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PART 2

KIEW, Ruth:

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yy KENG, Hsuan:

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49 SAW, Leng Guan and DRANSFIELD, John:

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4% Book Review

» ROBBRECHT, E.:

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Published by the Botanic Gardens

Parks and Recreation Department

Ministry of National Development

Cluny Road, Singapore 1025.

GARDENS’ BULLETIN

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THE GARDENS’ BULLETIN

SINGAPORE

VOL. 42 (Part 1 & 2) 1 December 1989

CONTENTS

PART 1

ARDITTI, Joseph:

aa ee Gls PAE AEMIST ATIC. OUGIET © o'snda vcs 5.2 ba ncaa ds cicchsc 0ecnSontaueasanetivescec. eS

FOONG, T.W., YANG, C.N. and MUSTAFA, binte Nashita:

An Atlas of Nutrient Deficiency Symptoms in

EMER MERIT AOIIGIN oan. caine vn aeae vga cna Gee ccateesteeenr ve yn cats usens rest sas tan dess 8 19-24

ADAM, Jumaat Haji and WILCOCK, C.C.:

A New Nepenthes from Gunong Besar,

eimeiedrsel aim? POTNEG) fi.) 220) 30. 2 RO) PO Le 25-28

TURNER, Ian:

An Enumeration of One Hectare of Pantai Aceh

nena EIN aE ERRATA OR ce Ae ae epoca. ec once wauwaiewsdacasse¢h<sueT<cndevens 29-44

PART 2

KIEW, Ruth:

Didymocarpus (Gesneriaceae) on Gunung Tahan, Malaysia .................... 47-64

KENG, Hsuan:

whe Genus Camelia (iheaceac) in’ Malesia 92) .05 A nde dee dee 65-69

SAW, Leng Guan and DRANSFIELD, John:

A New Species of Licuala (Palmae) from Peninsular Malaysia ................ 71-73

Book Review

ROBBRECHT, E.::

CO IGS OS Ta oe ee Oe eee 75-77

Published by the Botanic Gardens

Parks and Recreation Department

Ministry of National Development

Cluny Road, Singapore 1025.

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Singh, H. (1967). Sclereids in Fagraea. Gard. Bull. Sing. 22, 193-212.

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Kent; 242-255.

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Medinilla alternifolia Bl., Mus. Bot. Lugd.-Bat. I:2 (1849) 19.

Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524.

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THE GARDENS’ BULLETIN

SINGAPORE

Volume 42

Part 1

Kwan Koriba: Botanist and Soldier

JOSEPH ARDITTI

Department of Developmental and Cell Biology

University of California

Irvine, California 92717

USA

There is no nationality in the geen ee

field of science

icine Weil Charles Dickens

A Tale of Two Cities, Book I, Chapter |

Abstract

Kwan Koriba (1887-1957), a noted Japanese botanist, educator and academic administrator was

given the rank of Brigadier General and appointed Director of the Singapore Botanic Gardens from

December 1942 to August 1945 during the World War II occupation. As Director he performed his duties

exceptionally well, carried out first rate scientific research and treated his British colleagues with com-

passion and understanding. After the war Prof. Koriba was appointed to be the president of Hirosaki

National University.

Introduction

Dickens wrote a novel about a noble deed during times of strife and upheaval.

This is a true tale of noble deeds during a time of war. The deeds are those of Prof.

(and for a period) Brigadier General Kwan Koriba who was Director of the Singapore

Botanic Gardens from December 1942 until late August 1945.

When I was a graduate student at the University of Southern California (1960-1965)

my first Department Chairman was Prof. John L. Mohr. John was a marine biologist

and we did not have much in common scientifically (except that in the Summer of

1962 I worked as his data collector for him on Arlis II, an ice island near the North

Pole), but I took his course on the History of Biology. This was easily the best

course I ever took because John was an erudite and interesting teacher who taught

us to appreciate scientific integrity and scientists who held to their principles even

at a very high personal cost.

My last Chairman at USC before I left for UCI was Prof. Paul Saunders. He

was a biochemist whose research centred on poisons produced by marine animals.

Shortly before becoming Chairman he visited the Pacific to collect toxins and the

animals which produce them. Paul gave a seminar shortly after becoming Chairman

and never tired of telling us about his trips and the places he visited. One of these

was Singapore and he told us about the city, the harsh Japanese occupation (he was

there 10-12 years after the end of the war), and (for my sake) the Botanic Gardens,

their orchids and their war-time Japanese Director (whose name Paul had forgotten)

who was a very kind and well liked man. Paul’s accounts stimulated my interest in

the Director since he seemed to be like the principled universal men of science in

John Mohr’s lectures.

In 1969, approximately 15 years after taking John’s course and a decade follow-

ing Paul’s account, I visited Singapore for the first time. Before going to the Botanic

Gardens I went to the Botany Department of the University of Singapore to meet

Prof. A.N. Rao who already had (and still has) an impressive reputation as an

orchid anatomist. He introduced me to Prof. P.N. ‘Danny’ Avadhani. We spent a

2 Gard. Bull. Sing. 42(1) (1989)

very pleasant day together and the visit led to what is now a 30-year friendship and

scientific collaboration carried out during summer and sabbatical leave visits to

Singapore.

Rao and Danny took me to the Botanic Gardens where I saw on a wall the

photographs of all previous directors including those of H.N. Ridley who died at

the age of 101 and is primarily responsible for the introduction of rubber trees into

Malaya; R.E. Holttum who lived through the war in the Gardens and in writing a

book on the Orchids of Malaya wrote one of the best orchid floras ever written, and

the war-time Japanese Director Prof. Kwan Koriba (Fig. 1). His picture was (and

still is) there despite the fact that he was imposed on the Gardens by an occupation

force. I asked for more details about Prof. Koriba, but the only answers were that

he was a good director and a kind man.

Fig. 1. Professor Koriba in a field near Hirosaki. This photograph was taken by Professor Shigeo Chiba,

then a student at Hirosaki University on September 26, 1957. A copy, taken to Singapore in

1971 by Professor Chiba, is now displayed among the photographs of other former Directors

of the Singapore Botanic Gardens (courtesy Prof. Shigeo Chiba).

In the introduction to his book on the Orchids of Malaya Holttum wrote: “‘The

main part of the work for.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 Singapore by the Japanese Government to take charge of the Botanic Gardens

Kwan Koriba 3

here. I wish to express grateful thanks to Prof. Koriba, for the courtesy with which

he allowed me complete freedom to continue my studies, and for much personal kind-

ness during that period’’ (Holttum 1964, the first edition of his book was published

in 1953).

On initiating the Orchid Biology series I decided to follow the example of the

Annual Reviews of Plant Physiology and to invite eminent orchidologists past retire-

ment age to write the first chapter in each volume. My first choice for volume I was

Prof. Holttum who was kind enough to accept the invitation. He wrote in this

chapter that with ‘‘... the Japanese occupation of Singapore came Professor H.

Tanakadate who said ‘I conserve cultural institutions’ and asked me to remain in

charge of the Gardens. I did so, but was formally relieved of my charge after about

ten months, when Professor K. Koriba, a distinguished plant physiologist, came as

Director. He permitted me and Mr. Corner to continue our botanical studies. It was

then that I began to write a systematic account of the orchids of the Malay peninsula

woe (aolttumaty77):

Holttum’s statements, especially the first one, kept me wondering whether Koriba

was ever interested in orchids. My inquiries always elicited a negative answer and

I was told that he worked on forest trees. As it turns out these answers were in-

correct. When Dr. L.P. Nyman, Dr. R. Ernst and I decided to write a review on

resupination we surveyed the literature and found that most of it was by German

workers. However two often cited papers were by a K. Koriba (Koriba 1913, 1914).

I did not know whether this was merely a coincidence or if the author was Prof.

Kwan Koriba. The question was not resolved even after I obtained copies of the

papers. These papers dealt with resupination, but Prof. Koriba’s presumed interest,

at least while he was in Singapore, were forest trees and their phyllotaxis. At that

time I could see no obvious connection between phyllotaxis and resupination. In an

effort to obtain additional information I wrote Prof. Holttum. He replied suggesting

that I should consult E.J.H. Corner’s book The Marquis — A Tale of Syonan-to

and also write Prof. Akinori Ueyama at Kyoto University. Corner’s book proved

to be very interesting and to contain information on Prof. Koriba, but did not

resolve the puzzle.

I wrote Prof. Ueyama. His reply was delayed for a week or two due to ill health,

but it answered my question. The K. Koriba who wrote these papers was indeed a

Prof. Kwan Koriba. These papers dealt not only with resupination per se, but also

with the placement and subsequent positioning of flower buds on the stems of Spiran-

thes australis — a subject not far removed from phyllotaxis. I was pleased to find

that Prof. Koriba was interested in orchids after all and wrote Prof. Ueyama several

times asking for biographical information, photographs and any other details he

might have. Prof. Ueyama was patient, kind, gracious and generous. He sent me

photographs, copies of Koriba’s papers, all the information he had and translated

parts of a posthumous collection of Koriba’s writings.

I also obtained copies of several obituaries (Anonymous 1947; Asida 1943; Ashida

1958; Holttum 1958). Two of them (Asida 1943; Ashida 1958) listed a third paper

on orchids by Koriba, a contribution dealing with 7aeniophyllum (Koriba 1926).

From Koriba’s obituaries I learned that in his final years he was President of Hiro-

saki University (Fig. 2). I wrote to the current President Prof. Shuji Tono requesting

more information and received a very kind reply from Prof. Shigeo Chiba, one of

the last students to hear Prof. Koriba lecture at Hirosaki University in 1956 (S.

Chiba, Hirosaki University, personal communication), who sent me biographical

information and several photographs. Finally, almost 25 years after Paul Saunders

mentioned the war-time Director of the Singapore Botanical Gardens I had enough

information to write about Professor Kwan Koriba.

4 Gard. Bull. Sing. 42(1) (1989)

Fig. 2. An official photograph of Professor Kwan Koriba as President of Hirosaki University. This

photograph taken on September 6 1957 (according to Prof. Shigeo Chiba) marks Prof. Koriba’s

75th birthday, 26 September 1957 (courtesy Prof. Akinori Ueyama and Prof. Shigeo Chiba;

each supplied me with a copy).

Outline of Koriba’s Career

In one of his letters Professor Ueyama Akinori translated for me an outline of

Koriba’s life and career which was published first in a posthumous collection of his

works. Prof. Ueyama (1927- _ ) stated in his letter that he did not know much about

Prof. Koriba who was ‘‘grandfather’’ Professor to him ‘‘... because [the] late Prof.

Hamada [1910-1981] was his student ...’’ This outline is presented below with

minor modifications in language and additions based on information from several

sources (Asida 1943, 1958; Holttum 1958; S. Chiba, personal communication).

1882 (Meiji 15th year) September 6, 7:00 pm, Koriba was born at Aomori-city, in

the North of Honshu, North East Area of Japan, the 4th child and 2nd son of

Naoyo Koriba ‘‘... a ‘Samurai’ of the feudatory at the north end of Honshu ...”’

As a boy he was educated in the Middle School of the neighbouring city of Hirosaki

(Holttum 1958). His interest in plants was aroused by one of his teachers a plant

> an

Kwan Koriba 5

pathologist named Naoharu Hiratsuka. His mother Kumi was also interested in plants ©

(Asida 1943).

1903 (Meiji 36th year) September. Entered Tokyo Imperial University College of

Science, Department of Botany.

1907 (Meiji 40th year), Koriba, 25 years old, graduated from the University with an

undergraduate degree. Thesis: Uber die individuelle Verschiedenheit in der Entwicke-

lung einiger fortwachsenden Pflanzen mit besonderer Ricksicht auf die Aussen-

bedingungen. J. Coll. Sci., Imp. Univ. Tokyo 27(3): 1-86 (1909). His Professor was

Manabu Miyoshi (Asida 1943).

1907 (Meiji 40th year), entered graduate course at the University of Tokyo.

1912 [Meiji 45 = Taisho | (Meiji-Tenno passed away and the era in the Japanese calen-

dar was changed to Jaisho-Tenno), December 10th, Koriba, 30 years old, obtained

Rigaku-Hakushi (Dr. of Science, Tokyo University). Dissertation: Mechanisch-

physiologische Studien itber die Drehung der Spiranthes-Ahne. J. Coll. Sci. Tokyo

Imp. Univ. 36(3): 1-179 + 7 plates (published March 30, 1914).

1913 (Jaisho 2), 31 years old, appointed lecturer at the College of Agriculture,

Tohoku Imperial University, Sapporo.

1915 (Taisho 4), 33 years old, appointed Professor of Botany at the same University.

1917 (Taisho 6), December 28, appointed one of the designers of a New Institute

and a Department of Botany which he inaugurated at the College of Science, Kyoto

Imperial University.

1918 (Jaisho 8), March 21, became overseas research scientist for the Japanese

Government and visited the USA, UK, France, Italy and Sweden. During this trip

he had the good fortune of meeting Professor W. Pfeffer just before his death.

Koriba also attended Pfeffer’s funeral and laid a wreath in the name of the botanists

of Japan (Asida 1943).

1920 (Zaisho 9), August 8, returned via the port of Kobe. August 20, appointed

Professor at the Botany Department, College of Science Kyoto Imperial University.

1921 (Jaisho 10), Koriba, 39 years old, was one of the designers of a New College

of Agriculture at Kyoto Imperial University.

1921 (Zaisho 10), appointed Director of the Botanical Gardens, Kyoto Prefecture

and held the post until October 1928.

1921 (Taisho 10), he visited Ponape Island (Ashida 1958).

1924 (Taisho 13), Koriba was appointed member of the National Research Council

of Japan (Asida 1943).

1926 (Taisho 15), Koriba attended the Third Pan-Pacific Science Congress in Japan

and presented a paper on Jaeniophyllum (Asida 1943).

1929 [(Showa 4) Taisho-Tenno died in 1927 and a new Majesty, Hirohito-tenno

assumed the throne, he was very interested in biology], April-July, Koriba 47 years

old, visited and did research in Java during a Pan Pacific Congress. Prof. Akinori

Ueyama wrote me that he learned from Holttum in 1976 at Kew that Prof. K. Koriba

first met Prof. R.E. Holttum, then Director of the Singapore Botanic Gardens, at

this meeting. On the same line Ueyama added ‘‘1925(?)’’ without an explanation.

1932 (Showa 7), 21 June to January 13, 1933, Koriba 50 years old, was appointed

Overseas Scientist for the Japanese Government and visited Germany, Belgium, the

Netherlands, UK, France, South America (Brasil, Argentina, etc) and the USA

before returning to Kyoto. These visits may have started in 1931 (Ashida 1958). I

can not help but wonder whether these and his previous international travels gave

Koriba a good feeling for scientists of other nationalities. If so these feelings could

be partially responsible for his kindness in Singapore. At the same time I am sure

that the major reason for his attitude in Singapore must have been his personality

and character.

6 Gard. Bull. Sing. 42(1) (1989)

~ 1940 (Showa 15), 17 August-21 September, Koriba, 58 years old visits Manchuria,

Mongolia and Central China.

1942 (Showa 17), Koriba reached the age of 60 kanreki in Japanese and retired on

26 September. On 22 December he arrived in Shonon-to (Prof. Ueyama used this

spelling and not Syonan-to like Prof. Corner) as supreme cultural advisor in what

was then an occupied area.

1943 (Showa 18), 15 April, Koriba was appointed as Director of the Singapore

Botanic Gardens where, Prof. Ueyama writes, he ‘‘... co-studied with Holttum,

Cotmi,r, ctc.”’

1944 (Showa 19), 15 December, Koriba was given the additional post of Director of

the Singapore Museum.

1945 (Showa 20), Japan surrendered, Koriba left the Gardens in late August accor-

ding to Prof. Corner who kept Koriba’s research notes and a draft of a paper which

was eventually published in the Gardens Bulletin, Singapore (Koriba 1958).

1946 (Showa 21), the 64 year-old Koriba left Singapore on 26 January and arrived

in Kyoto on 13 February.

1948 (Showa 23), at the age of 66 Koriba became Professor Emeritus at Kyoto

University. According to one report (Ashida 1958) Koriba spent 1946-1954 reading

and writing. He published two books during that period (Holttum 1958).

1954 (Showa 29), 1 February, Koriba, age 72, was appointed President of Hirosaki

National University in his native prefecture of Aomori. Prof. Ueyama thinks that

he accepted the position as a means of returning to his birth area. His assumption

is accurate as indicated by a letter (dated 27 March 1954) from Koriba to Corner:

. This year I was elected to the President of the Hirosaki University.... As

Hirosaki is my native prefecture (Aomori) I could not deny the request, though |

get fairly old.’’

1956 (Showa 31), 3 May, the Aomori-ken Biological Society was formed and Prof.

Koriba, a founding member, became its first president (S. Chiba, personal com-

munication).

1957 (Showa 32), 14 December, Prof. Kwan Koriba passed away (fuki in Japanese,

meaning ‘‘no return’’ or ‘‘died’’ according to Prof. Ueyama) at the age of 75 in the

bathroom of his official residence. Another report (Ashida 1958) gives the date of

his death as December 15, 1957. His funeral was held on 21 December 1957 (S.

Chiba, personal communication).

1958, Koriba’s paper on the periodicity of tree growth in the tropics was published

posthumously in the Singapore Botanic Gardens’ Gardens’ Bulletin Volume 17,

No. 1, pages 11-81.

1958, a volume commemorating Prof. K. Koriba edited by Prof. Joji Ashida was

published in Japanese (S. Chiba, personal communication).

1983, 23 October, Prof. E.J.H. Corner visited Hirosaki University and put flowers

on Prof. Koriba’s grave (Chiba 1984).

The Botanist and Orchidologist

Young Koriba developed an interest in ecology and morphology. The writer of

one obituary (Ashida 1958) attributed this to the fact that he lived in the mountains.

At the Imperial University in Tokyo Koriba devoted attention to physiology because

he felt that ecology and morphology can be explained in physiological terms (Ashida

1958). For this reason his early research was on the effects of ecological and physio-

logical factors on morphology. He utilized his considerable knowledge of physics

for this work, his orchid research and the observations on trees in Singapore (Holt-

tum 1958). His work with Spiranthes is an extension of this interest.

Kwan Koriba a

Koriba referred to Spiranthes sinensis (Fig. 3; Table 1), the subject of his doc-

toral dissertation (Koriba 1914), as Spiranthes australis. This is not surprising at all

since this species has many synonyms perhaps due to its very wide distribution

which includes Japan, Taiwan, Philippines, Indo-Malaya (Pakistan, Himalayas, etc),

Afghanistan, Thailand, Indochina, China, Korea, Malaysia, Australasia (New Guinea,

Australia, New Zealand, Tasmania, etc) and parts of the Middle East (Gagnepain

and Guillaumin 1933; Holttum 1964; Garay and Sweet 1974; Lin 1976; Rechinger

1978; Seidenfaden 1978). The plants are terrestrial and very variable in form and

Size.

Wes,

il gE

Fig. 3. Spiranthes sinensis. A. Specimen from Taiwan; a, whole plant with two inflorescences; b, flower

with bract; c, lateral petal; d, dorsal sepal, e, lateral sepal; f, labellum and gynostemium;

g, gynostemium; h, pollinia; i, pollinium; j, labellum. B. Example from Taiwan; a, whole plant;

b, flower; c, labellum; d, petals and dorsal sepal. C. Plant from Indochina; a, side view of

flower, <4; hood formed by two lateral petals and the dorsal sepal, x6; c, lateral sepal x 6;

d, labellum and gynostemium x 6; e, anther x 10; f, pollinia x 10. D. Representative from Italy;

a, leaves and inflorescence; b, the spiral on the stem of the inflorescence; c, flowers on side

(bottom) and 45° (top) view, (sources: A, Lin 1976; B, Seidenfaden 1978; C, Gagnepain and

Guillaumin 1933; D, Alessandrini 1985).

