Bull. Singapore 62(1): 177-209.

Wong, S.Y. & Boyce. P.C. (2011) Studies on Schismatoglottideae (Araceae) of Borneo XVI: A

New Species and a New Informal Taxon (the Multinervia Complex) of Schismatoglottis

from Sarawak. Acta Phytotax. Geobot. 60(3): 131-137.

Gardens’ Bulletin Singapore 64(1): 271-272. 2012 D/A

BOOK REVIEW. Heritage Trees of Penang. Simon Gardner, Pindar Sidisunthorn

& Lai Ee May. 2011.

Penang, Malaysia: Areca Books. 27.3 cm x 21.7 cm, hard cover. 397 pp. ISBN 978-967-5719-

06-6. Price RM 100.

This is a superb piece of work. The book weaves strands of history, botany, culture

and the visual arts into a rich tapestry on the subject of the heritage trees of Penang.

The book will appeal to the informed layperson as well to professionals working with

plants. The well-thought through layouts of the pages successfully combine stunning

photographs and vivid botanical illustrations with just the right amount of text so that

almost every page that one flips through is a delight to the eyes. Perhaps an unintended

consequence of this is the relatively small font size for the text.

The professional will find that the book is well researched with up-to-date

botanical names. The work takes into cognizance relatively recent changes in the

family-level taxonomy of the trees treated. A prelude to the scholarship that the reader

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under the heading of “Orthography and Spelling” the authors set out how the Malay

names of trees and places are treated in this work in English.

As an entrée before embarking on a sylvan odyssey, the introductory chapter

presents convincing arguments on why Penang’s urban and natural forests are an

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Information on the history of tree planting, the Esplanade and the Botanic

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Under “Forest Trees” there is a simplification. In the first paragraph of page 24, the

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An interesting brief introduction to the inland forests of Penang is provided.

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order. The format for each species begins with the botanical name followed by

vernacular names, general comments, a section on uses, and a section on botany.

For the botanical names, the describing authorities have been omitted. The general

comments are very useful in highlighting exemplary specimens of the species treated

and provide well-researched and interesting information on these individual trees. One

of the key features of this book is that the authors have painstakingly provided stunning

photographs illustrating the habits, trunks, fruits and flowers of the trees treated. These

DGD. Gard. Bull. Singapore 64(1) 2012

are augmented by exquisite botanical illustrations usually of the flowers and leaves.

The magnificent trees of the forest dominate with 49 species treated under

their section. The photographs and illustrations in the section are among the best in the

book. On page 275, together with the flowers of Callerya atropurpurea, a photograph

is presented of a pair of perfectly camouflaged caterpillars which are dark purple in

colour with a cream stripe, just like the flowers of this tree!

In conclusion, I think that that the book has succeeded through words,

photographs and botanical illustrations in drawing attention to these most important

assets that Penang has in its heritage trees, urban forests and natural forests.

S.K. Ganesan

Centre for Urban Greenery & Ecology (Research),

National Parks Board

Gardens’ Bulletin Singapore 64(1): 273-275. 2012 273

BOOK REVIEW. The Wild Orchids of Hong Kong. G. Baretto, PJ. Cribb & S.

Gale. 2011.

Kota Kinabalu: Natural History Publications (Borneo), in association with Kadoorie Farm &

Botanic Garden. 26.3 x 19 cm, hard cover. 697 pp. ISBN 978—983-812—136—1. Price RM 240.

Hong Kong, situated on the coast of south-east China just south of the Tropic of Cancer,

is best known as a modern city of high-rise buildings and as an economic and financial

hub in the region. Much less known are its comparatively rich natural resources, of

which the orchids form an interesting part.