8 Gard. Bull. Sing. 42(1) (1989)

Table 1

Nomenclature of Spiranthes sinensis (S. australis)

Spiranthes sinensis (Pers.) Ames, Orch. 2: 53, 1908; Masamune in Sci. Kep. Kanazawa Univ. 9:

126, 1964.

Basionym: Neottia sinensis Pers., Syn. 2: 511, 1807.

Syn.: Ophrys spiralis Georgi, Reise 1: 232, 1775.

Aristotelea spirals Loui., Fl. Cochinch. ed. 1, 2: 522, 1790.

Epidendrum Aristoelea Raeusch., Nomencl. ed. 3, 265, 1797.

Neottia australis R. Br. Prodr. Fl. Nov. Holl. 319, 1810.

Neottia flexuosa Smith in Rees, Cyclop. 24: Neottia no. 9, 1813.

Neottia parviflora Smith in Rees, Cyclop. 24: Neottia no. 10, 1813, not Blume.

Neottia amoena M.v. Bieb., Fl. Taur. Cauc. 3: 606, 1819.

Spiranthes pudica Lindl., Coll. Bot. t. 30, 1821.

Neottia australis var.

chinensis Lindl. in Bot. Reg. 7: t. 602, 1821.

Spiranthes australis (R. Br.) Lindl. in Bot. Reg. 10: sub t. 823, 1824.

Spiranthes parviflora (Smith) Lindl. in Bot. Reg. 10: sub t. 823, 1824.

Spiranthes flexuosa (Smith) Lindl. in Bot. Reg. 10: sub t. 823, 1824.

Neottia crispata Bl., Bijdr. 8: 406, 1825.

Spiranthes amoena (M.v. Bieb.) Sprengl., Syst. Veg. 3: 708, 1826 (excl. syn. S. congesta)

Neottia pudica (Lindl.) Sweet, Hort. Britt. ed. 2, 485, 1830.

Spiranthes Wightiana Lindl. in Wall. Cat. no. 7378, 1832, nomen.

Calanthe australis Ait. ex Loud., Hort. Brit. Suppl. 2: 615, 1839, nomen.

Spiranthes longispicata A. Rich. in Ann. Sci. Nat. ser. 2, 15: 78, 1841.

Spiranthes densa A. Rich. in Ann. Sci. Nat. ser. 2, 15: 78, 1841.

Spiranthes indica Steud., Nomencl. ed. 2: 625, 1841.

Sarcoglottis pudica (Lindl.) P.N. Don in Hort. Cantabrig. ed. 13, 590, 1845.

Spiranthes Novae-Zeylandiae Hook. f., Fl. N. Zeal. 1: 243, 1853.

Spiranthes crispata Zoll., Syst. Verz. 89, 1854.

Spiranthes australis var. pudica (Lindl.) Lindl. in Hook. Kew Journ. Bot.

72.38, 18533

Spiranthes stylites Lindl. in Journ. Linn. Soc. Bot. 1: 178, 1857.

Gyrostachys australis (R. Br.) Bl., Fl. Jav. Orch. 107, t. 37, 1858.

Gyrostachys amoena (M.v. Bieb.) BI., Fl. Jav. Orch. 108, 1858.

Gyrostachys australis var. amoena (M.v. Bieb.) Bl., Fl. Jav. Orch. 129, 1859.

Gyrostachys australis var. flexuosa (Smith) BI., Fl. Jav. Orch. 130, t. 38, 1859.

Spiranthes australis var. suishaensis Hayata., Icon. Pl. Formos. 6: 86, 1916.

Spiranthes Aristotelea (Raeusch.) Merr. in Phil. Journ. Sci. 15: 230, 1919.

Spiranthes australis var. viridiflora Makino in Journ. Jap. Bot. 3: 5, 1926.

Spiranthes australis var. sinensis (Pers.) Gagnep. in Lecomte, Fl. Gen. Indo-Chin. 6:

546, 1933.

Spiranthes spiralis Makino in Journ. Jap. Bot. III, 7: 25, 1934.

Spiranthes spiralis var. albescens Honda in Bot. Mag. Tokyo 49: 791, 1935.

Spiranthes spiralis var. depauperata Honda in Bot. Mag. Tokyo 49: 698, 1935.

Spiranthes sinensis var. albescens (Honda) Honda in Bot. Mag. Tokyo 50: 669, 1936.

Spiranthes sinensis forma albiflora (Matuda) Iwabrechi in Tohuku Hakubutu-Kai |:

136, 1936.

Spiranthes spiralis forma albiflora Matuda in Tohuku Hakubutu-Kai 1: 136, 1936.

Spiranthes sinensis var. depauperata (Honda) Honda in Bot. Mag. Tokyo 50: 669, 1936.

Spiranthes sinensis forma viridiflora (Makino) Ohwi in Bull. Nat. Sci. Mus. Tokyo

33: 69, 1953.

Spiranthes sinensis subsp. australis (R. Br.) Kitamura in Act. Phytotax. Geobot. 21:

23, 1964.

Spiranthes lancea var. chinensis (Lindl.) Hatusima in Journ. Geobot. 16: 80, 1968.

Spiranthes sinensis var. amoena (M.v. Bieb.) Hara in Journ. Jap. Bot. 44: 59, 1969.

Garay, L.A. and Sweet, H.R. (1974). Orchids of Southern Ryukyu Islands. Bot. Mus. Harvard Univ.

Cambridge, Massachusetts. t

Kwan Koriba 9

The spiral arrangement of Spiranthes flowers, their position on the inflorescence

and orientation relative to the ground drew the attention of botanists at least 60

years before Koriba studied the phenomenon. Koriba was the first to study these

phenomena in detail. He carried out numerous observations and reported them in

great detail in an extremely long German language paper published in Japan and a

shorter one in a German journal (Koriba 1913, 1914). What he found was that

1. while the flowers open the axis of the inflorescence twists and this causes the

spiral appearance of the raceme.

2. this torsion is limited to the axis of the inflorescence.

3. the growth of the inflorescence influences the turning.

4. flowers change their position (Fig. 4) and undergo resupination either from

left to right or right to left.

5. the directions of twisting of the inflorescence [right to left or left to right

(clockwise and counterclockwise) if one looks at it from above] and resupina-

tion are the same. But I wonder if the flowers did not alternate in resupinating

in a clockwise and counterclockwise direction.

6. if the tip of the raceme is prevented from turning the rotation of the entire

inflorescence is affected.

7. the twisting of the inflorescence and movement by the buds are inhibited on

a horizontal clinostat.

As a result of his studies Koriba suggested that the twisting of Spiranthes in-

florescences is due to differential growth of tissues. To reach these conclusions

Koriba did a considerable amount of very painstaking work. I found it difficult reading

his papers due to a poor knowledge of German.

Dr. Robert Ernst and Dr. Franz Hoffman, both born in German speaking coun-

tries, helped me, but I could not, of course, expect them to translate both papers

Fig. 4. Development and positioning of Spiranthes sinensis flowers (A) and (B) directions of twisting

of the floral axis and torsion of flowers (Koriba; 1914, Prof. Akinori Ueyama was kind enough

to provide a copy of this paper).

10 Gard. Bull. Sing. 42(1) (1989)

in their entirety and this is the reason for the sketchy summary presented here. Be

all this as it may, Koriba’s dissertation explains his subsequent interest in phyllotaxy.

The jump between the arrangement of flowers on an inflorescence and the position

of buds on a stem is not all that big.

In addition to studying the twisting of Spiranthes inflorescences Koriba also com-

pared the process with resupination by flowers of Orchis aristata, Malaxis paludosa,

and four species of Platanthera. These comparisons are a clear indication that he

was aware of the general nature of the phenomenon. Most subsequent reviews and

research papers on resupination cite Koriba’s work which is indicative of scientific

importance.

Koriba’s work with orchids was not limited to Spiranthes. He was also interested

in 7aeniophyllum aphyllum or kumo-lan, the spider orchid, a Japanese shootless

species. He described the position of its roots and noted that they are dorsiventral

(Koriba 1926).

Another phenomenon studied by Koriba was the effect of light on the develop-

ment and morphology of the roots. During his observations Koriba carried out a

simple yet clever experiment: ‘‘... when the growing apical portion of the hypocotyl

is cut off, the rest can regenerate a new hypocotyl with the germinal point, whereas

the root has no such capability’? (Koriba 1926).

Vegetation and succession on volcanoes, high mountains and dunes were among

other interests pursued by Koriba and he published several papers on these subjects

(Ashida 1958). He also worked on the Podostemonaceae and the fossil Hydrodictyon.

In 1927 he suggested that morphogenesis and differentiation should be explained on

the basis of plant hormones and vitamins. This is remarkable since the first plant

hormone, auxin, was discovered in 1926 by Prof. F.W. Went in Utreht. Starting in

1931 Koriba was interested in phytoclimatology and between 1936 and 1942 worked

on evaporation and transpiration.

While in Singapore Koriba studied or discussed buds, trees, phyllotaxis,

deciduousness, transpiration, plant distribution, fossils, palms, pandans, seeds and

floral development. Koriba was very interested ‘‘... in the occult problem of phyllo-

taxis ...’’ (Corner 1981) and explained it at great length to Corner who wrote that

‘*’.. sO mature an intellect had not previously arrived on the scientific scene in

Malaysia.’’ Altogether Koriba had a ‘‘... a very broad scientific outlook ...”’

(Holttum 1958). When Koriba puzzled he put his head on one side and bent his arm

over it.

On Sunday mornings Koriba, Dr. Yata Haneda (Director of the Museum), Holt-

tum and others would go by car to inspect parts of Singapore and forest reserves

and to study trees. During these trips Koriba, who was past 60 at the time showed

considerable bodily vigour to match his strength of mind and ‘‘... never paused for

breath, stumbled or appeared tired ...’’ On other occasions he, Dr. Yata Haneda

and Corner walked in the Botanic Gardens after dark to search for luminous insects

and fungi. Koriba continued to lead students and colleagues on field trips (Fig. 5-7)

and to engage in teaching and observations in the field even after he became Presi-

dent of Hirosaki University. This was very unusual since Presidents of National

Universities become executive officials in Japan (the same is true in the US) and seldom

teach. Koriba had special permission from the Ministry of Education, Science and

Culture to continue as a teacher (S. Chiba, personal communication).

Koriba’s chief interest in Singapore was the abundance of different kinds of trees

there and the diversity of growth habits. He wrote a paper on the periodicity of tree

growth there (Koriba 1958), but after the Japanese surrender in 1945 he left the

manuscript in Singapore as the Gardens’ property. In due time the manuscript was

returned to him and on 27 June 1953 he sent it to Corner for corrections. Corner

Kwan Koriba 11

Fig. S-6. Prof. Kwan Koriba on field trips. 5. Prof. Koriba leading students on a field trip, 2 June 1954.

Prof. Shigeo Chiba wrote that Prof. Koriba went on field trips often despite the heavy demands

placed on him by the Presidency of the University. 6. With Professor Kanzo Wada (right) of

Hirosaki University on a field trip near Aomori on 27 June 1954 (courtesy Prof. Shigeo Chiba).

must have returned it not long after that since on 27 March 1954 Koriba thanked

him for the corrections and apologized for a lapse in their correspondence. Two days

after that, on March 29th of that year Koriba commented that the manuscript was

too long and complained about not having enough time to do anything about it.

More than three years later, on 19 July 1957 he wrote Corner that according to

H.M. Burkill, Director of the Botanic Gardens at that time, the manuscript had gone

to the printer. Koriba read and corrected the galley proofs in his office as President

Lz Gard. Bull. Sing. 42(1) (1989)

het ee Kev: re

Auk +

Fig. 7. Professor Kwan Koriba at Mount Iwaki on 26 September 1957 (courtesy Prof. Shigeo Chiba).

of Hirosaki University with Professor Zyun Nakazawa shortly before his death (S.

Chiba, personal communication). The paper was published in 1958, but Koriba never

saw it. It is fitting I think that his last work was published in the Bulletin of the

Gardens he directed, obviously loved and took pains to save.

In 1951 Koriba published a book on plant forms in which he discussed mor-

phology from the viewpoints of organization, development and function. A year later

he published a comprehensive book on plant physiology and ecology (Ashida 1958).

He also planned to write a book on plant evolution. His notes reveal that in this

book he intended to approach the topic through a synthesis of morphology, physio-

logy, ecology and genetics. Few people could have done that and unfortunately

Koriba never had a chance to write that book.

The Director

To fully appreciate Koriba’s behaviour it must be placed in context and compared

with that of the Japanese occupation forces in Singapore. The Japanese entered

Singapore (or Syonan, Light of the South as they called it) on the morning after

General Arthur Percival surrendered the city unconditionally to General Tomoyuki

Yamashita (then called the Tiger of Malaysia and hanged for his war crimes after

the war as the only Japanese war criminal whose sentence was either not commuted

by General Douglas McArthur or who was not given the privilege of a more

honourable execution by a firing squad). Most of the population stayed indoors on

that day and stores were boarded up. Only the Singapore Cold Storage food store

Fs a a

Kwan Koriba 13

was open (this store, now modernized, still exists; its major contribution to orchidology

was the extensive use of the Cold Storage Creameries Magnolia brand milk and

cream bottles as culture flasks for orchid seed cultures). The Japanese assembled the

European population on the next morning and decided who was to be interned. On

February 17th they assembled the Indian and Malay regiments and urged them to

renounce the British crown and transfer their allegiance to the Emperor of Japan.

A few did. Most did not and eighty Malay officers were executed for their refusal

to renounce the crown.

The next step for the Japanese was to stop the looting and then they made sure

the waterworks, gas and electricity workers would report to work. After that they

drowned or machine gunned large numbers of Chinese. Massacres continued for

most of the first week. The number of victims is estimated to have been between

5,000 and 25,000. Mass killings stopped after the first two weeks. Beatings, torture,

arrests, degradation and some killings continued. Brutality was common. Those

arrested often did not know why. Diseases, hunger, food shortages and malnutrition

were part of daily life.

Japan surrendered formally on 15 August 1945, but the official announcement

was made in Singapore only on the 21st. British warships arrived on September Sth

and on the 12th of that month five Japanese generals and two admirals went up the

steps of the municipal building and surrendered formally to Admiral Lord Louis

Mountbatten, Supreme Allied Commander in South East Asia. The very same Union

Jack which came down in 1942 (and remained hidden during the occupation) was

hoisted over the city. This meant that the nightmare was over for most Singaporeans

(for more details about the Japanese occupation of Singapore see Turnbull 1977

from which I summarized information for this short account). But, ‘‘... in the in-

terest of science, one must distinguish carefully between the ‘Japanese’ of popular

conception and the Japanese men of science who in Malaya, at least, endeavoured

to serve science with impartiality’? (Corner 1946).

Koriba, a Japanese man of science, arrived in Singapore in December 1942, almost

a year after the surrender, and took charge of the Botanic Gardens and the forest

reserves. He had just retired from the Chair of Botany and as Dean of the Science

Faculty at Kyoto University. Professor E.J.H. Corner who had close contacts with

Koriba both during the occupation and after the war has suggested that the prospect

of tropical research must have appealed to the botanist, traveller, explorer, naturalist

and philosopher in Koriba (Corner 1981). He was in good health on his retirement

and had ‘‘... a remarkable flow of spirits. His retirement from the professorship

meant the start ... [of] ... a still greater field of activity ...’’ (Asida 1943) which

included a major research project (Koriba 1957), botanical excursions throughout

Singapore and scientific discussion with other botanists (Corner 1981).

On arriving at the Gardens Koriba first learned about the workings of the office.

Having done so he delegated the task to clerks. He also learned about the Library,

Herbarium and scientific policies ‘‘... for he wished to continue along the same

lines’? (Corner 1981). Koriba was given the rank of Brigadier General, but wore his

uniform only on ceremonial occasions (Holttum 1958) and detached the sword ex-

cept when required to wear it by official circumstances. He was also given a revolver

and shown how to use it. However he left the gun on his desk where no one else

touched it either. Koriba simply devoted himself to his administrative duties at the

Gardens and to botanical studies (Holttum 1958).

As Director of the Botanic Gardens Koriba was dependent on the military for

funds and various supplies and made every effort to secure whatever amenities were

possible for the staff. Even under the military rule Koriba ‘*... was on the side of

Nature and science ...’’ (Ashida 1958) and took ‘‘... energetic action to prevent

14 Gard. Bull. Sing. 42(1) (1989)

encroachment on the Nature Reserves of Singapore in which he took great interest’’

(Holttum 1958). He also took great pains to maintain the Herbarium and Library

intact (Holttum 1958). As a result Koriba was well liked by his staff and workers.

When Garden wages were very low in 1944, staff and labourers could get better pay

working for the military, but they did not leave the man they called ‘‘orang yang

baik sakali’’ (the perfect gentleman). Nor did they pilfer the cigarettes he left on his

desk when going home at 3:30 pm daily with a handkerchief full of books and a brief-

case loaded with notes. Koriba found a unique use for cigarette packages in that he

made cards from them. He used these cards to make notes for future books. Notes

on such cards and an incomplete manuscript on plant physiology, ecology and evolu-

tion remained in the Hirosaki University President’s Office following Koriba’s death

(S. Chiba, personal communication).

Prof. Koriba’s intervention saved the lives of those who remained in the Gardens.

Holttum and Corner resumed their work. Much less fortunate were 49 other mem-

bers of the staff, including J.C. Nauen, Assistant Curator of the Gardens from 1935

to 1939 and in charge of the Waterfall Gardens in Penang between 1939 and 1941.

They were sent to work on the Siam-Burma Railway where 22 of them lost their lives

and Nauen died as a result of blood poisoning in October 1943 (Anonymous 1947;

Tinsley 1983). When the food shortages in Singapore became acute Koriba or Dr.

Yata Haneda, Director of the Museum invited Corner and Holttum for lunch after

the Sunday excursions to make sure they had at least one good meal a week. One

can imagine Koriba at these lunches by looking at a photograph of him taken in

November 1956 (Fig. 8). During these lunches or at other times Koriba never spoke

of the war (Corner 1981).

Probably as a result of such actions confidence in Koriba never waned (Corner

1981) and in turn he stayed in Singapore to the very end even though he could have

left earlier. His last message to Prof. Corner at that time was dated 28 August 1945

and read in part (Corner 1981):

*‘Dear Dr. Corner,

I leave now the Gardens ... I regret very much that the Gardens and the Forest Reserves

were disformed and contaminated lately, though it was inevitable for me at present....

Hoping that the Gardens will flourish forever!

Best regards to Dr. Holttum....

Kwan Koriba’’

On 29 September 1957 shortly before his death he wrote to Corner from Hirosaki:

‘‘! feel urgent necessity to preserve natural forests of the tropical regions.’’ Koriba

clearly had a special relationship with Corner (Fig. 9). I think that theirs was a warm

relationshp between two people who liked and admired each other as people and

scientists. Corner described it best by writing (Chiba 1984a, 1984b):

**In the footsteps of Kwan Koriba

his one-time enemy

his adopted son

his admirer

E.J.H. Corner

Em. Professor of Tropical Botany

University of Cambridge

23 Oct. 1983”’

Koriba died ten weeks after his last letter to Corner. His funeral was held in

University Hall, Hirosaki University on December 21 1957 (Fig. 10), and the ‘‘...

beloved son of Mt. Hakkoda’’ is interred in the family graveyard which ‘‘faces Mt. —

Hakkoda, on the slopes of which the great botanist was brought up’’ (Ashida 1958).

Kwan Koriba

—

Fig. 8-10.

The last years and the aftermath. 8. Prof. Koriba at meal time in November 1956.

9. Professor E.J.H. Corner and Mrs. Helga Corner in front of Prof. Koriba’s photograph

(which is displayed among the photographs of other Presidents of Hirosaki University) in

October 1983. 10. Funeral rites for Prof. Kwan Koriba held on 21 December 1957 in University

Hall, Hirosaki University (courtesy Prof. Shigeo Chiba).

16 Gard. Bull. Sing. 42(1) (1989)

Prof. Koriba’s life was devoted to science and education and he had the ability to

inspire all those who came into contact with him (Ashida 1958). ‘‘He was a man that

commanded respect by his simplicity of life and unfailing devotion to duty, his very

broad scientific outlook, and also by his sympathetic understanding of human nature,

an understanding that transcended racial boundaries, even in times of war and of

privation’’ (Holttum 1958).

In one of his letters to me Prof. Ueyama wrote: ‘‘I always think: We must become

humanist[s] through the hard training of Academic study. I learned this from Holt-

tum, Corner and Koriba in Kew and Kyoto.’’ Prof. Ueyama’s thought is noble and

| keep thinking that under the circumstances in Singapore during the Japanese

occupation the lesson in humanism was taught by a devoted and great teacher, the

Perfect Gentleman, Brigadier General Professor Kwan Koriba, one of the illustrious

Directors of the Botanic Gardens.

Acknowledgements

I am indebted to several people for information and photographs. In chrono-

logical order I would like to thank Prof. John L. Mohr, University of Southern

California (USC), for teaching me to appreciate integrity in men of science; the late

Prof. Paul Saunders, also of USC for telling me about Prof. Kwan Koriba; Pro-

fessors P.N. ‘Danny’ Avadhani, and A.N. Rao, Botany Department, National

University of Singapore, for taking me to the Singapore Botanic Gardens on my first

visit there; Mr. Arthur George Alphonso, former curator and Acting Director of the

Singapore Botanic Gardens for sharing his war-time recollections; Prof. R.E. Holt-

tum, former Director of the Singapore Botanic Gardens, Professor of Botany

Emeritus, University of Singapore and now at Kew Gardens for providing informa-

tion, suggesting additional sources and for reading the first draft of this article;

Prof. Akinori Ueyama for information, photographs, copies of Koriba’s papers, very

kind letters and for reading the first draft; Prof. Shigeo Chiba for biographical data

and photographs and for reading the first draft; Prof. Shuji Tono, current President

of Hirosaki University, for his courtesy and assistance; Prof. Robert Ernst for

translating parts of Koriba’s papers and reading and commenting on the manuscript;

Prof. Franz Hoffmann for translating portions of Koriba’s papers; Dr. Leslie Paul

Nyman for reading and commenting on the manuscript and Mr. Kevin J. Hackett

for writing computer software which makes writing and the elimination of errors

easier.

References

Alessandrini, A. (1985). Le Orchidee spontanee dell’emilia-Romagna. Grafis Edizioni.

40033 Casalecchio di Reno (BO), Italy.

Anonymous. (1947). The Singapore Botanic Gardens during 1941-1946. The Gard.

Bull. 11: 263-265.

Anonymous. (1947). Mr. J.C. Nauen. The Gard. Bull. 11: 266.

Ashida, J. (1958). Kwan Koriba. Bot. Mag. Tokyo 72: 165-166.

Asida, Z. (1943). Professor Kwan Koriba. Bot. Mag. Tokyo 57: 100-103.

Chiba, S. (1984a). On Professor E.J.H. Corner’s visit to our University, Part 1. The

Gazette of Hirosaki University 64: 9-10.

Kwan Koriba |

. (1984b). On Professor E.J.H. Corner’s visit to our University, Part 2. The

Gazette of Hirosaki University 64: 4-8.

Corner, E.J.H. (1946). Japanese Men of Science in Malaya during the Japanese

Occupation. Nature 158: 63.

. (1981). The Marquis — A Tale of Syonan-to. Heinemann Asia. Singapore.

Desch, H.E. (1946). Letter. Nature 158: 513.

Gagnepain, F. and Guillaumin, A. (1933). Flore Générale de |’Indo-Chine, Volume

6, Fascicle 4 Orchidaceae. Masson et Cie, Paris.

Garay, L.A. and Sweet, H.R. (1974). Orchids of Southern Ryukyu Islands. Botanical

Museum Harvard University, Cambridge, MA.

Holttum, R.E. (1958). Kwan Koriba. The Gard. Bull. Singapore 17: 339-340 (the

paper is actually signed ‘‘R.E.H.’’).

____. (1964). A Revised Flora of Malaya. Volume I. Orchids of Malaya, 3rd Ed.

Government Printing Office, Singapore.

Koriba, K. (1913). Uber die Drehung der Spiranthes-Ahre. Ber. Deut. Ges. 31:

1957-165.

_____. (1914). Mechanisch-physiologische Studien uber die Drehung der Spiranthes

Ahre. J. Coll. Sci. Tokyo Imp. Univ. 36: 1-180.

___. (1926). Observations on a Japanese species of 7aeniophyllum. Proc. Third

Pan-Pacific Sci. Congress, Tokyo 2: 1900-1901.

Lin, T.-P. (1976). Native Orchids of Taiwan. Taipei, Taiwan.

Renz, J. (1978). Flora des Iranischen Hochlandes und der Umrahmenden Gebirge.

Orchidaceae. Akademische Druck- und Verlagsanstalt. Graz, Austria.

Seidenfaden, G. (1978). Orchid Genera in Thailand VI. Neottioideae. Dansk Botanisk

Arkiv. 32: 1-195.

Tinsley, B. (1983). Singapore Green. Times Books International. Singapore.

Turnbull, C.M. (1977). A History of Singapore, 1819-1975. Oxford Univ. Press,

Kuala Lumpur.

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An Atlas of Nutrient Deficiency Symptoms

in Hibiscus rosa-sinensis

T.W. FOONG, C.N. YANG and NASHITA BINTE MUSTAFA

Botanic Gardens

Parks & Recreation Department

Singapore

Abstract

Deficiency symptoms of essential nutrient elements were induced in Hibiscus rosa-sinensis in a nutrient-

omission trial in acid-washed sand. An atlas of deficiency symptoms was presented with description on

individual deficiency symptoms. Dry matter of above- and below-ground parts from control and deficient

plants as well as their foliar elemental contents were determined and compared.

Introduction

Hibiscus is noted for its colourful and attractive flowers. Owing to this trait, it

is highly regarded as a candidate for hedges, road-side planting and landscape in

parks, gardens and private homes.

In recent years, the Singapore Botanic Gardens has initiated a breeding programme

for Hibiscus resulting in the procreation of an array of hybrids with great ornamental

value. After screening their potential, selected hybrids are being introduced for mass

planting. There is, therefore, a need to better understand the nutrition of Hibiscus.

Nutrient deficiency symptoms have only been scantily described without specific

reference to Hibiscus (Anonymous 1979; Chin, 1986; Beers & Howie, 1986 and

Howie, 1980). The nutrient-omission trial was to establish the nutrient deficiency

symptoms in Hibiscus rosa-sinensis so as to provide a useful cross reference for

deficiency symptoms in the wide spectrum of possible hybrids. This would enable

Hibiscus breeders, growers and enthusiasts alike to trouble-shoot nutritional problems

in Hibiscus.

Materials and Methods

Sand was loaded onto a fine sieve and washed continuously with tap water to rid

soluble salts, silt, clay and other colloidal materials. Sand free of fine particles was

leached several times with IN HCI to remove adsorbed ions if any and rinsed with

deionised water to remove all residual acid. Plastic pots, approximately 20 cm in

diameter and 25 cm deep, were packed with the treated sand.

Uniform bag-grown Hibiscus plants about | m tall were selected. The root systems

were bared carefully and washed free of all soil contaminants. After rinsing with

deionised water, they were transplanted into the pots of sand. The potted plants were

left to establish under shade for a few days before being exposed to full sun in an

open cement ground.

Liquid nutrient formulations were according to Foong et al. (1982). Salinity of

20 Gard. Bull. Sing. 42(1) (1989)

each liquid feed was checked and the pH adjusted to S-5.5 with dilute HCI before

application. The control was raised with a complete nutrient feed whereas each treat-

ment was fertilised with a nutrient feed lacking the nutrient element in question.

Treatments and control were replicated six times. Fertilising was carried out every

alternate day at the rate of 200 ml per pot, the amount required to soak the pot

completely. On other days, 200 ml of deionised water was added to each pot.

Deficiency symptoms were recorded photographically as they became evident.

After 6 months, trial plants were harvested and divided into leaf, stem and root.

There was no flowering over the trial period. The three fractions were washed free

of contaminants. The foliar fraction was further subdivided into the mature and

young portions, washed with a non-ionic detergent and rinsed several times with

deionised water. All plant samples were dried at 80°C until constant weight for

dry matter analysis. Young and mature foliar samples were subjected to elemental

analysis. The mature fractions were used for the analysis of mobilisable nutrient

elements while the young fractions were used for the analysis of non-mobilisable

nutrient elements.

Total nitrogen was determined by the micro-Kjeldahl method, phosphorus by the

vanadate-molybdate-yellow method after dry ashing with magnesium nitrate, boron

by the Azomethine-H method after dry ashing with calcium oxide, and the remaining

elements by atomic absorption using the PU9000 Spectrometer after dry ashing and

dissolution of ash in dilute HCI (Cantle 1982; Chapman & Pratt 1961 and Evenhuis

1978). Sulphur and molybdenum were not determined.

Results and Discussions

Results of dry matter analysis and elemental analysis are tabulated in Table 1 and

Table 2 respectively. Statistical inference was based on the least significant difference

(Gomez & Gomez 1976). Nutrient deficiency symptoms are presented in Plates I to

VII and described as follows:-

(a) N Deficiency

Deficient plants were dwarfed. Leaves were very much reduced in size

compared with the counterparts of the control. Uniform chlorosis became

evident initially on the older leaves and slowly spread to the younger leaves in

advanced deficiency. Even the veins became chlorotic, giving a total chlorotic

appearance (Plate I). Premature leaf fall was common. Dry matter production

of above- and below-ground parts was significantly reduced in N deficiency

(Table 1). Foliar analysis showed a distinct difference in the N content between

control and deficient plants (Table 2).

(b) P Deficiency

Leaf size was not affected and deficiency symptoms were only mildly

expressed over the 6-month trial period. Deficient plants became slightly

chlorotic with chlorosis concentrating along the leaf margin (Plate II). Overall

dry matter production was curtailed (Table 1). Foliar P was markedly reduced

in deficient plants (Table 2) and should be used in conjunction with foliar

symptoms to confirm P deficiency in Hibiscus.

There were little tell-tale signs of K deficiency except for minor marginal

chlorosis as in the case of P deficiency (Plate III). Leaf size and plant growth

were altogether unaffected (Plate III and Table 1). However, there was a

se,

Nutrient Deficiency Symptoms in Hibiscus rosa-sinensis 21

pronounced reduction in foliar K (Table 2) and diagnosis of K deficiency

should be based on measurement of this parameter.

(d) Ca Deficiency

(e)

Symptoms were manifested as chlorosis in the young leaves followed by

necrosis, culminating in the die-back of growing tips (Plate IV). Gradually,

the deficient plants assumed a rather bare appearance. Both the foliar Ca

content and dry matter of affected plants were significantly lower than those

of the healthy control plants (Tables 1 & 2).

Mg Deficiency

In early Mg deficiency, the mature leaves exhibited yellow spotting mainly

in the middle of the lamina. As deficiency progressed, the yellow spots linked

to form chlorotic patches. Eventually, the whole blade became chlorotic with

the main veins retaining a light green tinge (Plate V). Reduction in the foliar

Mg content in Mg-deprived plants was associated with smaller leaves and

subdued growth (Tables 1 & 2).

(f) Fe Deficiency

Fe deficiency symptoms were characterized by interveinal chlorosis on the

young leaves especially on the newly-emerged leaves, resulting in a very distinct

““netting’’ effect (Plate VI). However, Fe contents in healthy and deficient

Treatment

Control

—N

—Pp

—K

= Oe)

— Mg

—Fe

—B

—Mn

—Cu

= Zn

—Mo

Table 1

Dry weights of above- and below-ground parts

Dry matter

of above-

ground parts

(g/pot)

Difference

from

Control

(48.8)**

(27.82)*

(13.52)

(45.03)**

(44.05)**

28.44

33.71

(29.82)**

41.34

24.24

44.73

56.91

Dry matter

of below-

ground parts

(g/pot)

Values tabulated are the averages of 6 replicates

Bracketed values are below those for the control

* Significant at P

** Significant at P

0.05

0.01

Difference | Total Dry

from

Control

(4.91)*

(4.92)*

(1.92)

(6.84)**

(8.29)**

8.65

3.68

(6.84)**

7.13

2.07

7.66

5.09

Matter

(g/pot)

Percentage

Difference

from

Control

(56.75)

(34.59)

(16.31)

(54.81)

(55.30)

39.19

39.51

(38.74)

a 7

27.80

55.36

65.51

Plate I: Foliar symptoms of Plate II: Foliar symptoms of

N deficiency P deficiency

Plate III: Foliar symptoms of Plate IV: Foliar symptoms of

K deficiency Ca deficiency

bs | :

Plate V: Foliar symptoms of Plate VI: Foliar symptoms of

Mg deficiency Fe deficiency

Plate VII: Foliar symptoms of

, B deficiency

Key: C connotes control

Nutrient Deficiency Symptoms in Hibiscus rosa-sinensis 23

Treatment

—Mg

Treatment

—Ca

—Fe

—B

—Mn

—Cu

—Zn

Table 2

Foliar elemental contents in deficient and control Hibiscus

Difference from

control

(%)

Treatment means

for mature foliage

(%)

Control mean for

mature foliage

(%)

(1.08)**

(O.27}*>

Fle 8 4 Lan

(0.29)*

Difference from

control

(ppm)

Treatment means

for young foliage

(ppm)

Control mean for

young foliage

(ppm)

(fzZ235}""

(35)

(21)**

(157)**

(5)**

Values tabulated are the averages of 6 replicates

Bracketed values are below those for the control

* Significant at P = 0.05

** Significant at P = 0.01

Deficient levels of elements do not necessarily connote the critical values at which symptoms just begin

to appear.

young leaves were similar (Table 2). It has been reported that iron chlorosis

in a number of plant species could only be resolved by examining the ‘‘active

Fe’’ fraction instead of the total Fe content (Foong & Yang 1987 and Takkar

& Kaur 1984). Hibiscus may well fall into this category. Dry matter accumula-

tion was also unaffected over the period of the trial (Table 1).

(g) Boron Deficiency

B deficiency symptoms were manifested at the growing points by a marked

reduction in the size of young leaves and very much shortened internodes,

generating the ‘‘rosette’’ appearance (Plate VII). Both dry matter and foliar

B were adversely affected by B deficiency (Tables 1 & 2).

No characteristic deficiency symptoms were induced in the omission of S, Cu,

Zn, Mn or Mo. In these cases, deficient plants performed as well as the control

plants (Table 1). Traces of these elements as remaining contaminants in the sand even

after cleansing could have sustained normal growth over the 6-month trial period.

Conversely, this finding may be taken to imply that Hibiscus is not very sensitive

to the deficiency of these trace elements. Nevertheless, plants deprived of Cu or Mn

did show a reduction in the foliar level of the element omitted (Table 2).

24 Gard. Bull. Sing. 42(1) (1989)

Conclusion

Deficiency symptoms of N, P, K, Ca, Mg, Fe and B were successfully induced

in Hibiscus rosa-sinensis cultured in sand and further confirmed by foliar analysis.

The characteristic symptoms of N, Ca, Mg, Fe and B deficiency can be employed

with a high degree of certainty as diagnostic tools. Symptoms of suspected P or K

deficiency are, however, best ascertained by foliar analysis.

Fe deficiency is by far the most commonly observed in field- and pot-grown

Hibiscus as a result of overliming or inadvertent overdoses of phosphatic fertiliser.

References

Anonymous, (1979). What every Hibiscus grower should know. American Hibiscus

Society.

Beers, L. & Howie, J. (1986). Growing Hibiscus. Kangaroo Press, Australia.

Cantle, J.E., ed. (1982). Atomic absorption spectrometry. Elsevier Scientific

Publishing Company, N.Y.

Chin, H.F. (1986). The Hibiscus — Queen of Tropical Flowers. Tropical Press Sdn.

Bhd. Kuala Lumpur.

Evenhuis, B. (1978). Simplified methods for foliar analysis. Communication 70,

Department of Agricultural Research, Product Research Division, Koninklijk

Institut voor de Tropen, Amsterdam.

Foong, T.W. & Yang, C.N. (1987). Resolving iron deficiency in Wrightia religiosa

by foliar analysis and its amelioration using an iron chelate as a soil additive.

Gard. Bull. 40(2), 133-137.

Foong, T.W., Ong, C.S. & Bakar, S.F.S.A. (1982). Induced deficiency symptoms

of nitrogen, phosphorus, potassium, magnesium and iron in Axonopus com-

pressus cultured in sand. Gard. Bull. 35(1), 33-44.

Gomez, K.A. & Gomez, A.A. (1976). Statistical procedures for agricultural research

with emphasis on rice. The International Rice Research Institute, Manila.

Howie, J. (1980). Hibiscus — Queen of the flowers. James Malcolm Howie, Australia.

Takkar, P.N. & Kaur, N.P. (1984). HCl method for Fe?* estimation to resolve iron

chlorosis in plants. J. Plant Nutr, 7(1-5), 81-90.

A New Nepenthes from Gunong Besar,

Kalimantan Selatan, Borneo

JUMAAT HAJI ADAM* and C.C. WILCOCK

Department of Plant & Soil Science, University of Aberdeen,

St. Machar Drive, Aberdeen AB9 2UD, Scotland, UK

EFFECTIVE PUBLICATION DATE: 15 MAR 1990

Abstract

One new species of Nepenthes, N. borneensis, from Gunong Besar in Kalimantan Selatan Borneo is

described.

Introduction

The new Nepenthes described is based on a single duplicate female specimen

distributed to L from KYO. The species according to the collectors note on the

herbarium label was found at an altitude of 1300-1880 m. The altitudinal zone falls

within the range of mossy forest in Kalimantan.

The species is related to N. boschiana Korthals by its prominent decurrent petiole

base, expanded peristome of upper pitcher and delicately wavy outer peristome

margin. The species, however, is distinguished from N. boschiana as listed in Table 1

below:

Table 1

The diagnostic characters of N. borneensis and N boschiana

N. borneensis N. boschiana

. Glandular crest on lower lid absent. Glandular crest on lower lid surface

prominent.

. Inside surface of upper pitcher wholly Inside surface of upper pitcher glandular

glandular. on lower ovate part only.

. Upper pitcher infundibulate. Upper pitcher + cylindrical in upper 2/3,

ovate lower 1/3.

. Peristome flattened. Peristome cylindrical.

. Tendril insertion to leaf tip sub- Tendril insertion to leaf tip apical.

peltate.

We have seen the type specimens of N. boschiana in L; and the original description

of the species by Korthals (1839). The differences between the 2 taxa indicate their

separation as distinct species.

* Permanent address — Jabatan Biologi, Fakulti Sains & Sumber Alam, Universiti Kebangsaan Malaysia

Kampus Sabah, Beg Berkunci No. 62, Kota Kinabalu 88996, Sabah, Malaysia.

26 Gard. Bull. Sing. 42(1) (1989)

Nepenthes borneensis Adam & Wilcock sp. nova. Plate 1, Fig. 1

Type: Murata, Kato & Mogea, B-3455 (HLB. 478518), Kalimantan Selatan, Gunong

Besar, alt. 1300-1880 m, 18.2.1979, holotype L.

Caulis angularis. Folia adulta alternantia, coriacea, petiolata; lamina lanceolata apex acutus sub-peltatus

RISKSHERBARIUM, LEIDEN

(Herb, Lugd. Bat.)

BEESUBLIMA TiSckaus

Ne 475518

{eta iy

780 300

280 270 280

il HMMA NNN MT MT MM

UT TL

ey cg gu

= ~~

EX HERBARIO UNIVERSITATIS KYO =

FLORA OF BORNEO Ny.

L4DIO ONERSIN BCTAIOCD. LAOEIATON TO KML OMANIAN. 1044-9

a ug ORAS WU THE WEBEL BOG52 £898

Lele,

Loo, Kalimantan Salstan: Guaang Basar. 5999-1980 mm alt,

Loan nr.