A large part of Hong Kong is mountainous with rugged peaks and gorges, and

although there has been a very long history of deforestation, the forest cover is still a

surprising 13 % (although now mainly found at higher elevations). In addition, much

of the countryside is regenerated secondary forest. Grassland is a human-induced

environment (in many cases through accidental hill fire) and is also a widespread

vegetation type in Hong Kong, accounting for 14% of the total land area. The small

territory of some 1100 km? is home to 126 species and varieties of orchids which are

placed in 58 genera, and this is a remarkably high number for a small area situated

near the northern limit of tropical South-east Asia. In fact, 9% of all orchids of China

are found in Hong Kong, although this small territory only amounts to 0.01% of its

land area. About two thirds of native Hong Kong orchids (83 taxa) are terrestrial,

while the remainder (43) are primarily epiphytic or lithophytic. The native orchids are

most abundant on the north-facing slopes and in ravines above 300 m. Affinities to

the orchid floras of southern China and northern Indochina are obviously the greatest,

while a few species are more widespread. Five species from temperate Asia are also

found in Hong Kong. In addition, eight taxa are currently considered endemic to Hong

Kong. Several of Hong Kong’s orchids have been decimated in nature to such an

extent that they can now be considered as being at the risk of extinction in the wild.

In view of this, conservation aspects are of particular importance in the present book.

The book is a complete and richly illustrated enumeration of all orchid species

recorded in Hong Kong so far. It brings to fruition the work of Hong Kong resident

Gloria Barretto (1916-2007) who, in over 40 years of painstaking and dedicated work,

produced draft descriptions of all orchids of Hong Kong she had intended to use for

a comprehensive treatment of the native orchids. Sadly, Gloria never saw the finished

result of her studies, and co-authors Drs. Phillip Cribb (former curator of the Orchid

Herbarium, Royal Botanic Gardens Kew, London, UK) and Stephan Gale (Kadoorie

Farm and Botanic Garden, Hong Kong) concluded her work to publish this book.

An introductory part on more general aspects of the territory of Hong Kong

and its orchid flora forms the first 74 pages of the book. A detailed and informative

coloured map is provided in Fig. | (unfortunately rather difficult to read, and a black-

and-white map labelled in larger font would probably be clearer; but in any case this

is the only shortcoming of the book noted here). Chapters deal with the geology,

topography, climate and vegetation of Hong Kong. A brief account of the history of

exploration of the orchid flora is given, ranging from the early collections made in

the 1830s and 1840s to the recent work by Gloria Barretto. As is commonly found in

274 Gard. Bull. Singapore 64(1) 2012

publications of this kind, a description of the structure of vegetative and reproductive

parts of the orchids is given, and their life cycle is briefly explained. A chapter is

devoted to the cultivation of the orchids of Hong Kong which will be of special

interest to many readers, given the fact that the cultivation of orchids in China dates

back 2500 years to pre-Confucian times. Careful plant selection, mounting, potting,

watering, fertilising as well as pests and diseases are explained and illustrated by

photographs. The affinities of the orchid flora of Hong Kong are discussed, and a list

of the species found in Guangdong Province (the Chinese province directly adjacent

to Hong Kong) but not yet recorded in Hong Kong is given—t is estimated that some

of these may be found inside the territory as well one day. In view of the decline of

the orchids as a consequence of human activities, the discussion of various aspects

of conservation 1s receiving particular attention. Threats to the continued survival of

the orchids are various, and include man-made habitat destruction as a consequence

of urban development and accidental hill fire, and illegal collecting by plant thieves.

Effects of climate change are expected to have an additional adverse effect on the

orchids as well. Natural phenomena like extreme weather conditions play a role too,

and add to the man-made threats. The Hong Kong government has recognised the need

for conservation of the natural environment, and has set aside about 40% of the total

land area of the territory as ‘Country Parks’. The comprehensive legal framework with

its several ordinances is here well explained to the reader. On the basis of previously

made herbarium collections, reputable field notes, contemporary orchid surveys and

an estimation of threat levels, conservation assessments are made for all of the native

orchids of Hong Kong. Currently, nearly three quarters of all orchid species native in

Hong Kong are at risk of extinction in the wild and fall into the Vulnerable, Endangered

or Critically Endangered categories (IUCN 2001). Seven species have not been seen

for several decades and are therefore presumed extinct. In addition to conservation

assessments using the IUCN criteria, an easy-to-use Hong Kong-centric system was

applied where relative rarity (based on recent sightings) and distribution were used.