Hab. is svergoan mossy forest v

Cate Fab. 18, 1973

Coli, Gen Murata, Masshiro Kero & Yohene # Manes

5 44 i No. & 3455

Plate 1 Nepenthes borneensis Adam & Wilcock. From Murata, Kato & Mogea, B-3455. Upper stem

with leaves and pitchers.

New Nepenthes from Gunong Besar

CMTC

Fig. 1. | Nepenthes borneensis Adam & Wilcock. From Murata, Kato & Mogea, B-3455. Upper stem

with leaves and pitchers.

28 . Gard. Bull. Sing. 42(1) (1989)

(<1 mm), basis obtusa; petioli canaliculati alati, semi-amplexicaules decurrentes; nervi longitudinales

utroque latere 4, provenientes e variis locis costae; nervi pinnati inconspicui vel distincti, subhorizontales.

Ascidia superiora infundibuliformia, costis 2 prominentibus; pagina interior tota glandulosa; oris pars

anterior horizontalis pars posterior elevata posterioribus elevatum; peristomium applanatum, breviter

expansum, margo exterior breviter sinuatus; operculi facies inferior sine appendice basali glandulosa.

Inflorescentia feminei racemosa.

Upper stem angular, 9 mm thick, internodes 1-3 cm long, buds 5 mm long c.

10 mm above the node. Leaves of upper stem coriaceous, alternate, petiolate; lamina

lanceolate, 9-20 x 2-5 cm, apex acute, tendril insertion on leaf apex sub-peltate

(<1 mm), base obtuse; petiole 3-5 cm long canaliculate, narrowly winged, semi-

amplexicaul, decurrent into 2 opposite wings, extending c. 3/4 of the length of the

internode; longitudinal nerves 4 on each side, originating on lower half of the lamina

and from different points on the midrib; pinnate nerves distinct in inner 1/3 then

obscured or inconspicuous; tendrils of pitchers curled near the tip, longer than

lamina, 18-25 cm long, tendrils without pitchers not curled normally shorter than

lamina.

Upper pitchers infundibulate, 13-17 cm high, 4-5 cm wide immediately below the

mouth, with 2 prominent ribs running the entire length; mouth almost horizontal

in front, elevated towards the lid; peristome flattened and slightly expanded, outer

margin delicately wavy, 3-12 mm thick, broader in the middle, peristome ribs

0.2-0.5 mm apart, inside surface of the pitcher wholly glandular with numerous

minute glands; lids sub-orbiculate, base cordate, glandular crest absent, inner sur-

face almost wholly glandular with numerous, small, round-elliptic and rimmed

glands. Spur simple, flattened, 4-7 x 1-1.5 mm. Female inflorescence racemose,

peduncle 24 cm; sepals 4. Male inflorescence unknown. Fruits unknown.

Indumentum of stem of short whitish branched and unbranched hairs, denser in

the younger parts. Upper leaf surface glabrous, midrib covered with dense long

whitish branched and unbranched hairs; lower leaf surface glabrous with numerous

small dark spots. Tendrils densely hairy when young. Young pitcher densely covered

with short branched, and long branched and unbranched hairs, becoming glabrous.

Inflorescence pubescent.

Distribution: Borneo, Gunong Besar only.

Altitudinal range: 1300-1880 m.

Acknowledgements

We wish to thank Universiti Kebangsaan Malaysia & Dept. of Plant & Soil

Science, University of Aberdeen, for financing the project; Rijksherbarium (L) for

the use of the specimen; Mr. T.E.V. Pearce of Classics Department, University of

Aberdeen for the Latin description; Mr. N. Little for the photographs taken.

References

Danser, B.H. (1928). The Nepenthaceae of the Netherlands Indies. Bull. Jar. Bot.

Buit. Serie HI, Vol. IX, LIVR. 3-4: 249-428.

Korthals, P.W. (1839). Over het geslacht Nepenthes. In C.J. Temminck, Verhandel-

ingen over de natuurlijke geschiedenis der Nederlandsche overzeesche bezittingen,

p. 25-28.

Kurata, S. (1976). Nepenthes of Mt. Kinabalu. Sabah National Parks Publication,

No. 2. Sabah National Parks Trustees.

An Enumeration of One Hectare of Pantai

Aceh Forest Reserve, Penang

IAN TURNER*

Oxford Forestry Institute

Department of Plant Sciences

University of Oxford

Abstract

One hectare of forest in Pantai Aceh F.R. was enumerated for stems >10 cm d.b.h., and toa 1 cm

d.b.h. lower limit in a 0.05 ha subplot.

The large plot was found to contain 374 stems belonging to 70 or so species. The Dipterocarpaceae

was the most abundant family (34% of stems >10 cm d.b.h. Shorea curtisii dominated the largest size

class (> 30 cm d.b.h.) with 48 out of 81 stems. The most numerous species in the sub-canopy were: Shorea

maxwelliana, Porterandia anisophylla and Pternandra coerulescens.

The floristic composition indicates that Pantai Aceh is a coastal hill dipterocarp forest of the Shorea

curtisii-Eugeissona tristis segregate.

Analyses of stand tables, together with the known history of logging on the site, lead to the conclusion

that the composition of the forest is changing. More shade-tolerant dipterocarp species such as Shorea

maxwelliana are taking over the upper canopy as Shorea curtisii fails to regenerate.

Introduction

Investigations into the seedling ecology of tree species in Pantai Aceh Forest

Reserve, Penang, Peninsular Malaysia (Turner 1988) made it evident that there were

considerable differences in the demography of seedlings of various Shorea species.

Shorea curtisii seedlings showed a very high mortality in their first year (93%),

significantly higher than in the more shade-tolerant Shorea multiflora (72%). It was

therefore decided to look in detail at a small area of forest to see if these differences

in present-day regenerative capability were reflected in the composition and structure

of the forest.

Materials and Methods

One hectare of forest on the east-facing hillside behind the Muka Head Field

Station of Universiti Sains Malaysia was enumerated in November 1987. A 100 m

x 100 m square was marked out by walking on a bearing for 100 metres, then on

a perpendicular bearing for another 100 m and so on. The end of the fourth side

was some 10 m out from the starting point; to compensate for this the line was

angled in from the 70 m point on this fourth side. The plot is, therefore, only an

approximation for one hectare.

* Present Address: Department of Botany, National University of Singapore, Lower Kent Ridge Road,

Singapore 0511.

30 Gard. Bull. Sing. 42(1) (1989)

All trees within the plot were measured for girth at breast height and those 32 cm

or greater (1.e. > 10cm d.b.h.) were tagged and identified as far as possible by using

binoculars and collecting and matching fallen leaves.

In addition on five 10m x 10m sub-plots along the lower edge from the north-

east corner all stems of 1 cm d.b.h. or greater were enumerated and all dipterocarp

seedlings were tagged and measured for height.

Results

A total of 374 stems with d.b.h. =>10 cm were recorded from the 1 ha plot;

comprising some 65 recognised, but not in all cases named, species (Table 1). Twenty-

six stems remained indetermined because no distinguishing features could be seen

from the ground so the appropriate fallen leaves could not be picked out from the

litter. This group may contain up to ten extra species, but five or six is probably a

fairer estimate, thus giving an overall species count of approximately seventy.

The forest is dominated by the Dipterocarpaceae: 34% of all stems greater than

10 cm d.b.h. (see Table 2 and Fig. 1) with the Rubiaceae the next most abundant

family with 16%. The Dipterocarpaceae are also represented by the highest number

of species, eleven in all. As is usual with tropical rain forests many species are rare

(Jacobs 1988, Whitmore 1988), in this case more than two-thirds of the species are

represented by under 1% of the total number of stems each.

The nature of the forest is elucidated by breaking down the results by size class

(Class 1 = 1-10 cm d.b.h., Class 2 = 10.1-20 cm d.b.h., Class 3 = 20.1-30 cm

d.b.h. and Class 4 = >30cm d.b.h.). Class 4, the canopy-top trees, is dominated

by the dipterocarp Shorea curtisii with 48 out of 81 stems. Gluta curtisii (Anacar-

diaceae) is the only other species to make up more than 10% of the total in this class.

Dipterocarpaceae — 34.2%

Anacardiaceae — 8.3%

Guttiferae — 4.5%

Indetermined — 7.5%

Rubiaceae — 16.3%

Others — 14.7%

‘Melastomataceae — 10.2% '

Euphorbiaceae — 4.3%

Fig. 1. Relative abundance of families for all stems >10 cm d.b.h. on | ha of forest in Pantai Aceh

F.R., Penang.

Enumeration of One Hectare of Pantai Aceh Forest Reserve

A list of the tree species found on 1 ha of hill forest in Pantai Aceh F.R.

Table 1

(Class 2: 10.1-20 cm d.b.h., Class 3: 20.1-30 cm d.b.h., Class 4: >30 cm d.b.h.)

Species

Shorea curtisii Dyer ex King

Shorea maxwelliana King

Porterandia anisophylla

(Jack ex Roxb.) Ridley

Pternandra coerulescens Jack

Gluta curtisii (Oliv.) Ding Hou

Timonius corneri K.M. Wong

Shorea multiflora (Burck) Sym.

Elaeocarpus sp.

Glochidion superbum Baill.

Anisoptera curtisii Dyer ex King

Cryptocarya ferrea Bl.

Gironniera parvifolia Planch.

Calophyllum calaba L.

Eugenia sp.

Garcinia sp. A

Gluta aptera (King) Ding Hou

Homolanthus populneus

(Geisel.) Pax

Swintonia spicifera Hk.f.

Austrobuxus nitidus Miq.

Calophyllum teysmannii Miq.

Calophyllum wallichianum

Planch. et Triana

Kokoona reflexa

(Laws.) Ding Hou

Psydrax sp. 1

Shorea ovata Dyer ex Brandis

Shorea sp.

Vatica sp.

2311

Artocarpus lowii King

Brackenridgea hookeri

(Planch.) A. Gray

Buchanania arborescens (Bl.) Bl.

indet.

Calophyllum molle King

Dacryodes longifolia (King) Lam

Dillenia grandifolia

Wall. ex Hk.f. et Thoms.

Elaeocarpus ferrugineus Steud.

Eugenia attenuata

(Miq.) Koord. & Val.

Family

Dipterocarpaceae

Rubiaceae

Melastomataceae

Anacardiaceae

Rubiaceae

Dipterocarpaceae

Elaeocarpaceae

Euphorbiaceae

Dipterocarpaceae

Lauraceae

Ulmaceae

Guttiferae

Myrtaceae

Guttiferae

Anacardiaceae

Euphorbiaceae

Anacardiaceae

Euphorbiaceae

Guttiferae

Celastraceae

Rubiaceae

Dipterocarpaceae

unknown

Moraceae

Ochnaceae

Anacardiaceae

Burseraceae

Guttiferae

Burseraceae

Dilleniaceae

Elaeocarpaceae

Myrtaceae

Class 2 Class 3 Class 4 TOTAL

WwWNe NPN DN NHN Ff

— W Ww ho

mes

h@ S&S BRA aa aA =a &

ive) ww ph f

NO WwW WwW Ww Ww

NNN WN N

Species

Gardenia tubifera Wall.

Gluta elegans Kurz

Hopea pedicellata (Brandis) Sym.

indet.

Lithocarpus wallichianus

(Lindl. ex Hance) Rehd.

Ryparosa sp.

Symplocos adenophylla

Wall. ex Don

Rengas sp.

Aporusa benthamiana Hk.f.

Artocarpus sp.

Bhesa paniculata Arn.

Buchanania sessifolia Bl.

Canarium littorale Bl.

Canarium sp.

Canthium confertum Korth.

Dacryodes rugosa (Bl.) Lam

Dipterocarpus caudatus Foxw.

Dipterocarpus grandiflorus

(Blanco) Blanco

Drypetes sp.

Garcinia sp. B

Gordonia penangensis Ridley

Heritiera sumatrana

(Mig.) Kosterm.

Hopea beccariana Burck

Mallotus sp.

indet.

Payena obscura Burck

indet.

Schoutenia accrescens

(Mast.) Curtis

Indetermined

TOTAL

(Total number spp identified)

Table 1 (contd)

Family

Rubiaceae

Anacardiaceae

Dipterocarpaceae

Lauraceae

Fagaceae

Flacourtiaceae

Symplocaceae

Anacardiaceae

Euphorbiaceae

Moraceae

Celastraceae

Anacardiaceae

Burseraceae

Rubiaceae

Burseraceae

Dipterocarpaceae

Euphorbiaceae

Guttiferae

Theaceae

Sterculiaceae

Dipterocarpaceae

Euphorbiaceae

Myristicaceae

Sapotaceae

Rosaceae

Rubiaceae

Tiliaceae

Gard. Bull. Sing. 42(1) (1989)

Class 2 Class 3 Class4 TOTAL

m= NN NY

a ee ee

(56)

= oes 2

= oe 2

at = 2

1 a 2

2 = 2

Wa = 2

oe ae 2

bse we I

“i al I

= I I

~ I 1

a Ws I

ch sae 1

= 1 1

ee =. I

at so 1

1 ~ I

ae 5 1

a a I

at a 1

in a} I

= en 1

aa I 1

as 1

= a 1

at Bre I

ate tse I

= ne I

sa os I

3 3 26

51 81 374

(21) (18) (65)

Enumeration of One Hectare of Pantai Aceh Forest Reserve 33

Table 2

The families of trees found on | ha of coastal hill

dipterocarp forest (stems >10 cm d.b.h.)

Family Number of stems Species per Family

Dipterocarpaceae 128 1

Rubiaceae 61

Melastomataceae 38

Anacardiaceae 31

Guttiferae 17

Euphorbiaceae 16

Burseraceae

Elaeocarpaceae

Lauraceae

Myrtaceae

Ulmaceae

Celastraceae

Moraceae

Dilleniaceae

Fagaceae

Flacourtiaceae

Ochnaceae

Symplocaceae

Myristicaceae

Rosaceae

Sapotaceae

Sterculiaceae

Theaceae

en ee ee oe ee LO ee DO ee b> © > © en © i ©

ee ee ee ee ee Oe © ee Oe © en © O° mG ©, ao) i. i. a |

Tiliaceae

Indetermined

TOTAL 374

The subcanopy has a markedly different composition, with Shorea maxwelliana,

Porterandia anisophylla (Rubiaceae) and Pternandra coerulescens (Melastomataceae)

being the commonest species. This is reflected in the familial composition of Class 2

(Fig. 2) with the Rubiaceae the most abundant (22.5%) followed by the Dipterocar-

paceae (17%) and the Melastomataceae (15%). Porterandia anisophylla is common

in the small gaps formed in the canopy where the large trees die standing. Raich

(1987) has shown that Porterandia anisophylia is restricted to canopy gaps at Pantai

Aceh and that its seeds germinate better in a small gap compared to a closed-canopy

site or large clearing.

The stand table for Shorea curtisii (Fig. 3) is heavily skewed to the biggest size

Gard. Bull. Sing. 42(1) (1989)

‘Sueudd “YA Yyaoy lequed Ul jsasOj JO BY [ UO ("Y'Q’p WD OZ< = pH-£

sseID “U'G'P WO OZ-I'OI = Z SSB[D) SASSBID OZIS JUSJAJJIP Ul SaI[IwWey Jo saouepuNge oaneey “7 “SI4

“Gy — a0sa}1}}N5 %6°7) — 90800IPsDO0UY %6°02 — SJ24}0

% — aoa0Igny

%) — 90390}DUU0}SOja/\

YS" | — adao01qs0ydn}

%8°6 — Ss2UI0

%S'S - aDdad0Iquoudn}

= %SL — eda00}DWO}SDjaK

lik

ZL°6 — Peuluejapul

ZB'C — Peulwe}apul

25°70 — 8D9DIGNY —

%y'p9 — aDa20d1090I0)0iq “GG — 9D90/PIOOOUY

“LY — B0JOPN}NY

Wee OSV G Se

Enumeration of One Hectare of Pantai Aceh Forest Reserve 35

Ss Ss Ss

—

Number of stems in tha

() ae a N K K s N WS 0

S. curtisit S. maxwelliana P. anisophylla

Fig. 3. Stand tables for the three commonest species, Shorea curtisii, Shorea maxwelliana and

Porterandia anisophylla from 1 ha of forest in Pantai Aceh F.R., Penang. Bars from left

to right represent Class 2 (10.1-20 cm d.b.h.), Class 3 (20.1-30 cm d.b.h.) and Class 4

(>30 cm d.b.h.).

class whereas Shorea maxwelliana has more small trees than large. No individuals

of Porterandia anisophylla reach the largest size class.

The more detailed enumeration of the 0.05 ha area shows that the understorey,

Class 1, has a more mixed composition (Table 3 and Fig. 4), the Guttiferae being

the commonest family. In addition to the 190 tree and shrub stems recorded in

Class 1 there were 7 stems of rattans and other woody climbers with a d.b.h. greater

than | cm. Large climbers do not appear to be abundant at Pantai Aceh. There

were 62 clumps of the stemless palm, bertam (Eugeissona tristis Griff.) in the 0.05 ha

area, each with a mean of 6.1 leaves (standard error = 0.5). Only two clumps

showed signs of having ever flowered.

The stand table (Fig. 5) for the 0.05 ha shows the usual reverse-J curve of a

primary rain forest with many small saplings and fewer large trees. Fig. 6 is a species-

area curve for the five 100 m? sub-plots for all stems >1 cm d.b.h. The points

represent the average of five random compilations of the data from the five sub-

plots. The curve is still rising at 500 m?.

A total of 379 individual seedlings of six species of dipterocarp were recorded

from the five 100 m? sub-plots (Table 4). The commonest was Hopea beccariana

which had a greater mean height than the other two common seedlings, Shorea

curtisii and Shorea multiflora. Many of the Shorea curtisii seedlings were from the

fruiting of September 1986.

Gard. Bull. Sing. 42(1) (1989)

A list of species found in Class 1 (1-10 cm d.b.h.)

on 0.05 ha of Pantai Aceh F.R., Penang

Species

Urophyllum glabrum Wall.

Calophyllum wallichianum Planch. et Triana

Hopea beccariana Burck

Ardisia korthalsiana Scheffer

Brackenridgea hookeri (Planch.) A. Gray

Eugenia attenuata (Miq.) Koord. & Val.

Pternandra coerulescens Jack

Swintonia spicifera Hk.f.

Gluta elegans Kurz

Gironniera parvifolia Planch.

Mesua kunstleri (King) Kosterm.

Rhodamnia cinerea Jack

Austrobuxus nitidus Miq.

Garcinia eugenaeifolia/rostrata

Macaranga lowii King ex Hk.f.

Microtropis bivalvis (Jack) Wall.

Canarium littorale Bl.

Eugenia sp. B

Shorea curtisii Dyer ex King

Shorea maxwelliana King

Shorea ovata Dyer ex Brandis

Anisophyllea corneri Ding Hou

Ardisia sp.

Baccaurea sumatrana (Miq.) M.A.

Calophyllum teysmannii Miq.

Cryptocarya ferrea Bl.

Eurycoma longifolia Jack

Garcinia sp.

Guioa fuscidula (Kurz) Radlk.

Hopea pedicellata (Brandis) Sym.

Lophopetalum floribundum Wight.

Semecarpus sp.

Shorea sp.

Timonius corneri K.M. Wong

Aidia cochinchinensis Lour.

indet.

Arthrophyllum diversifolium Bl.

Barringtonia macrostachya (Jack) Kurz

Buchanania sessifolia Bl.