It is estimated that this system would be more meaningful to land managers than a

more scientific approach based on the [UCN criteria. One of the basic requirements of

plant conservation in general is a good knowledge of the plant group concerned, and

the authors of the present book have certainly succeeded in making comprehensive

information available on the native orchids of Hong Kong.

The main part of the book consists of the genus and species accounts, with a

short introduction on orchid classification and identification keys to subfamilies and

genera. Genera are arranged in phylogenetic order, 1.e., after groups of related ones,

which is much more meaningful and easy-to-use than an alphabetic arrangement. For

all accepted genera and generic synonyms, the name and place of description are cited,

together with their typification. Following the generic description, the distribution of

the genus and the number of its species (worldwide and in Hong Kong) are given. The

derivation of the genus name is explained. This is followed by the treatment of the one

to several species. An identification key is provided in all cases where more than one

species is found in Hong Kong. Each species account starts with historical notes on the

discovery, and gives other relevant information on identity and nomenclatural history.

Book Review a5

The currently accepted name and the relevant synonyms are cited, together with

references, type specimens and the acronym of the herbarium where these are kept.

Species descriptions are very detailed and are mostly based on Hong Kong material

(rarely on material from adjacent Chinese provinces). For all species the worldwide

distribution, habitat and occurrence in Hong Kong, phenology, vernacular names

and the derivation of the species epithet are given. Detailed notes on conservation

issues, area of occupancy and abundance are provided, and the category of relative

rarity is displayed (colour-coded for easy reference). It is emphasised that the authors

have on purpose omitted distribution maps to discourage theft of orchids which is

still a significant threat. The species accounts are generously illustrated. One to three

black-and-white line drawings are provided for each species, showing the habit of

the plant as well as a dissection of the flower. Excellent high-quality photographs are

reproduced in nearly all taxa (one to few per taxon) and show the plant, leaf characters

(particularly in the ‘Jewel Orchids’) as well as floral details. Most of these photographs

were taken in Hong Kong. In addition, fine colour paintings by General John Eyre are

also reproduced to illustrate 25 species.

A comprehensive bibliography list and a detailed glossary follow the main

text. Three appendices are provided. Many readers will find Appendix I very useful,

containing brief biographies of plant collectors and botanists who are associated

with Hong Kong since the 18th century. Appendices II and III are assessments of the

conservation status of all native orchids (based on IUCN criteria and relative rarity in

a Hong Kong-centric approach, respectively). Two indices are given in the end of the

book, a General Index and an Index to Scientific Names.

This book is certainly an invaluable addition to the orchid flora treatments

of continental Asian orchids, and as such will be used for decades to come. The text

is very detailed, clearly written and well laid-out, and the numerous black-and-white

drawings, as well as the magnificent colour photographs and fine paintings, make it a

clearly outstanding achievement. The authors and the publisher can be congratulated

to the production of this comprehensive handbook on the native orchids of Hong

Kong. The book is well suited both to the botanist who studies Hong Kong orchids

and to the amateur who is growing native orchids and is trying to name orchids in the

wild. It will also prove an indispensable resource for researchers studying orchid floras

elsewhere in continental Asia.

Hubert Kurzweil

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 64(1): 277-278. 2012 277

BOOK REVIEW. Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 2.

(Malayan Forest Records No. 49) R. Kiew, R.C.K. Chung, L.G. Saw, E. Soepadmo &

PC. Boyce (eds). 2011.

Kepong: Forest Research Institute Malaysia. 25.7 cm = 18 cm, hard cover. 235 pp. ISBN 978-

967-—5221-53-8. Price RM 100 / USS 75.

Barely a year after the much anticipated first volume of the Seed Plants series, Flora of

Peninsular Malaysia, the second volume was published. The contents of this volume

are well structured and typically conform to the format preset for the first volume (see

Review in Gardens’ Bulletin, Singapore 62(2): 331-332).

This second volume is entirely devoted to the family Apocynaceae, with special

emphasis on two of the five subfamilies, namely, Apocynoideae and Rauvolfioideae.

The author of this momentous volume, Dr. David Middleton (Royal Botanic Garden,

Edinburgh), is an accomplished apocynologist who specialises in the two subfamilies of

Southeast Asian Apocynaceae. His past significant achievements in the region include

revisions for the Flora of Thailand (1999), Tree Flora of Sabah and Sarawak (2004).

and Flora Malesiana (2007). The other three subfamilies not treated in this volume

are the Asclepiadoideae, Periplocoideae and Secamonoideae. These three subfamilies

were formerly included in their own family, Asclepiadaceae, and rejoined Apocynaceae

only recently based on phylogenetic findings. In this account, a total of 35 genera and

112 species, namely, 21 genera and 55 species for subfamily Apocynoideae, and 14

genera and 57 species for subfamily Rauvolfioideae, are enumerated.

The taxonomic account begins with a brief family description, followed by

general distribution, ecology, general uses with some highlights of native species used

as ornamental plants, and a brief taxonomic explanation of the family, in a broad sense,

that includes Asclepiadaceae. Adding a phylogenetic tree to illustrate the evolutionary

relationships of the five subfamilies in the more broadly defined Apocynaceae would

have made it more informative. Conservation assessment was briefly provided by L.S.L.

Chua with an accompanying table listing Critically Endangered (CR), Endangered (EN)

and Vulnerable (VU) species in the two subfamilies. Arrangement of the taxonomic

account in this volume is based on the position of the two subfamilies reflected in the

key to subfamilies of Apocynaceae, beginning with subfamily Rauvolfioideae, and

later Apocynoideae. However, genera enumerated were arranged alphabetically under

each respective subfamily. The general enumeration format is as follows: accepted

name, etymology, key references and type citation, description, vernacular name,

distribution, species distribution map, conservation status, ecology, and uses.

In general, this volume is easy and friendly to use, not just for botanists but

also horticulturists. There are two identification keys provided at the beginning of the

account, namely, key to subfamilies of Apocynaceae (p. 8), and key to genera for the

two subfamilies revised in this volume (pp. 9-13), before the enumeration of genera

begins. Then, there are also identification keys to species provided in the account

for the following genera: subfamily Apocynoideae: Anodendron, Chonemorpha,

Epigynum, Holarrhena, Kibatalia, Micrechites, Parameria, Parsonsia, Strophanthus,

Urceola, and Wrightia; and subfamily Rauvolfioideae: Al/stonia, Alyxia, Cerbera,

278 Gard. Bull. Singapore 64(1) 2012

Chilocarpus, Kopsia (also key to varieties of Kopsia griffithii), Leuconotis, Melodinus,

Rauvolfia, Tabernaemontana, and Willughbeia. In addition to that, this volume is also

well illustrated with 15 colour plates provided at the end of the account and also 36

line drawings distributed throughout the book for the various genera.

As the Flora of Peninsular Malaysia team is picking up its pace to document and

disseminate information of Peninsular Malaysian plants to the scientific community,

we eagerly look forward to the next volume.

Y.W. Low

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 64(1): 279-280 2012 279

BOOK REVIEW. Wild Orchids of Peninsular Malaysia. PT: Ong, P. O'Byrne,

W.S.Y. Yong & L.G. Saw. 2011.

Kepong: Forest Research Institute Malaysia. 28.9 cm = 22.8 cm, hard cover. 196 pp. ISBN

978-967-5221-66-8. Price RM 150.

It is estimated some 10 per cent of all flowering plants are orchids, with perhaps 22,500

species worldwide. The Malay Peninsula has an impressive diversity, with around 900

species and 143 genera enumerated for the political part that is Peninsular Malaysia, a

tally now expected to reach c. 1000 species as exploration proceeds through to many

areas previously much less accessible. This orchid flora has 400-plus species in common

with that of the wetter territories around—each of Borneo, Sumatra, Thailand—and

less so with regions more seasonal or farther away, thus some 300 species with Java, c.

160 species with the Philippines, and just 60 species shared with Papua New Guinea.

A fifth of Peninsular Malaysian orchids are endemic, not found elsewhere.