Dacryodes longifolia (King) Lam

Eugenia sp. A

Garcinia malaccensis/penangiana

Gluta aptera (King) Ding Hou

Family

Rubiaceae

Guttiferae

Dipterocarpaceae

Myrsinaceae

Ochnaceae

Myrtaceae

Melastomataceae

Anacardiaceae

Ulmaceae

Guttiferae

Myrataceae

Euphorbiaceae

Guttiferae

Euphorbiaceae

Celastraceae

Burseraceae

Myrtaceae

Dipterocarpaceae

Rhizophoraceae

Myrsinaceae

Euphorbiaceae

Guttiferae

Lauraceae

Simaroubaceae

Guttiferae

Sapindaceae

Dipterocarpaceae

Celastraceae

Anacardiaceae

Dipterocarpaceae

Rubiaceae

Annonaceae

Araliaceae

Lecythidaceae

Anacardiaceae

Burseraceae

Myrtaceae

Guttiferae

Anacardiaceae

Number of stems

=e eee KR hh Kh KRCULU MU NNN NNNNNNNN WN WWW WwW Lf HL HL Hh NN NA ~I C CO CO CO CO

Enumeration of One Hectare of Pantai Aceh Forest Reserve 37

Table 3 (cont'd)

Species Family Number of stems

Gluta curtisii (Oliv.) Ding Hou Anacardiaceae l

Gordonia penangensis Ridley Theaceae l

Heretiera sumatrana (Miq.) Kosterm. Sterculiaceae l

Hydnocarpus sp. Flacourtiaceae l

Kokoona reflexa (Laws.) Ding Hou Celastraceae 1

Lepisanthes senegalensis (Poir.) Leenh. Sapindaceae l

Lithocarpus wallichianus (Lindl. ex Hance) Rehd. Fagaceae l

Mallotus sp. Euphorbiaceae ]

Mangifera sp. Anacardiaceae l

Pandanus sp. Pandanaceae l

Porterandia anisophylla (Jack ex Roxb.) Ridley Rubiaceae 1

Psydrax sp. 1 Rubiaceae l

Schoutenia accrescens (Mast.) Curtis Tiliaceae 1

Streblus elongatus (Miq.) Corner Moraceae 1

Teijsmanniodendron coriaceum (Clarke) Kosterm. Verbenaceae ]

Ternstroemia wallichiana (Griff.) Engler Theaceae l

Tristaniopsis merguensis

(Griff.) Wilson & Waterhouse Myrtaceae ]

Vatica sp. Dipterocarpaceae l

Indetermined -

TOTAL 190

Lianas and rattans 7

Dipterocarpaceae — 12.2% Anacardiaceae — 10.7%

Melastomatacece — 4.1%

Guttiferae — 13.7%

Rubiaceae — 9.6%

Indetermined — 2%

Euphorbiaceae — 5.6%

Others — 42.1%

Fig. 4. Relative abundance of families in Class 1 (1-10 cm d.b.h.) from 0.05 ha of forest in Pantai

Aceh F.R., Penang.

Mi6es-es

Wessss

K6Sr-SP

Gard. Bull. Sing. 42(1) (1989)

H6ee-ce

16 2-2

V6 12 +2

681-81

iB SI-Si

6 ri-vi

PASCI-E1

PAGzI-21

(ZZA6 hh

PAG OI-O1

466-6

VZZZZL60-8

(ZZA6 L-L

PZA69-9

(ZZZLLLL ALLL LAOS S

(ZLIZLLLAAALAALL LALLY

CZZLLLLLLL LALLA ALLA AMAAMAAMAAA LALLA LOE E

(ZZZZZIL LLL LLL ALLL LALLA ALLL ALLL ALLA LALLA ALAA ALLL LLM Oe

UZZ77L LLL ALLA LLAMA NAA ALAS Sh

=) jam) a) Cc (=) co) ja) _

™ co) T=) =e Lae — —

Swa}s JO Jaquinu

1cm increment classes

d.b.h

Stand table for all stems >1 cm d.b.h. on 0.05 ha of forest in Pantai Aceh, F.R., Penang.

Fig. 5.

S Ss —

saidads 40 Jequun\

AREA m?

Species-Area curve for all stems >1 cm d.b.h. from 0.05 ha of forest in Pantai Aceh, F.R.,

Penang.

Fig. 6.

Enumeration of One Hectare of Pantai Aceh Forest Reserve 39

Table 4

The numbers and heights of dipterocarp seedlings

(<1 cm d.b.h.) on 0.05 ha of forest

Species No. indivs mean height + s.e. (cm)

Hopea beccariana 124 14: 7]

Shorea curtisti 108 a0 <3

Shorea multiflora 100 22 tk

Shorea maxwelliana 30 82 +9

Hopea pedicellata 2 ry +86

TOTAL 379

Discussion

The enumeration has shown that the forest at Pantai Aceh is made up of Shorea

curtisii with some Gluta curtisii, overtopping a subcanopy of Shorea maxwelliana,

Pternandra coerulescens and various other Rubiaceous and some Euphorbiaceous

trees; with Porterandia anisophylla being common in the small canopy gaps. The

understorey is rife with bertam; with tree seedlings and saplings, mostly those of

dipterocarp species, occupying the spaces between the clumps. The floristic composi-

tion is clearly that of the subformation of coastal hill forest known as the Shorea

curtisii-Eugeissona association-segregate as defined by Symington (1943) which was

later called seraya-ridge-forest (Wyatt-Smith 1963). Wyatt-Smith (1963) pointed

to Anisoptera curtisii, Calophyllum teysmannii var. inophylloide, Calophyllum

wallichianum, Hopea beccariana and Shorea multiflora as species characteristic of

this subformation. They are all common at Pantai Aceh.

In general Pantai Aceh comes out as being species-poor in comparison with other

lowland forests in Peninsular Malaysia. Fig. 7 compares Pantai Aceh with the data

given by Wyatt-Smith (1963) for various coastal hill forests and that from Kemasul

F.R. (Wong 1967) and the species-area curve from Sungei Menyala (Wyatt-Smith

1966) as examples of lowland forest. It becomes clear that coastal hill forest is species-

poor compared with lowland forest and that Pantai Aceh is highly depauperate as

coastal hill forest goes. The area of the Pantai Aceh plot may not be accurately

known but it cannot be overestimated to the extent to bring Pantai Aceh towards

the same level of species-richness as the other plots. The reasons for this poverty are

not obvious but may involve the climatic and topographic propensities of the site

to drought.

The most interesting comparator for this data set would be the work of Wong

(1987) for Bukit Timah Nature Reserve, Singapore; another example of coastal hill

forest. Unfortunately the data are not directly comparable because those from Bukit

Timah involved a systematic inventory with regularly spaced clusters of circular plots

rather than the contiguous plots generally used for ecological enumerations of forest.

Also the published data consist largely of lumped data from plots where different

measuring criteria were used. However, it is clear that Bukit Timah is considerably

richer in species than Pantai Aceh (Bukit Timah has c. 80 species per hectare trees

>20 cm d.b.h. whereas Pantai Aceh has only c. 30). Shorea curtisii, whilst the

commonest large tree at Bukit Timah, is not as abundant there as at Pantai Aceh

but it is regenerating successfully (Wong & Teo 1978); there are no heavy-hardwood

40 Gard. Bull. Sing. 42(1) (1989)

IN Sunget Menyalo

ri Kemesul FR. es

2 I ea

a —

2 IN a

= tae Pali doh ER

/ % Coastal Hill Forest

De Db orllie cai tle iileediee a: ean

Area. (ha]

Pigac; Species-Area plots for various Malayan forests. Data from Pulau Jarak, Pulau Rumbia,

Pulau Pangkor, Kampong Gajah F.R., Banang F.R., Soga F.R., Gunung Raya F.R. and

Bukit Enggang F.R, (Wyatt-Smith 1963); Sungei Menyala F.R. (Wyatt-Smith 1966) and

Kemasul F.R. (Wong 1967).

Shorea species i.e. the Balau group (Shorea section Shorea) such as Shorea max-

welliana, and the Euphorbiaceae rather than the Rubiaceae dominate the understorey.

The most notable points to come out of the Pantai Aceh data concern comparisons

between the stand tables of the two common dipterocarps, Shorea curtisii and Shorea

maxwelliana. Shorea curtisii exhibits a stand table with few middle-sized trees com-

pared to the canopy-top individuals whereas Shorea maxwelliana has few big trees

and many small ones. A regeneration sampling carried out in Compartment 6 of

Pantai Aceh in 1949 (Wyatt-Smith 1963) also showed a difference in stand structure

between dipterocarp species, though different sizes were considered. A milliacre

sampling technique was used which included all trees <one foot g.b.h. (c. 10 cm

d.b.h.). The results are plotted in Fig. 8 with the individuals divided into four size

Enumeration of One Hectare of Pantai Aceh Forest Reserve 41

Shorea curtistt Shorea glauca « Shorea marwell‘ona

Class A = < 1.5m tal

Class B = 1.5m < 3m (al

Class C= 3m loll < Sem dbh

Class ) = Sem dbh < 10cm dbh

Frequency %

Chass A Chiss B Chis C Closs 0

Fig. 8. Results of a linear regeneration sampling of Pantai Aceh F.R. in 1949 (from Wyatt-Smith 1963).

classes. The two heavy hardwood Shorea species, Shorea glauca and Shorea max-

welliana, were well represented in the advanced regeneration of large seedlings,

saplings and poles while Shorea curtisii, a light hardwood, had many small seedlings

but very few individuals in the larger size classes.

Foresters generally regard the wood density of dipterocarps as a reflection of their

relative regeneration capabilities or ‘tolerance’. Those of a high density, the heavy

hardwoods, are known to be very shade tolerant as juveniles; persisting and growing

at lower radiation levels than the light hardwood species e.g. as Becker (1983) found

in a comparison of Shorea maxwelliana and Shorea leprosula seedlings in Pasoh

F.R., Peninsular Malaysia. Silviculturally this means that the light hardwoods require

greater canopy opening (disturbance) to release the seedlings from the suppressed

state in the deeply shaded forest understorey than the heavy hardwoods, but respond

more rapidly when released (Whitmore 1984).

At present there is more demand for light hardwood timber but in the past,

before the advent of chemical wood-preservatives, the heavy hardwoods found

greater favour. As Curtis (1894) wrote in the first flora of Penang:

“*Shorea utilis King [Shorea maxwelliana] ‘Damar laut No[mbor] satu’ Undoubtedly the

best timber in the island [Penang]. Waterfall and Muka Head, too much in demand to be

common’’.

The naturally strong and durable timbers of the Balau group were put to many uses

especially heavy structural work such as bridge-building (Desch 1941). Pantai Aceh

was exploited for its heavy hardwoods from around 1915 (and probably before) to

the late 1930s (Ong & Dhanarajan 1976); though felling of such timber for boat-

building may have been going on long before this (J. Wyatt-Smith pers. comm.). The

large Balau trees were felled selectively and dragged out by man or buffalo. A few

signs of this activity are still evident in the forest in the very occasional tree stump

or gully caused by the erosion of a drag line.

It would appear from the stand tables that Pantai Aceh is in a state of composi-

tional change; that Shorea curtisii will be unable to replace itself in the canopy as

42 ~ Gard. Bull. Sing. 42(1) (1989)

the biggest trees die because there are insufficient small trees to grow to take their

place. The likely usurper is Shorea maxwelliana as it has a large number of middle-

sized (Class 2 and 3) trees. Generally it is inadvisable to draw deductions about forest

dynamics from stand tables (Whitmore 1984 p. 86) but in this case the evidence does

seem convincing.

1. The present day stand tables which show Shorea curtisii to be shade-intolerant,

with little regeneration, but Shorea maxwelliana to be tolerant with a strongly

positive stand table.

2. The 1949 regeneration sampling which showed few Shorea curtisii seedlings taller

than 1.5 m but a considerable number of large seedlings, saplings and poles of

the heavy hardwood Shorea species. This indicates that the regeneration of Shorea

curtisii has been problematic for at least 40 years.

3. The general perceived wisdom of nearly a century of practical forestry in the

Malay Peninsula which has developed a system of ecological groups amongst the

dipterocarps based on silvicultural responses with wood density as the shorthand

means of reference.

4. The known history of Pantai Aceh F.R., which experienced a period of light and

selective logging that ceased some fifty years ago.

A hypothesis must be put forward to answer the questions: How did Shorea curtisii

come to dominate the forest?, and; why did it regenerate successfully in the past but

fail to do so now?

The hypothesis presented here is built upon an understanding of the ecologies of

these two species (species groups) whose main difference lies in their responses to

canopy disturbance via the effects of understorey light levels. Whilst there are no

reports of major changes in the frequencies of fierce storms or other cataclysmic

natural phenomena there has, as has been stated above, been a change in anthro-

pogenic interference in that the selective felling of heavy hardwood species stopped

approximately fifty years ago. It can be hypothesised that the logging activities would

have produced canopy disturbance of a magnitude great enough to produce rapid

growth of the light hardwood species such as Shorea curtisii; which would have been

favoured over other light hardwoods because of its probable drought-tolerance

(Whitmore 1984) on the steep hillsides of Pantai Aceh (90% of the forest is on slopes

of more than 18° (Ong & Dhanarajan 1976)) with their coarse, granite-derived soils.

It would seem reasonable to deduce that the present abundance of Shorea curtisii

is the result of the pre-war selective felling of heavy hardwoods which both selected

against these species at maturity and gave the competitive advantage at the juvenile

stages to the light hardwoods because of increased illumination of the forest floor.

The cessation of logging would have reduced this artificial selection in favour of

Shorea curtisii. The canopy would have closed over and the average solar radiation

levels on the forest floor would fall, favouring the shade-tolerant heavy hardwoods

(mostly Shorea maxwelliana) over the light hardwoods. Therefore for the last fifty

years Shorea maxwelliana has successfully regenerated and grown to the small tree

size seen today but Shorea curtisii has failed because the radiation levels have been

too low. Burgess (1975) found that Shorea curtisii typically regenerates poorly from

seed and that its seedlings do not persist for many years in the shade of the forest

understorey. The abundance of bertam at Pantai Aceh, the proliferation of which

would also be favoured by logging, may also be deleterious to Shorea curtisii regenera-

tion, as Burgess showed in the hill forests he studied. Symington, in his seminal

work on the dipterocarps of the Malay Peninsula (1943), gives figures for the mean

annual girth increment of the two species; that for Shorea curtisii he gives as 1.54

inches year~! and for Shorea maxwelliana as 0.92 inches year~'. Therefore in fifty

Enumeration of One Hectare of Pantai Aceh Forest Reserve 43

years a tree of Shorea curtisii could grow to 60 cm d.b.h. but Shorea maxwelliana

would be some 40% smaller in diameter, at c. 35 cm d.b.h. Given the fact that the

growth rates on the drought-prone Penang coastal hills are probably below average

(Symington 1943), the present stand structure of the forest with large numbers of

Shorea curtisii stems around 40-50 cm d.b.h. and Shorea maxwelliana around 20-25

d.b.h. are a plausible reflection of a change in disturbance pattern about fifty years

ago.

It can be predicted that as long as Pantai Aceh remains in a state of low distur-

bance frequency and intensity Shorea maxwelliana will increase in abundance in the

largest size classes as the mature Shorea curtisii stand senesces. Possibly Shorea

curtisii may maintain some presence through rare, large disturbances allowing waves

of new regeneration. It may also hold its own on the ridge crests where its drought

tolerance is at the greatest advantage. The simplest way of testing this hypothesis

will be to see if its predictions come to pass, and it is to be hoped that the forest

will be allowed to develop naturally and be the subject of future studies.

Acknowledgements

This research was conducted with permission from the Malaysian Government

and assistance from the authorities of Universiti Sains Malaysia for which I am most

grateful. I was supported financially by a studentship from the Natural Environment

Research Council (UK). My thanks go to Drs T.C. Whitmore and Gong Wooi Khoon

for their supervisorial skills. Koay Tian Soo, Lynn Davy, Janet Simms and Wolfgang

Wiuster helped with the fieldwork and Wong Khoon Meng of the Forest Research

Institute of Malaysia with the identification of specimens. J. Wyatt-Smith kindly read

and commented upon a draft of this paper.

References

Becker, P. (1983). Effects of insect herbivory and artificial defoliation on survival

of seedlings of Shorea, pp. 241-252 in Tropical Rain Forest: Ecology and Manage-

ment. Eds. A.C. Chadwick, S.L. Sutton and T.C. Whitmore. Blackwell Scientific

Publications, Oxford.

Burgess, P.F. (1975). Silviculture in the hill forests of the Malay Peninsula. Research

Pamphlet, Forest Research Institute, Kepong No. 66.

Curtis, C. (1894). A catalogue of the flowering plants and ferns found growing wild

in the island of Penang. J. Straits Br. Roy. As. Soc. 25, 67-169.

Desch, H.E. (1941). Dipterocarp timbers of the Malay Peninsula. Malay. Forest Rec.

No. 14.

Jacobs, M. (1988). The Tropical Rain Forest. Springer-Verlag, Berlin.

Ong Jin Eong and Dhanarajan, G. (1976). On a national park for Pulau Pinang.

Malay. Nat. J. 29, 277-281.

Raich, J.W. (1987). Canopy openings, seed germination, and tree regeneration in

Malaysian coastal hill dipterocarp forest. Thesis (Ph.D.), Duke University, North

Carolina.

Symington, C.F. (1943). Foresters’ manual of dipterocarps. Malay. Forest Rec. No. 16.

Turner, I.M. (1988). The response of tree species to canopy gaps in a tropical forest.

Thesis (D.Phil), University. of Oxford.

44 _ Gard. Bull. Sing. 42(1) (1989)

Whitmore, T.C. (1984). Tropical Rain Forests of the Far East (Second edition),

Claredon Press, Oxford.

. (1988). The influence of tree population dynamics on forest species composi-

tion, pp. 271-291 in Plant population ecology. Eds. A.J. Davy, M.J. Hutchings

and A.R. Watkinson. Blackwell Scientific Publications, Oxford.

Wong Yew Kwan (1967). Some indications of the total volume of wood per acre in

lowland dipterocarp forest. Research Pamphlet, Forest Research Institute, Kepong

No-..53:

. (1987). Ecology of the trees of Bukit Timah Nature Reserve. Gard. Bull.

Sing. 40, 45-76.

and Teo Teck Seng (1978). Girth class distribution of Seraya Shorea [Shorea

curtisii] as an indication of its regeneration and recruitment in Bukit Timah

Nature Reserve. Malay. Nat. J. 31, 149-156.

Wyatt-Smith, J. (1963). Manual of Malayan silviculture for inland forests. Malay.

Forest Rec. No. 23.

. (1966). Ecological studies of Malayan Forests. Research Pamphlet, Forest

Research Institute, Kepong No. 52.

THE GARDENS’ BULLETIN

SINGAPORE

Volume 42

Part 2

‘OLA e

pstes f

Rt}

Didymocarpus (Gesneriaceae) on Gunung Tahan, Malaysia

RUTH KIEW

Department of Biology, Universiti Pertanian Malaysia,

43400 UPM Serdang, Selangor, Malaysia

EFFECTIVE PUBLICATION DATE: 15 MAR 1990

Abstract

A checklist for the 20 Didymocarpus species on Gunung Tahan is given. Three new species, D. codo-

nion, D. polyanthoides and D. yongii are described. The species Ridley called D. kompsoboea is renamed

D. lithophilus. D. filicifolius is reduced to synonomy with D. salicinus and reasons for not recognising

var. montanus of D. flavobrunneus are given.

Introduction

Botanical exploration of Gunung Tahan, the highest mountain in Peninsular

Malaysia reaching 2,188 m a.s.l., has been confined to the Tahan valley, the lower

reaches of the Teku Gorge, the ridge between the Tahan and Teku rivers and the

padang vegetation on the summit (Fig. 1). The first botanical collections were made

along the Tahan valley in 1891 by Ridley (who failed to ascend G. Tahan because

of a shortage of supplies and the prevalence of malaria among the porters) and from

G. Tahan by Robinson and Wray in 1905 and then by Ridley in 1911. These expedi-

tions yielded a large number of botanical novelties, which were described by Ridley

(1893, 1908, 1915).

Since the turn of the century most botanists working in Malaysia have visited G.

Tahan following the same route pioneered by Skeat and by Robinson and Wray. Pre-

war expeditions were made by boat up the Sungai Tahan to Kuala Teku. Because

of the expense of hiring boats and boatmen, post-war expeditions are made overland

following the east bank of the Sg. Tahan, which in fact is just as quick as the river

journey, which is hampered by rapids and at some times of the year by shallow water.