There are seven chapters: Introduction, Diversity, Habitats, Pollination,

Classification, Scientific Research & Conservation, and The Path Ahead (the last

essentially concluding statements centring on propagating the orchid message and

contributing to orchid protection and conservation). Conservation discussion provides

a variety of snippets, dealing even with orchid thievery that all too often feeds an

obsession in the bid for unique possessions. Good summary accounts of the orchids

of various vegetation types found in Peninsular Malaysia, from the lowlands to

mountaintops, and on specialised substrates such as limestone, are given.

The key emphasis is on the impressive diversity in size, habit and floral structure

and display. Some key facts are referenced by numbers referring to sources listed at

the end of the book. There is, throughout, an attempt to explain would-be technical

terms, from “species” and “genus” across a wide range to, for example, “heterotrophic

mycotrophs”, the annointed term for non-chlorophyll-producing (so typically leafless)

orchids that rely on the absorption of materials broken down by their fungal associates

for their nutrition. Interesting snippets of information are carefully infused, covering

such aspects as the adaptations of epiphytes, CAM photosynthesis and, of course,

pollination. The information on pollination definitely provides many highlights and

perhaps is the single most original aspect of this book, even though practically all

areas covered have been wonderfully, albeit briefly, managed. Original information

on pollination is included, revealing a newly emphasised area of investigation that

not many, except the patiently observant, methodically curious and well-equipped,

can partake. This fascination emerges early, as the very first photo with the opening

main text already mentions the “rather nasty, fishy scent” of Bulbophyllum virescens

flowers.

Text style is creative (a few section titles may appear to be an effort) and

imaginatively crafted ‘stories’ teach a great deal about orchid specialisations and

biology. One could certainly say that no part of this book tends to the verbose.

After just some 60 pages of the most interesting primary information, the survey of

representatives from the different orchid subfamilies takes the form of a s/ab of richly

illustrated pages—evoking a curiously rebellious deja vu as it piles up an impressive

280 Gard. Bull. Singapore 64(1) 2012

compilation of richly coloured illustrations somewhat reminiscent of all big illustrated

accounts, yet with the biological and ecological sense they are organised, and by the

time the reader reaches this portion of the book, it is all too clear that a great deal of

knowledge and experience have been behind this.

A lot of effort has been put into photographing orchids and assembling

illustrative images for the many topics covered, so essentially a variety of useful

illustrated glossaries aid the beginner’s initiation into a systematically woven (and

may I add, eagerly pursued) introduction to orchid diversity. The authors are also

the main photographers (you can tell that they enjoy orchids). There is a case for

wondering if glossy art paper was the best choice for this book but colour printing

is well served. Heavy use is made of black page backgrounds, to provide maximum

contrast. Some extreme bursts of colour are cleverly choreographed into the account.

_ And—the bizarre view of pollinating blowflies crawling all over a Bulbophyllum

lasianthum inflorescence on the back cover simply closes this account with a thump.

K.M. Wong

Singapore Botanic Gardens

Instructions for contributing authors (continued)

Title and authorship. The title should concisely describe the contents. If a scientific name is used, its

authority is normally excluded, but the family name would be provided. Authors” names, affiliations

and postal/ e-mail addresses are stated below the title. If more than one author, indicate “corresponding

author”. Avoid footnotes. A short running title (up to six words) should also be provided.

Abstract. The abstract is at most 100—300 words. It should concisely indicate the article’s contents without

summarising it: mentioning novelties and name changes. Keywords: Suggest at most eight keywords, in

alphabetical order.

Scientific names and author abbreviations. Genus and species names of organisms must be italicised and

followed by the authority (with family name in parentheses) when first mentioned in the text or diagnoses.

Standards for author abbreviations include:

Brummitt, R.K. & Powell, C.E. (1992) Authors of Plant Names. Kew: Royal Botanic Gardens, Kew.

Kirk, P.M. & Ansell, A.E. (1992) Authors of Fungal Names [Index to Fungi Supplement]. Wallingford:

CAB International.