The difficulty in mounting an expedition to G. Tahan has always been the porter-

ing of supplies. Carrying a full load, it takes about three days to reach Kuala Teku

and another two to reach the summit. Most visits have of necessity been short and

there has been little collecting away from the beaten track.

Burkill (1927) noted how thorough Ridley’s collecting trips had been. Ridley

recorded 17 species of Didymocarpus (including those species he included in Paraboea

sect. Campanulati) from G. Tahan and the Tahan valley, of which 15 were new. Of

these, eight are endemic to the G. Tahan area, viz. D. ericiflorus, D. flavobrunneus,

D. leucocodon, D. lilacinus, D. grandiflorus (now called D. ridleyanus), D. robinsonii,

D. rubiginosus and D. salicinus. No further species were described from the area

until our expedition in 1987, when another three species were discovered of which

D. polyanthoides and D. yongii are endemic to the Tahan valley. The third, D.

codonion, has also been collected from Trengganu.

In total 20 species are now recorded from G. Tahan and its foothills representing

about a quarter of the 85 odd species of Didymocarpus in Peninsular Malaysia. Only

three of these species are widespread throughout Peninsular Malaysia (D. crinitus,

48 Gard. Bull. Sing. 42(2) (1989)

Gunung Tohan

Fig. 1. | Map of botanical collecting sites on Gunung Tahan, Peninsular Malaysia.

which is also known from Borneo and Sumatra; D. platypus, which also grows

in Sumatra; and D. quinquevulnerus, a peninsula endemic). D. fasciatus and D.

lithophilus are not so common but have a wide distribution in the peninsula excluding

Johore. All five species belong to sect. Heteroboea and it is very rare to find a

lowland forest without at least one species of this section.

Ten species are endemic to the G. Tahan area. The other five have an easterly

distribution and have not been collected from the Main Range. D. atrosanguineus,

D. codonion and D. tahanicus have been collected from central Pahang and Treng-

ganu, while D. heterophyllus and D. pyroliflorus have also been collected further

south from Johore. The majority of the endemic species are allied to species with

an eastern distribution: D. flavobrunneus is closely related to D. falcatus Kiew from

Johore; D. lilacinus to D. corneri Kiew from Trengganu and D. niveus (Kiew) Kiew

from Johore; D. salicinus to D. salicinoides Kiew from southern Kelantan and Treng-

ganu; and D. polyanthoides to D. atrosanguineus Ridley. Only D. robinsonii and

D. ridleyanus, both montane species, are related to species that grow on the Main

Range: D. robinsonii to D. hispidus Ridley and D. ridleyanus to D. flavescens Ridley.

(D. ericiflorus, D. leucocodon, D. rubiginosus and D. yongii do not show a close

relationship with any other Malayan species).

The aim of our expedition was to collect complete material (including material

preserved in spirit) and to make field notes on habitat, morphology and variation

within populations. We also spent a day each along the Sg. Tahan and Sg. Teku and

it was along the Sg. Teku that two new species were found.

It is unfortunate that the only expedition from the north, by Waterstradt in 1901,

which was the first successful ascent of G. Tahan, was a zoological one and did

not produce any herbarium specimens. Although orchids were collected they were

abandoned on the descent (Waterstradt 1902). Today there is no trail to G. Tahan

either from the north or from the west and these areas remain ferra incognita to

the botanist.

Didymocarpus on Gunung Tahan 49

The majority of Didymocarpus species in the G. Tahan area are found in the

shaded understorey of lowland forest. Species such as D. platypus and D. quin-

quevulnerus form dense patches that dominate the ground layer. On the banks of

the Sg. Tahan several species are common on lightly shaded rocky or steep earth

banks. D. salicinus is a rheophyte and grows down to the water’s edge, D. hetero-

phyllus and D. pyroliflorus grow higher up and are presumably less regularly sub-

jected to flooding. D. yongii finds a foothold on the sloping edge of lightly shaded

cliffs, while D. lithophilus clings to crevices in shaded rock faces along the tributaries.

In hill forest, D. flavobrunneus grows on the vertical faces of damp cliffs and

large boulders, while D. tahanicus is common on steep slopes below the ridge at

Wray’s Camp. Along the ridges leading up to G. Tahan, D. rubiginosus grows in

crevices on the vertical sides of dry scarps — an unusual habitat for Didymocarpus

species, the majority of which cannot withstand exposed conditions. In upper mon-

tane forest D. leucocodon and D. robinsonii are common and D. salicinus persists

along the Sg. Teku up to the edge of the padang giving it the widest altitudinal

amplitude (70-1,700 m a.s.1l.) of any species in the peninsula. Didymocarpus species

are absent from the padang, a harsh environment for plant life where low stunted

vegetation is scattered over exposed rock slopes, which are infilled with bleached

silver sand.

The literature cited below gives that of the type description and those that involve

nomenclatural changes, as well as Ridley’s flora (1923), which remains the most

recent complete account for Peninsular Malaysia. Only specimens collected from the

National Park (Taman Negara) are cited.

Checklist of Didymocarpus species

. Didymocarpus atrosanguineus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 328; Fl. Mal. Pen. 2 (1923) 518.

Lectotype: Ridley s. n. Tahan River (SING).

Distribution: Kelantan — Kuala Aring; Pahang — Tahan and Kenyam valleys;

Trengganu — Batu Biwa, Sekayu F.R., Ulu Brang.

Habitat: In deeply shaded lowland forest, locally common.

Specimens examined: Tahan River Ridley s. n. (SING); Kuala Kenyam | Oct 1982

Kiew B.H. RK121] (UPM), 6 Oct 1984 R. Kiew RK1458 (UPM).

Notes: D. atrosanguineus is the only species in sect. Heteroboea with red flowers.

The corolla is pale cream outside with blood-red lobes and inside the throat is

yellow with two golden-yellow nectar guides aligned inwards from between the lower

lobes. In addition, it differs from other species in this section in its particularly long

capsule, which is 9-11 cm long.

There are two forms both of which are found within Taman Negara and which

have identical flowers. One form has leaves tightly clustered at the top of the stem,

the leaves are rather broad (9-10 cm wide), the margin is less jaggedly toothed and

the indumentum is finely velvety. The other form has leaves spaced up to 2 cm

apart, the leaves are narrower (6-8 cm), the margin is more jagged particularly

towards the leaf base and the indumentum is hispid rather than velvety.

. Didymocarpus codonion Kiew sp. nov.

Didymocarpus floribundus (M.R. Henderson) B.L. Burtt et D. heterophyllus Ridley affinis sed petiolo

longioribus et lamina basi anguste cuneatis differt.

50 Gard. Bull. Sing. 42(2) (1989)

Type: Kiew Bong Heang RK1204 (holo UPM: iso SING).

Stem woody c. 6 mm thick, to 17 cm tall, flowering at c. 5 cm tall. Leaves

crowded at the top of the stem, spirally arranged, leaf bases persistent. Lamina

oblanceolate, 11 by 4 cm to 19 by 8.5 cm, apex acute, base narrowly cuneate,

glabrous above, in life upstanding and bullate, dark green above, young leaves

pinkish or whitish at base, in dried state rough and minutely pustulate beneath,

margin crenate. Midrib and veins plane above, prominent beneath, in life red, secon-

dary veins 9-15 pairs, tertiary veins prominent beneath. Indumentum of stem apex,

petioles and lower surface of midrib and veins densely matted with long uniseriate

hairs, sparse on peduncle. Petiole 2.5-7 cm long, 1.5 mm thick.

Inflorescence a cymose panicle with 3-4 order branching and up to 14 flowers,

peduncle slender (3-) 7-12 cm, branches 2.5-3 cm long. Pedicels slender 2-10 mm

long. Bract pairs ligulate, 3-4 mm long. Calyx divided to base, lobes narrowly acute,

2-3 mm long, + minutely hairy with few long hairs. Corolla narrowly campanulate,

pale lilac or pink, tube 2.5-4 by 2 mm, lobes similar, oblong with slightly acute apex,

1.5 by 1 to 2 by 1.5 mm, reflexed, inner surface and margin densely papillose with

minute glandular hairs. Stamens 2, filament 1 mm long, anthers enclosed within

corolla tube, 2 by 1.5 mm, connivent. Ovary 2 by 0.8 mm long, style slender 3-4 mm

long projecting beyond corolla, ovary and style minutely pubescent, stigma discoid

0.5 mm across, disc none. Capsule narrowly linear, straight, 14-18 mm long.

Distribution: Pahang — Tahan and Kenyam valleys; Trengganu — Batu Biwa.

Habitat: Lowland forest (below 200 m), locally common in deep shade on earth

banks.

Specimens examined: Kuala Kenyam 30 Sept 1982 Kiew B.H. RK1204 (UPM, SING);

2 Oct 1984 R. Kiew RK1399 (K); 6 Oct 1984 R. Kiew RK1457 (L); Batu Biwa 22 Oct

1986 R. Kiew RK230] (UPM); Tahan Valley 21 March 1987 R. Kiew RK2419 (UPM).

Notes: This species is closely related to D. heterophyllus (see below) and D. floribun-

dus, the latter has been collected only from the Bukit Kajang area in Kemaman,

Trengganu. All three are similar in their leaves, which are oblanceolate, acute or

rounded at the apex and are glabrous above, in the crenate margin and the pink or

white colour of the unexpanded leaves, in their cymose inflorescences, small cam-

panulate flowers 3-9 mm long in which the upper two lobes are reflexed and in the

short fruits, which are 1-3 cm long. D. heterophyllus differs from the other two

species in its few-flowered inflorescences and narrower leaf (Table 1). D. codonion

and D. floribundus have cymose panicles more than twice branched and are generally

more robust plants with larger leaves. Compared with D. floribundus, D. codonion

has much smaller bracts and calyx lobes.

In addition, D. codonion differs from both D. floribundus and D. heterophyllus

(Table 1) in its leaves, which in life are bullate above, in its leaf base, which is

narrowly cuneate rather than rounded or cordate, and in its longer petioles and in

its flowers which are much smaller and narrower (hence the epithet, which means

little bell in Greek). Although both D. codonion and D. heterophyllus grow in the

foothills of G. Tahan, they do not grow together as D. codonion is a plant of the

shaded undergrowth while D. heterophyllus grows on lightly shaded river banks.

. Didymocarpus crinitus Jack

Mal. Misc. I (2) (1820) 1; Ridley Fl. Mal. Pen. 2 (1923) 519, fig. 124.

Distribution: Johore — G. Pulai; Kedah — G. Jerai (Kedah Peak); Kelantan —

Chaning, Kota Baru, K. Lebir; Negri Sembilan — Bk. Sulu, Sg. Ujung; Pahang —

Didymocarpus on Gunung Tahan Si

Table 1

Diagnostic characters to distinguish

Didymocarpus codonion, D. floribundus and D. heterophyllus

D. codonion D. floribundus D. heterophyllus

No. flowers/inflorescence 10-14 8-18* 2-4

Stem height (cm) 5-17 2-13 (O-) 3 (-7)

Lamina width (cm) 4-8.5 3.5-6 (2-) 2.5 (-4)

Lamina length (cm) 11-19 10-18 (S-) 8 (-13.5)

Lamina base narrowly rounded, rounded

cuneate + cordate

Petiole length** (cm) 2.5-7 0.5-2 (0.5—-) 1.5 (-3)

Peduncle length (cm) (3-) 7-12 4.5-9 (2.5-) 5.5 (-8)

Calyx length (mm) 1-1.5 3-5 (1-) 2

Corolla length (mm) 2.5-4 raw ee. | 3-4

Corolla width (mm) Z 4-5 3-4

Corolla colour pink or pale-deep purple-violet

pale lilac purple

* Henderson (1933) from cultivated plants.

** young leaves are sessile, measurement is from the oldest leaf.

Cameron Highlands, Sg. Tahan; Penang Hill; Perak — Thaiping Hills; Selangor —

Kanching, G. Bidai; Trengganu — Bk. Kajang. Singapore, Borneo and Sumatra.

Habitat: Although it is frequently a plant of hill forest, in Taman Negara it has been

collected only in the lowlands on shaded rocky stream banks at c. 70 m azss.l..

Specimens examined: Sg. Tahan 27 July 1936 Kiah SFN31910 (K), 29 March 1987

R. Kiew RK2482 (UPM).

Notes: This is a distinctive species in its leaves, which are narrowly lanceolate and

are deep purple beneath.

. Didymocarpus ericiflorus Ridley

J. Fed. Mal. States Mus. 6 (1915) 166; Kiew Blumea (in press).

Synonym: Codonoboea ericiflora (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 533.

Type: Ridley 16283 G. Tahan (Wray’s Camp) (holo K; iso SING).

Distribution: Endemic to G. Tahan.

Specimens examined: Wray’s Camp July 1911 Ridley 16283 (K, SING), 29 August

1928 Holttum SFN20709 (SING).

Notes: This is a very rare species, which has only been collected twice. In spite of

a search I was unable to refind it. Ridley (1923) included this species in his new

genus, Codonoboea, on account of its epiphyllous flowers. This genus and its species

are now returned to Didymocarpus (Kiew, in press).

52 Gard. Bull. Sing. 42(2) (1989)

Reference:

Kiew, R. Blumea (in press). Reassessment of the generic status of Codonoboea (Gesneriaceae) and its

species.

. Didymocarpus fasciatus Ridley

J. Roy. Asiatic Soc. Str. Br. 43 (1905) 50; Fl. Mal. Pen. 2 (1923) 520.

Lectotype: 1891 Ridley 2169 Tahan River (K).

Distribution: Kedah — Kedah Peak; Negri Sembilan — Bk. Tangga; Pahang —

G. Tapis, Sg. Tahan.

Habitat: Rare and local, it grows on rocky banks well above the flood zone of

Sg. Tahan.

Specimens examined: Sg. Tahan 10 June 1922 Md. Haniff & Md. Nur 8105 (K),

Kiah SFN31910 (K).

Notes: This species most resembles D. crinitus in its narrow lanceolate leaves. It

differs in its indumentum of persistent hair bases, which give the dried leaf the rough

appearance of lizard skin; in its leaf margin, which is more finely toothed; and in

leaf colour, which is malachite green with a broad central pale band. Not a common

species, it nevertheless has a widespread distribution. Apparently rarely in flower,

this may account for it being undercollected. Its leaf coloration renders it a striking

plant even when sterile. Both D. fasciatus and D. crinitus have been collected from

Sg. Tahan and Kedah Peak; otherwise their distribution does not overlap.

Ridley (1923) gives D. crinitus var. tuberculatus C.B. Clarke as a synonym for

D. fasciatus, but this name was never published. Clarke did annotate a Sarawak

specimen (Beccari PB 1557 from G. Matang, K, FI) as var. tuberculatus, but Clarke

(1883) listed this specimen as D. crinitus var. exasperatus, which was characterised

by ‘pilis e tuberculo ortis exasperatis’. Ridley (1905) noted his species ‘is allied to

a Bornean plant occurring on Mt. Matang [i.e. Beccari’s specimen], which it much

resembles’. Beccari’s specimen is indeed similar in its rough indumentum and leaf

coloration to the Malayan D. fasciatus. However, until a wider range of specimens

of both these taxa is available, I hesitate to consider them conspecific.

. Didymocarpus flavobrunneus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 329; Fl. Mal. Pen. 2 (1923) 515.

Typus: Ridley 2163 Sg. Tahan (holo K, iso SING).

Synonym: D. flavobrunneus var montanus Ridley

J. Fed. Mal. States Mus. 6 (1915) 167; Fl. Mal. Pen. 2 (1923) 515.

Type: Ridley 16384 Wray’s Camp (holo K, iso SING).

Distribution: Endemic to Gunung Tahan area.

Habitat: It is confined to shaded low cliffs or large boulders from 300 m on Bk.

Teresik to 1,200 m at Wray’s Camp and beyond.

Specimens examined: Tahan Woods Nov 1891 Ridley 2163 (K); Wray’s Camp 2 June

1905 Wray & Robinson 5367 (K), July 1911 Ridley 16384 (K), 10 June 1922 Md.

Haniff & Md. Nur 8103 (K), 21 July 1936 Kiah SFN31754 (K), 23 March 1987 R.

Kiew RK2428 (UPM), 28 March 1987 RK2476 (UPM).

Notes: This species is most closely related to D. falcatus Kiew from G. Janing,

Johore. Both species grow on dry, shaded rock faces and produce a woody stem

Didymocarpus on Gunung Tahan 53

with a tuft of soft, hairy leaves at the top. Their cymose inflorescences have long

peduncles supporting a condensed cluster of flowers. They differ, amongst other

characters (Kiew, 1987), in flower colour, which is white with yellow lines in the throat

in D. falcatus and brownish-red with yellow lobes and a yellow throat with brownish-

red lines in D. flavobrunneus.

According to Ridley, var. montanus differs from the typical lowland variety in

its greater height, its more softly woolly leaves, and the colour of the corolla (var.

montanus has broad red-brown bands in the throat compared with a few streaks in

the typical variety). Examination of plants in the field and the wider range of her-

barium specimens now available shows that var. montanus cannot be maintained

because plants of both varieties grow to 30 cm tall, the majority of the montane

plants have identical indumentum with the lowland plants i.e. they are sparsely hairy

as opposed to velvety, and some lowland populations (e.g. at Bk. Teresik, 300 m)

also have broad maroon lines in the throat. For these reasons, var. montanus is not

recognised as a distinct taxon.

Reference:

Kiew, R. 1987. The herbaceous flora of Ulu Endau, Johore-Pahang, Malaysia, including taxonomic

notes and descriptions of new species. Mal. Nat. J. 41: 201-234.

. Didymocarpus heterophyllus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 329; Fl. Mal. Pen. 2 (1923) 522.

Type: Ridley 2170 Tahan valley (SING).

Distribution: Johore, Bk. Pengantin (Labis), G. Janing, G. Lesong, Sg. Kinchin

(Ulu Endau); Pahang, Bk. Cheras (Kuantan), Sg. Tahan.

Habitat: Lightly shaded stream banks in the lowlands, growing on rocks or vertical

earth banks above the normal water level.

Specimens examined: Tahan valley Ridley 2170 (SING), 22 March 1987 R. Kiew

RK2423 (UPM).

Notes: This is a common plant all along the steep banks of the Sg. Tahan, where

it grows with D. pyroliflorus. Although it grows above the normal water level, it

will nevertheless be subject to periodic flood waters.

. Didymocarpus leucocodon (Ridley) Kiew comb. nov.

Synonym: Paraboea leucocodon Ridley

J. Fed. Mal. States Mus. 6 (1915) 167.

Codonoboea leucocodon (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 533.

Type: Ridley 1604] (holo K; iso SING).

Common name: The Bell of Tahan.

Distribution: Endemic to Gunung Tahan.

Habitat: This species grows along damp mossy ridges near Tangga Dua Belas and

in gullies below the padang at about 1,500-1,700 m a.s.1., often with Pentaphragma

aurantiacum Stapf (Pentaphragmataceae). The only other species of Didymocarpus

that grows at this altitude is D. robinsonii (see below).

Specimens examined: G. Tahan July 1911 Ridley 16041 (K, SING), 30 Aug 1928 Md.

Nur SFN20951 (SING), 25 March 1987 R. Kiew RK2448 (UPM).

54 Gard. Bull. Sing. 42(2) (1989)

Notes: Didymocarpus leucocodon is a strikingly beautiful species with pure white,

large, bell-like flowers (codon = bell, Greek). Corner suggested its common name

be The Bell of Tahan (Corner 1937, unpublished field notes). In March when I

visited G. Tahan all plants were in bud. Flowering specimens have been collected

in July and at the end of August and September, which suggests it might be a

seasonally flowering species.

Ridley (1923) included this species in Codonoboea although it does not have

epiphyllous flowers. For this reason, it is not included in sect. Codonoboea within

Didymocarpus (Kiew, in press). It is not closely allied to any other Malayan species

as it is unusual in possessing a corky bark, stiff fleshy leaves (which Corner described

as like thin cardboard) with veins plane above and below, and a corolla with acute

lobes and a matted indumentum outside.

Reference:

Kiew, R. Blumea (in press). Reassessment of the generic status of Codonoboea (Gesneriaceae) and its

species.

. Didymocarpus lilacinus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 330; J. Roy. Asiatic Soc. Str. Br. 43 (1905) 56; Kiew Blumea

(in press).

Synonym: Codonoboea lilacina (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 534.

Type: Tahan Valley Ridley 2165 (holo K; iso SING).

Distribution: Endemic to the Tahan Valley.

Habitat: It grows in lowland forest at c. 100 m a.s.l. on steep earth banks devoid

of other vegetation on the slopes above Sg. Tahan.

Specimens examined: Tahan valley 1893 Native coll. (K), July 1911 Ridley 2165

(K, SING), Sept 1929 Md. Nur s. n. (SING), 7 June 1931 Henderson SFN24847

(K, SING); Mohd Shah MS2717 (SING), 5 Sept 1982 R. Kiew RK1216, 20 March 1987

RK2407 (UPM).

Notes: It is a rare and local species, known only from a few populations where it

grows in dense clumps. Its leaves are unequal, of a pair the leaf produced away from

the earth bank on which it grows is longer than the one produced towards the bank.

The leaves are very soft and in dry weather, e.g. in March 1987, its leaves were

wilted, although other undergrowth herbs, such as Sonerila species and Didymocarpus

platypus showed no signs of water stress.

. Didymocarpus lithophilus Kiew nom. nov.

Synonym: D kompsoboea Ridley Trans. Linn. Soc. 2nd. Ser. 3 (1893) 328; Fl. Mal. Pen. 2 (1923) 518

— non D. kompsoboea C.B. Clarke in D.C. Mon. Phan. 5 (1883) 92, t. 10.

Type: Ridley 2152 Kuala Tanah (holo K; iso SING).

Distribution: Endemic to Peninsular Malaysia: Kedah — G. Bintang; Kelantan —

G. Setong; Pahang — Tahan Valley; Perak — G. Inas, Temangok; Selangor —

Bk. Hitam; Trengganu — Bk. Kajang.

Habitat: It grows on shaded vertical rock faces (usually granite) on stream banks

or cliffs, usually in the lowlands’ but at Wray’s Camp it grows at c. 1,000 m a.s.l..

Specimens examined: Sg. Tahan Aug 1891 Ridley 2152 (K), 21 June 1922 Md. Haniff

& Md. Nur 8037 (K, SING), Aug 1928 Holttum 20953 (SING), Seimund 873 (SING),

Didymocarpus on Gunung Tahan 55

March 1987 R. Kiew RK2425 (SING), RK2483 (UPM); Wray’s Camp 8 July 1911

Ridley s. n. (K).

Notes: This species has a woody stem 6-21 cm long, its leaves are 17-33 cm long

and 4-10.5 cm wide, they are plane above with 33-50 pairs of secondary veins, the

surface is not bullate and its capsule is c. 3 cm long. Ridley (1893) identified it as

D. kompsoboea, which was described from Borneo (type Beccari PB 3440, FI). Apart

from possessing the same rectangular venation pattern (Fig. 2), D. kompsoboea

differs in its rosette habit and its smaller, conspicuously bullate leaves (14 by 6-7 cm),

which are rugose beneath and its longer fruits (S.5-6 cm long). The rosette habit is

rare among Malayan species of sect. Heteroboea to which D. lithophilus belongs and

is seen only in D. rugosus Ridley. All other Malayan species in this section are robust

plants with a woody stem bearing a mop of large leaves at the top. Since it is not

conspecific with D. kompsoboea, Ridley’s species is renamed D. lithophilus.

Among Malayan species, D. lithophilus most resembles D. platypus in its thin

leaves, which are not densely pubescent on the upper surface. It differs from D.

platypus in the tertiary venation, which forms a pattern of regular rectangles, and

the areoles, which are flat and are not surmounted by a raised hair base. This

arrangement gives the dried leaf the appearance of crocodile skin. In contrast, in

D. platypus (Fig. 2) the tertiary veins form a polygonal pattern, and in addition the

areole is mammilate with a large hair base arising from its centre. D. lithophilus is

also different in being shorter, in having a thinner and more wiry stem, in its lower

leaves, which are spaced on the stem, and in bearing fewer leaves in the crown

(about 7 compared with about 10 in D. platypus), and in life the corolla appears

thinner, more delicate and almost transparent (the corolla of D. platypus is thicker

and more fleshy). The corolla is white suffused with pale lilac, the lobes are more

deeply tinted. Flower colour is variable in D. platypus depending on locality. Usually

it is white with two lemon-yellow nectar guides in the throat, but in some popula-

Fig. 2. Tertiary venation patterns in a. Didymocarpus lithophilus (RK2425), b. D. kompsoboea

(Beccari 3440) and c. D. platypus (Henderson 11422).

11.

56 Gard. Bull. Sing. 42(2) (1989)

tions these lines are lacking, and in a few populations the corolla is tinged purple

or cream or yellow. The fruit of D. lithophilus is shorter, c. 2.5 cm long, compared

with that of D. platypus, which is c. 5 cm long.

D. lithophilus is named for its habitat. It grows on vertical rock faces, a unique

habitat for Malayan species of sect. Heteroboea. Its wiry root system splays out over

the rock surface and becomes woody, while the fine roots penetrate cracks and

crevices. Its less robust habit, shorter stem and fewer leaves may be an adaptive

feature in reducing the weight of the top of the plant. D. platypus and the other

species in sect. Heteroboea are rooted in the soil.

Didymocarpus platypus C.B. Clarke

in D.C. Mon. Phan. 5 (1883) 94; Ridley Fl. Mal. Pen. 2 (1923) 517.

Distribution: Throughout Peninsular Malaysia. Sumatra.

Habitat: It is the commonest species of Didymocarpus in Peninsular Malaysia and

is found in almost every undisturbed shady lowland and hill forest up to about

1,000 m a.s.l., where it grows on the forest floor, frequently gregariously.

. Didymocarpus polyanthoides Kiew sp. nov.

A Didymocarpus atrosanguineus tubo corollae rubineo-purpureo et lobis carmesino-purpureo et fructis

brevioribus differt.

Type: Kiew RK2480 (UPM).

Stem woody 30-85 cm tall and | cm thick. Leaves sessile, crowded at top of stem.

Lamina oblanceolate, 25-30 by 6-9 cm, apex acute to attenuated, narrowed and

winged to base, thin, drying chartaceous, margin doubly serrate. Midrib plane above,

prominent beneath. Secondary veins c. 28 pairs, plane above, prominent beneath,

tertiary veins obscure above, prominent beneath, areoles forming a polygonal pattern

beneath. Indumentum softly hispid, dense above and below, hairs uniseriate, 4-6

celled, + appressed to leaf surface, hair base raised, hairs tufted on margin, hairs

on lower surface of midrib longer and thicker.

Flowers solitary, several per leaf axil. Pedicel 4-6 cm slender, hirsute with uni-

seriate, multicellular hairs c. 1 mm long, and shorter glandular hairs. Bract pairs

ligulate, 4 mm long, finely hairy. Calyx divided to base, lobes linear, 4-5 by 1-2 mm,

densely hirsute. Corolla infundibular, 3.5-5 cm long, 3.5 cm wide at base, enlarging

to 9 mm at the mouth, lobes c. 1 by 1 cm, broadly oblong, apex rounded, finely

and sparsely pubescent outside, tube ruby-purple outside, lobes crimson-purple

with 2 golden yellow nectar guides in throat, inner surface of upper two lobes with

glistening hairs. Stamens 2, filament slender, 11 by 1.5 mm, anthers oblong, 3 by

1 mm, connivent. Ovary narrowly cylindric, 2.5 cm long, style slender 1.5 cm long,

ovary and style densely pubescent, stigma white, discoid, slightly bilobed, 2 mm

across, nectary cylindric, shallowly lobed, 1.5 mm long. Capsule 7 cm, finely

pubescence when young, hairs glandular with thin multicellular stalk.

Distribution: Pahang — endemic to Sg. Teku in the foothills of G. Tahan.

Habitat: Growing in lowlands at c. 100 m a.s.l. on steep shaded slopes above the

river.

Specimens examined: Sg. Teku 21 June 1922 Md. Haniff & Md. Nur 8042 (SING);

29 March 1987 RK2480 (UPM).

Notes: Along the Tahan valley, grow seven species of sect. Heteroboea. While D.

lithophilus is restricted to rock faces, the others are plants of the forest floor. D.

Didymocarpus on Gunung Tahan 57

platypus is the most common and widespread; the others are either widespread

but less common, e.g. D. atrosanguineus, or are locally common, such as D. quin-

quevulnerus, which grows in dense patches, or they are rather rare, such as D.

crinitus, D. fasciatus and D. polyanthoides. In some cases, more than one species

grow together, as for example populations of D. quinquevulnerus which often grow

intermixed with plants of D. platypus. The few plants of this new species were also

found growing with plants of D. quinquevulnerus.

This new species is distinguished by its flower colour, which recalls that of the

purple garden polyanthus with its yellow-orange eye. Among Malayan Didymocarpi,

red corollas are seen only in D. atrosanguineus, which however has blood-red rather

than ruby-purple corolla lobes as does D. polyanthoides. Among species in sect.

Heteroboea only D. quinquevulnerus has deep purple lobes, which are an imperial

purple untainted by red; the other species have white corollas which may be suffused

with pale purple. It is not known what pollinates any of these species but a circular

hole at the base of the corolla of one flower of this new species indicates that nectar

thieves were at work.

Apart from flower colour, D. polyanthoides can be distinguished by its shorter

fruits (7 cm long) from D. atrosanguineus, which has exceptionally long fruits

(8.5-10 cm long). It can be told apart from D. quinquevulnerus not only by flower

colour but also by indumentum, as that of D. quinquevulnerus is longer and more

silky. It is also not as floriferous as D. quinquevulnerus.

It is obviously a very rare plant. I was only able to find a single population

consisting of a few plants along the Sg. Teku above Kuala Teku, which is also the

area from where Md. Haniff and Md. Nur collected it.

. Didymocarpus pyroliflorus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 330

Synonym: Paraboea pyroliflora (Ridley) Ridley

J. Roy. Asiatic Soc. Str. Br. 43 (1905) 67; Fl. Mal. Pen. 2 (1923) 529.

Type: Ridley 2164 (holo K, iso SING).

Distribution: Johore — G. Belumut, G. Panti, Kota Tinggi, Sg. Pelepah Kiri; Pahang

— Tahan Valley.

Habitat: It grows on rocks on lightly shaded river banks above the normal water

level but within the flood zone along the Sg. Tahan.

Specimens examined: Tahan River 1891 Ridley 2164 (K, SING); 5 August 1905 Wray

& Robinson 5545 (K), 20 Dec 1920 Seimund s. n. (K), 10 June 1922 Md. Haniff &

Md. Nur 8126 (K), March 1987 R. Kiew RK2422 (UPM), RK2470 (UPM).

Notes: Its flower is unique among Malayan species of Didymocarpus in its shape (its

tube is very narrow at the base, then opens broadly, the lower lobes are scarcely

longer than the upper and the lobes are not recurved), and in its very long style,

which is up to 1.7 times as long as the corolla. (The large anthers are included within

the corolla tube.) D. heterophyllus has a style that projects a few millimetres beyond

the upper corolla lobes, which are reflexed, but it is shorter than the lower lobes,

which project beyond the upper.

In 1905 Ridley transferred this species from Didymocarpus to Paraboea sect.

Campanulati, which he erected to accommodate species of Didymocarpus with short-

tubed campanulate flowers, short stamens and a style longer than the stamens. Burtt

(1971) abolished this section as he considered it contained short-flowered species of

14.

58 Gard. Bull. Sing. 42(2) (1989)

Didymocarpus. Apart from D. cordatus and D. tahanicus, he placed the remaining

species in section Salicini in Didymocarpus. However, sect. Salicini is best used in

a narrow sense to include only those species with willow-shaped leaves. Neither does

D. pyroliflorus fit comfortably into any other of Ridley’s sections, for while in habit

and foliage it resembles some species in sect. Didymanthus, species in this section

have tubular flowers with long thin filaments and the style is included within the

corolla tube. Didymocarpus pyroliflorus is therefore an isolated species within the

genus and until the sections are revised (almost all of which at present include

anomalous species), it is best not to assign this species to a section or to erect a new

one to accommodate it.

Didymocarpus quinquevulnerus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 328; Fl. Mal. Pen. 2 (1923) 518.

Type: Ridley 2153 August 1891 Tahan Woods (holo K, iso SING).

Distribution: Johore — Ulu Endau; Kelantan — Kg. La; Malacca; Pahang —

Fraser’s Hill, Semangok Pass, Tahan valley; Selangor — Klang Gates, Batu Tiga,

Kanching F.R..

Habitat: Shaded slopes in lowland forest up to 1,000 m a.s.l..

Specimens examined: Tahan valley Aug 1891 Ridley 2153 (K, SING); Aug 1928

Holttum 20550 (CGE, SING), 20955 (SING); July 1929 Henderson 21877 (SING),

22012 (SING); 22 March 1987 R. Kiew RK2426 (UPM); Kuala Teku July 1911 Ridley

16219 (K); Dec 1920 Seimund 508 (SING), 870 (SING); June 1922 Md. Haniff & Md.

Nur 8042 (K).

Notes: Apparently sharing the same habitat as D. platypus, with which it frequently

grows, it is nevertheless not a common species and its distribution is very local.

Where it occurs, it grows in dense patches. It is easily the most beautiful species in

sect. Heteroboea not only by virtue of its deep purple corolla lobes but also by its

floriferous habit (up to 40 flowers may be open at any one time). Ridley (1912)

described it as a beautiful plant ‘worth going far to see’. In 1893, he noted that it

is well worthy of cultivation. Unfortunately, these soft-leaved forest species do not

thrive away from the cool, humid conditions of the forest floor.

Colour of the corolla lobes is usually deep imperial purple but a few populations

or a few plants within a population may have paler lilac lobes (they are still, how-

ever, a deeper purple than those of D. lithophilus for example). One specimen from

Fraser’s Hill (Burkill & Holttum 8611, SING) is a white form collected within a

population of purple-lobed plants.

The original description of the species described the corolla lobes as ‘lovely dark

violet-purple’ and as ‘crimson edged white’ and among the specimens cited by Ridley

(1893), the one from Kota Gelanggi (Ridley 2154, K, SING) is described as ‘the pink

variety’. This specimen resembles the others of this species in all other respects,

although of course flower colour is not preserved. Unfortunately description of

colour is sometimes not consistent especially for the purple spectrum which may

variously be described as pink, purple or blue depending on the collector. In this

case since Ridley collected both specimens there is likely to be some difference in

colour between them. Unfortunately the Kota Gelanggi population no longer exists

as the surrounding forest has been converted to oil palm estate. The status of the

pink variety therefore remains in doubt. D. quinquevulnerus also differs from other

species in sect. Heteroboea in its densely silky indumentum.

16.

Didymocarpus on Gunung Tahan 59

Didymocarpus ridleyanus B.L. Burtt

Notes Roy. Bot. Gard. Edinb. 23 (1960) 99.

Synonym: D. grandiflorus Ridley

J. Fed. Mal. States Mus. 6 (1915) 167; Fl. Mal. Pen. 2 (1923) 523.

Type: Ridley s. n. July 1911 (K).

Distribution: Endemic to Gunung Tahan (Wray’s Camp).

Habitat: Collected from streams in hill forest at c. 1,100 m azss.l..

Specimens examined: Ridley s. n. July 1911 (K); 27 Aug 1928 Holttum SFN20587 (K).

Notes: This is a very rare species indeed, known from two collections from the

type site. It certainly does not belong to sect. Boeopsis where Ridley placed it, as

this section is defined as including plants with small flowers and a rosette habit,

whereas D. ridleyanus has pairs of leaves 2 cm apart and its flowers are, according

to Ridley, 5 cm long (although buds on Holttum SFN20587 are 2 cm long). Vege-

tatively it resembles some species in sect. Didymanthus and in particular D. flavescens

Ridley, which also has solitary flowers. However, its flower is unusual for species

of Didymocarpus. Ridley described them as violet purple, a colour more commonly

encountered in species of Didissandra or Chirita.

Didymocarpus robinsonii Ridley

J. Linn. Soc. 37 (1908) 318; Fl. Mal. Pen. 2 (1923) 513.

Type: 5 July 1905 Wray & Robinson 5470 (K)

Distribution: Endemic to G. Tahan at 1,700-2,000 m.

Habitat: This species is very common on lightly shaded slopes in upper montane

forest, e.g. in the gully at Tangga Dua Belas. It was through this gully that H.C.

Robinson found a route from the south to the top of G. Tahan in 1905. It is

therefore appropriate that this species should be named for him.

Specimens examined: G. Tahan 5 July 1905 Wray & Robinson 5470 (K); July 1911

Ridley 5470 (K, SING), 1/6040 (K); 11 June 1922 Md. Haniff & Md. Nur 8140 (XK),

30 August 1928 Holttum 20658 (K); 2 March 1973 F:S.P Ng FRI020963 (K, KEP);

24 March 1987 R. Kiew RK2432 (UPM).

Notes: Didymocarpus robinsonii is very similar to D. hispidus Ridley, which is a

common species in montane forest on the Main Range, in its erect, shrubby habit

with distant, opposite leaves and in its flowers, which are produced on long peduncled

cymes. It differs in flower colour — in D. hispidus the lower three lobes are some-

times streaked with purple, whereas in D. robinsonii there is a well-defined pattern

of purple lines alternating with the yellow nectar guides. D. robinsonii is also distinct

in its shorter fruit stalks (S-6 cm long) as compared with 8-12 cm in D. hispidus.

It differs from the other montane species with white flowers with violet stripes, D.

albinellus Ridley, which has larger leaves with c. 12 pairs of veins compared with

9 pairs in D. robinsonii.

At Tangga Dua Belas bumblebees were observed visiting its flowers.

. Didymocarpus rubiginosus (Ridley) B.L. Burtt

Notes Roy. Bot. Gard. Edinb. 31 (1971) 44

Synonym: Paraboea rubiginosa Ridley

J. Linn. Soc. Bot. 37 (1908) 319; Fl. Mal. Pen. 2 (1923) 530.

Type: Wray & Robinson 5390 (K).

Distribution: Endemic to G. Tahan between 1,300-2,000 m.

60 Gard. Bull. Sing. 42(2) (1989)

Habitat: It is most unusual for a species of Didymocarpus to grow rooted in fissures

on dry, exposed, vertical rock faces. It is found in this habitat on ridges and below

the edge of the padang.

Specimens examined: G. Tahan 3 June 1905 Wray & Robinson 5390 (K); July 1911

Ridley 16043 (K); 11 June 1922 Md. Haniff & Md. Nur 7865 (K, CGE); March 1987

R. Kiew RK2446 (UPM), 2467 (UPM).

Notes: This species is conspicuous in its dense velvety indumentum, which covers the

stem, petioles, upper leaf surface, and veins on the lower leaf surface. It is tempting

to suggest that this thick indumentum is an adaptation to reduce water loss. In life,

the upper leaf surface is dark green and the stem, petiole and veins on the lower

surface are densely covered by purple-brown hairs. In the dried state, the leaf is

reddish-brown. Plants growing in a more exposed position higher up the rock face

are more or less stemless and their compact rosette of leaves is appressed to the

rock surface. Those in a more shaded position may have stems up to 6 cm tall with

internodes between 2.5-10 mm long.

Ridley (1908) described this species under Paraboea sect. Campanulati as it has

campanulate as opposed to trumpet-shaped flowers. Burtt (1971) in transferring it

to Didymocarpus placed it in sect. Salicini. However, this section consists of a well-

defined group of narrow-leaved species, which usually live by rocky streams. Among

Ridley’s sections, D. rubiginosus with its rosulate habit and small flowers fits best

within sect. Boeopsis.