Herbarium abbreviations (http://sciweb.nybg.org/science2/IndexHerbariorum.asp) follow

Holmgren, P.K., Holmgren, N-H. & Barnett, L.C. (eds) (1990) Index Herbariorum. Part 1: The Herbaria

of the World [Regnum Veg. vol. 120]. New York: New York Botanical Garden.

Journal and book title abbreviations. For journals:

‘Bridson, G.D.R., Townsend, S.T., Polen, E.A. & Smith, E.R. (eds) (2004) BPH-2: periodicals with

botanical content: comprising a second edition of Botanico-Periodicum-Hunlianum. Pittsburg: Hunt

Institute for Botanical Documentation.

For books:

Stafleu, F.A. & Cowan, R. S. (eds) (1976-88) Taxonomic Literature. 2nd edition. 7 vols [Regnum Veg.

vols 94, 98, 105, 110, 112, 115, 116].

Stafleu, F.A. & Mennega, E.A. (eds) (1992—) Taxonomic Literature. Supplements [Regnum Veg. vols

125, 130, 132].

(A useful source of verify ing names of publications is

<http://asaweb.huh.harvard.edu:8080/databases/publication_index.html>. If in doubt, list full titles.

Other abbreviations and units of measurement. If using standard abbreviations and acronyms, give the

full term on first mention. Dates are cited as: 1 Jan 2000. SI (metric) units of measurement are used and

spelled out except when preceded by a numeral; they are abbreviated in standard form: g, ml, km, etc.

Tables. Tables are numbered in arabic numerals in the order they are first mentioned in the text and carry

an indicative legend at the head. Tables are given at the end of the manuscript.

Illustrations. All drawings, maps, graphs and photographic images (individually or collected in a plate)

are to be numbered in arabic numerals in the order they are first mentioned in the text, as Fig. 1, Fig. 2,

etc. (plate components would be referred to in the text as Fig. 1A, 1B, Fig. 1A—D, etc.). If relevant, scale

bars should be used to indicate magnification.

When grouping photographs, the maximum page area 19.5 x 13 cm must be heeded. High resolution digital

images may be submitted as separate files (line drawings in black and white at 600 dpi, photographs at

300 dpi) sent electronically or in a CD. Do not embed images into the main text file.

References in the text. Citation in the text should take the form: King & Gamble (1886) or (King &

Gamble 1886), or King et al. (1886) if more than three authors to a work. Use 2000a, 2000b, etc. if several

papers by the same author(s) in one year are cited.

References listed at the end. There, works mentioned in the text are listed alphabetically as follows:

Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999) User “s Guide to the DELTA Editor. http:

biodiversity.uno.edu/delta/ (accessed on 2 Aug. 2010).

Persson, C. (2000) Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the

rps 16 intron and trnL(UAA)-F(GAA) intergenic spacer. Nordic J Bot. 20: 257-269.

Ridley. H.N. (1930) The Dispersal of Plants Throughout the World. Ashford, U.K.: L. Reeve.

Smith, A.C. & Darwin, S.P. (1988) Rubiaceae. In: Smith, A.C. (ed) Flora Viliensis Nova, A New Flora of

Fiji 4: 143-193.

References to web-based resources should include either a doi (digital object identifier) specification

or full URL mentioning also the date it was accessed. Use of DNA sequences from GenBank should be

acknowledged and the studies for which the sequences were generated should be cited.

Style of nomenclatural summaries. The following style is required:

Gardenia anisophylla Jack ex Roxb., Fl. Ind. ed. Carey & Wall. 2: 561 (1824).

Medinilla alternifolia Blume, Mus. Bot. 1: 19 (1849).

If authors include full bibliographic data for these works in the list of references at the end of the paper,

they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include

Blume (1849) and Roxburgh (1824).”

Homotypic synonyms should be provided in a block, stating the type at the end.

Front cover picture: Amischotolype monosperma, Peninsular Malaysia (Photo by A. Schuiteman)

Back cover: Gardens’ Bulletin covers. 1912?—1969 (Montage hv Bazilah M. Ibrahim & S. Lee)