. Didymocarpus salicinus Ridley

Trans. Linn. Soc. 2nd Ser. 3 (1893) 329.

Synonym: Paraboea salicina (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 530.

Type: Ridley 2166 (K).

Didymocarpus filicifolius Ridley syn. nov.

J. Fed. Mal. States Mus. 6 (1915) 166

Paraboea filicifolia (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 530.

Type: Ridley 16059 (holo K; iso SING).

Distribution: Pahang, endemic to G. Tahan and Tahan valley.

Habitat: Rocky river banks growing at water level and on rock faces above the river,

from 70 to 1,700 m a:.s.l|..

Specimens examined: Tahan River 1891 Ridley 2166 (K, SING), 1893 Mat s. n.

(SING); 5 Aug 1905 Wray & Robinson 5544 (K); Dec 1920 Seimund s. n. (K); 21 July

1922 Md. Haniff & Md. Nur 8087 (K); 18 July 1936 Kiah SFN31705 (K); 20 March

1987 R. Kiew RK2409 (UPM); Kuala Teku July 1911 Ridley 16271 (K, SING); G.

Tahan July 1911 Ridley 16059 (K, SING).

Notes: Didymocarpus salicinus is one of the few rheophytes in the Gesneriaceae.

Along the Sg. Tahan, it grows on rocky banks within or immediately above the flood

zone. It is typical of rheophytes in having woody roots, which are firmly attached

to crevices in the rocks, wiry stems (the older plants branch and become bushy), and

willow-shaped leaves.

At Latah Berkoh on Sg. Tahan, plants just above the normal water level and

which are presumably subject to regular flooding have narrower leaves (c. 1 cm wide)

than plants that grow 2-4 m above the water level and which are presumably less

Didymocarpus on Gunung Tahan 61

regularly subject to flood waters, which have broader leaves up to 2.3 cm wide.

Plants growing in the small shaded tributaries also have broader leaves.

Didymocarpus filicifolius is based on one specimen collected from a stream bank

on G. Tahan at 1,700 m a.s.l.. Ridley (1905) distinguished it in his key from D.

salicinus by its white corolla and oblong parallel-sided leaves. However, flowers of

D. salicinus fade from pink to white and Ridley (1915) noted that the plant of D.

filicifolius was nearly out of flower suggesting its flowers were old. The leaves of

D. filicifolius fall within the range of size and shape of those plants of D. salicinus

that grow in more shaded or sheltered positions and have larger and more oblong

leaves. Although Ridley (1925) gives petiole length in D. salicinus as half an inch and

that of D. filicifolius as an eighth of an inch, examination of specimens shows that

the petiole length of D. filicifolius (S-12 mm) falls within the range of that of D.

salicinus (6-18 mm). The midrib of D. filicifolius is curiously wrinkled, but this

appears to be an abnormality of growth. Apart from its larger corolla, (7 mm

long compared with that of D. salicinus, which is about 4 mm long), D. filicifolius

falls within the range of variation of D. salicinus and with which it is considered

synonymous.

. Didymocarpus tahanicus B.L. Burtt

Notes Roy. Bot. Gard. Edinb. 31 (1971) 46.

Synonym: Didymocarpus grandifolius Ridley

J. Linn. Soc. Bot. 37 (1980) 318.

Paraboea grandifolia (Ridley) Ridley

Fl. Mal. Pen. 2 (1923) 531.

Type: Wray & Robinson 5369 (holo BM, iso SING).

Robust plant, stemless or with short stout woody stem c. 4 cm long and 1 cm thick.

Leaves crowded, young leaves purple beneath. Lamina oblanceolate, up to 28 cm

by 9 cm, apex acute, narrowing to base, margin crenate. Veins 17-19 pairs, arching

towards margin, midrib and veins plane above, prominent beneath, tertiary veins

conspicuous beneath. Indumentum of petiole, lower surface of midrib and veins,

peduncle, pedicel and outside of bracts and calyx with a dense layer of long multi-

cellular uniseriate hairs, appearing woolly on young petioles and peduncles. Petiole

5-6 cm long, fleshy.

Inflorescence a cymose panicle with up to 3rd order branching and up to 14

flowers orientated in the same direction. Peduncle 12-21 cm long, stout (up to 4 mm

thick in infructescence), branches 3-4 cm long. Bract pairs lanceolate, lowermost 10

by 1.5 mm, upper bracts 5 by 1 mm. Pedicel c. 5 mm long, pale violet. Flowers

pendant, 4-4.5 cm long, corolla lobes contorted in bud. Calyx divided to base, lobes

narrowly lanceolate, 7-9 mm by 2 mm, pale violet. Corolla tube trumpet-shaped,

25-30 mm long, flattened dorsiventrally, 3 mm wide at base expanding to 8 mm at

mouth, pale rosy-lilac outside with 2 yellow nectar guides in throat, sparsely covered

by stalked glandular hairs outside, inner surface glabrous, lobes broadly oblong, apex

rounded, pale lilac to almost white, upper two lobes recurved, 5 by 5 mm, each with

a large lemon-yellow spot which coalesces at their juncture, lower 3 lobes 8 by 6 mm

and projecting 5-10 mm beyond the upper lobes. Stamens 2, filaments slender 8 mm

long, attached halfway up corolla tube, anthers narrowly ovoid, 2 by | mm, conni-

vent. Ovary narrowly cylindric 12 by 1.2 mm, glabrous, style slender 12 mm tong,

stigma discoid 1 mm across. Disc cylindric c. 1 mm long. Capsule narrowly cylindric,

6.5 cm long and 2 mm thick.

Distribution: Pahang — Gunung Tahan; Trengganu — G. Padang.

20.

62 Gard. Bull. Sing. 42(2) (1989)

Habitat: On G. Tahan, it is common in a very local area on steep damp wooded

slopes in hill forest around Wray’s Camp at c. 1,100 m a.s.l..

Specimens examined: Wray & Robinson 5369 (SING), Holttum SFN20942 (SING),

March 1987 R. Kiew RK2427 (UPM), RK2472 (UPM).

Notes: In flower it is a very fine plant with up to 14 purple flowers each about 4 cm

long on a single inflorescence and often with several inflorescences per plant. As the

original description is incomplete because Ridley (1908) had only fruiting material

available, a complete description is provided here. Its flowers are unusual in several

respects. The bud is pointed because the unopened lobes are twisted. (In most species

the lobes are imbricate and the bud is broadly rounded.) The open flowers hang

down as though the pedicel is too weak to support the long corolla, and the flattened

corolla results in the mouth of the tube being narrowly oval rather than circular as

it is in most species.

Ridley (1908) first described it as a species of Didymocarpus with uncertain

affinities within the genus but in 1923 he transferred it to Paraboea, on the grounds

of habit as its flowers were then unknown. (Paraboea he characterised by its small,

campanulate flowers.) With flowering material available, Burtt (1971) returned this

species to Didymocarpus and, pointing out that ‘grandifolius’ had already been used

as an epithet for another species of Didymocarpus, renamed it ‘tahanicus’. (It has

also been collected from G. Padang in Trengganu, Moysey & Kiah SFN33924,

SING.) Burtt assigned it to sect. Heteroboea, which is, however, a section comprising

species with one-flowered inflorescences. It belongs with a group of montane species

with cymose panicles and foliaceous bracts that includes D. calcareus Ridley, D.

pubiflorus Ridley, D. castaneifolius Ridley and D. venustus Ridley, the latter two

Ridley (1923) placed in sect. Didymanthus.

Didymocarpus yongii Kiew sp. nov.

Species venatione alba conspicuis a congeneribus Malaya diversa.

Typus: R. Kiew RK2481 (holo UPM; iso L, K, SING).

Common name: The Kale-leaved Didymocarpus.

Stemless plant with woody rootstock and congested rosette of leaves + appressed

to ground. Lamina obovate, sometimes lanceolate, (9-) 14.5 (-15) by (4-) 5.5 (-6)

cm, apex acute or rounded, base narrowed, cordate, usually unequal, slightly rough

above. Indumentum on both surfaces of lamina and petiole of minute, erect, uni-

cellular hairs, dense in young leaves, leaf margin densely fringed, lower surface of

midrib and veins with dense covering of longer hairs. In life light or dark green above

with white veins, in some plants purple below. Margin finely serrulate. Midrib and

veins plane above, prominent below. Lateral veins (6-) 8 (-9) pairs, widely spaced,

subopposite, marginal vein 4-5 mm distant from margin, tertiary venation obscure

above, forming non-polygonal reticulation beneath. Petiole 1-3 cm long, slender.

Inflorescence a long-scaped cymose panicle. Peduncle slender 14-17 cm long (to

18 cm long in infructescence), branches 1.5-3.5 cm long, 2- to 3-times branched with

c. 12 flowers. Bract pairs ligulate, lower 2 mm long, upper 1 mm long. Pedicel

1-2.5 mm long, slender, flowers nodding. Indumentum of peduncle, pedicel, bracts

and calyx densely and minutely hispid. Calyx 1.5 mm long almost divided to base,

lobes narrowly acute, c. | mm long. Corolla campanulate, in bud minutely hispid

outside, 7 by 4 mm, pale lilac to almost white outside, lobes rosy-lilac, oval, 1.5 by

1 mm, apex rounded, +°equal in size. Stamens 2, filaments short c. 1.5 mm long,

thick; anthers yellow, oblong, 1.5 by 0.75 mm, connivent. Ovary cylindric 1 mm

long; style 2 mm long, curved upwards, minutely hispid; stigma minute, discoid, off-

Didymocarpus on Gunung Tahan 63

white in colour. Disc circular, 1 mm high. Capsule narrowly linear, 2-2.5 cm long,

glabrous.

Distribution: Pahang — endemic to Sg. Tahan and Sg. Teku at base of G. Tahan.

Habitat: Edge of overhanging rocks or cliffs where there is a covering of earth,

75 m a.s.l..

Specimens examined: Kuala Tahan 22 June 1922 Md. Haniff & Md. Nur 8301] (K);

Sg. Teku 29 March 1987 R. Kiew RK248] (K, L, SING, UPM).

Notes: This is a remarkable species of Didymocarpus with striking leaves that recall

those of kale (Brassica oleracea Acephala group, Cruciferae) in being glossy green

with the veins finely delineated in white. Hence its common name, the Kale-leaved

Didymocarpus. This feature is unique in Malayan Didymocarpi, for while several

species have leaves with a pale band down the centre, as for example plants of D.

curtisii Ridley, D. fasciatus, D. malayanus Hook. f., and D. puncticulatus Ridley,

only D. marginatus Ridley has a pale band along the midrib that extends partway

along the veins. Another herbaceous species with this same pattern of white-veined

variegation is Argostemma pictum Wall. (Rubiaceae).

The population of D. yongii is polymorphic for the presence of purple undersides

to the leaf. Leaves without this purple coloration are grass-green above, those with

purple undersides are dark green above, but in both cases the veins are white.

Didymocarpus yongii was discovered at the top of a low shaded cliff on the banks

of Sg. Teku above Kuala Teku by Yong Ghong Chong for whom it is named. It is

a precarious species to collect as it grows on loose soil on the downward sloping edge

of the cliff. Only one population was found, although there is an earlier collection

of a small plant from Kuala Tahan.

The species is not closely related to any other in Peninsular Malaysia. From its

habit, it falls within Ridley’s sect. Boeopsis, which he defined as having ‘leaves

crowded in a tuft at the top of a woody root stock, peduncles slender, flowers

usually small’. However, it differs from the species that Ridley included in this

section in its panicle of nodding flowers. It somewhat resembles D. rubiginosus

(see above) in that both have panicles of rosy-purple flowers. However, they are

strikingly different vegetatively: D. yongii is a larger plant with leaves with widely

spaced veins that in life are conspicuously white; the leaves of D. rubiginosus are

densely pubescent with many closely spaced veins and the leaves dry a rusty brown

colour. In addition, D. yongii is a lowland plant; D. rubiginosus is a montane plant

which grows above 1,000 m.

Acknowledgements

I am most grateful to the Director of Wildlife and National Parks for permission

to carry out research in Taman Negara; to Universiti Pertanian Malaysia which

funded the expedition and to St John’s College, Cambridge, which sponsored my

visit to Florence; to Yong G.C. and J. Dawn, without whom I should not have

reached G. Tahan at all; to our Batik guides and Temuan porters for their cheerful

co-operation; and to the curators of herbaria at CGE, FI, K and SING for permis-

sion to examine specimens in their keeping.

References

Burkill, I.H. (1927). Botanical collectors, collections and collecting: Gunung Tahan.

Gards’ Bull. Str. Settl. 4: 158-159.

64 Gard. Bull. Sing. 42(2) (1989)

Ridley, H.N. (1893). On the flora of the eastern coast of the Malay Peninsula. Trans.

Linn. Soc. 2nd Ser. 3: 267-408.

. . (1908). On a collection of plants from Gunung Tahan, Pahang. J. Linn.

Soc. Bot. 37: 301-336.

. (1915). The Botany of Gunung Tahan, Pahang. J. Fed. Mal. States Mus.

6: 127-202.

. (1923). Gesneriaceae. Fl. Mal. Pen. 2: 495-547.

Waterstradt, J. (1902). Kelantan and my trip to G. Tahan. J. Roy. As. Soc. Str. Br.

372do27:

The Genus Camellia (Theaceae) in Malesia

Florae Malesianae Precursores — LVIII, Part Three

HSUAN KENG

c/o Department of Botany,

National University of Singapore, Singapore

EFFECTIVE PUBLICATION DATE: 15 MAR 1990

Abstract

This is a taxonomic treatment of Camellia lanceolata (Bl.) Seem., the only Camellia species found

in the Malesian region.

Introduction

The genus Camellia was described by C. Linnaeus in the first edition of Genera

Plantarum (1737) in honour of Georg Josef Kamel (or Geogius Josephus Camelus

after latinization) (1661-1706), a Moravian Jesuit missionary. It was based on two

species then known: C. japonica L. and C. sasanqua L. Meanwhile, Linnaeus adopted

K. Kaempfer’s genus 7hea, which was based on 7: sinensis L., in the same publica-

tion. Owing to their androecial difference, these two genera were allocated under

two separate Linnaean ‘classes’, namely Camellia under Klass Monadelphia, and

Thea, under Klass Polyandria. Both genera were validated, according to our present

nomenclatural rules, in Linnaeus’ first edition of Species Plantarum (1753).

During the second part of the 18th and the early 19th centuries, as more species

of Camellia and Thea were collected from eastern Asia and subsequently described,

the androecial and other differences between Camellia and Thea became obsolete.

R. Sweet finally united them in 1818 and selected Camellia as the generic name. This

has been almost universally accepted today. For a detailed account of history and

discussions, see Cohen Stuart (1918) (in Bull. Jard. Bot. ser. III, pp. 232-238) and

Sealy (1958) (Rev. Gen. Camellia, pp. 1-6 and p. 14).

Among the synonyms of the genus Camellia, Calpandria (Gr. calpa — an urn,

andra — the male sex) is especially noteworthy. The sole Malesian species, Camellia

lanceolata (Bl.) Seem., was first described under Calpandria by Blume (1825). The

filaments of the outer stamens of this plant, as depicted in the original generic name,

are united to form a narrow fleshy tube. For this reason, it was regarded by Blume

as allied to Carapa of the Meliaceae. Calpandria was reduced to a section of Thea

by Pierre (1887), and to a section of Camellia by Cohen Stuart (1916). The latter

view is generally followed by Melchior (1925), Sealy (1958), Chang (1981), and many

others. Section Calpandria, according to Sealy, contains two known species, one,

C. lanceolata in Malesia, and another C. connata Craib, in northern Thailand.

In Sealy’s revision of Camellia, 82 species (plus a number of doubtful and im-

Part One: The genus Pyrenaria (Theaceae) in Malesia, Gard. Bull. Sing. 33 (1980) 264-289.

Part Two: The genus Gordonia (Theaceae) in Malesai, |.c. 37 (1984) 1-47.

66 Gard. Bull. Sing. 42(2) (1989)

perfectly known ones) are elaborately described and accurately illustrated. They are

classified under 12 sections. Later, another monographic study of Camellia was

carried out by Chang (1981), who enumerated a total of 196 species, distributed in

4 subgenera and 19 sections.

Taxonomic Treatment

Camellia Linnaeus

Camellia L. Gen. Pl. ed. 5 (1754) 311, et Sp. Pl. (1753) 698; Sweet, Hort. Suburb. Lond. (1818) 157;

Benth. in B. & H. Gen. Pl. 1 (1862) 187; Coh.-Stuart, Med. Proefst. Thee 40 (1916), et Bull. Jard. Bot.

Btzg. ser. 3, 1 (1919) 193; Melch. in E. & P. Nat. Pfl.-Fam. ed. 2, 21 (1925) 129; Sealy, Rev. Gen.

Camellia (1958) 14; Chang, Bull. Sun Yatsen Univ. (Nat. Sc.) 1 (1981) 1.

Thea L. Gen. PI. ed. 5 (1754) 232, et Sp. Pl. (1753) 515; Szyszyl. in E. & P. Nat. Pfl.-Fam. 3, 6 (1889)

183.

Calpandria Bl. Bijdr. 1 (1825) 178; Nakai, J. Jap. Bot. 16 (1940) 666.

Shrubs or small trees, rarely large trees, evergreen. Leaves alternate, spirally

arranged, serrate, mostly coriaceous. Flowers bisexual, terminal and axillary, solitary

or 2-3 in a cluster, pedunculate or sessile. Bracteoles 1 or 2 or more, distinguishable

or indistinguishable from the sepals. Sepals mostly 4-6. Petals 4-6 (-8), often more

or less united below. Stamens numerous, in several rows, briefly or highly connate

at the base and often adnate to the corolla, rarely free (in C. lanceolata); anthers

versatile. Ovary generally 3- or 5— loculate; ovules 3-4 (4-8 in C. /anceolata) in each

locule; styles 3-5, fused to varying extent proximally, rarely totally free. Capsule

woody, dehiscing loculicidally along a persistent columella and usually remaining

attached to it at the base. Seeds usually 1 or 2 (3-8 in C. lanceolata) in each locule,

generally globose or partly wedge-shaped, or irregularly shaped by mutual pressure;

testa crustaceus or woody; exalbuminous; embryo straight, with 2 large, often

hemispheric cotyledons.

Over 100 species distributed from Nepal, E. India, Burma, N. Thailand, Indo-

China to S. China and S. Japan and to Malesia. A great majority of species is con-

centrated in S. and S.W. China and Indo-China. Only one species, C. lanceolata,

is found in western Malesia, from Sumatra, Java, Borneo to the Philippines and

Sulawesi. In addition, two East Asiatic species, namely, C. sinensis (L.) OK., from

S. China, and C. japonica from S. China and Japan, are introduced and planted

at higher elevations in Malesia. The former species is cultivated in plantations in

Java, the Malay Peninsula, Luzon and elsewhere, and the latter is occasionally planted

as an ornamental.

Camellia lanceolata (B\l.) Seem.

Camellia lanceolata (Bl.) Seem., Trans. Linn. Soc. Lond. 22 (1859) 345; F.-Vill., Nov. App. (1880)

19; K. & V., Bijdr. 3 (1896) 303; Coh.-Stuart. Med. Proefst. Thee 15 (1916) 120, £/0, //, Bull. Jard. Bot.

Btzg. ser. 3, 1 (1919) 238, 281, f/2, 13; Sealy, Rev. Gen. Camellia (1958) 141, £66; Back. & Bakh. f.,

Fl. Jav. 1 (1963) 319; H.T. Chang, J. Sun-Yatsen U. (Nat. Sci.) 1 (4981) 108.

Calpandria lanceolata B\., Bijdr. 1 (1825) 178; Korth. Kruidk. (1839) 148, 4.337; Miq., Fl. Ind. Bat.

2 (1857) 492.

Calpandria quiscosaura Korth., Kruidk (1840) 149, 43/7; Miq. Fl. Ind. Bat. 2 (1857) 493.

Salceda montana Blanco, Fl. Filip. ed. 2, (1845) 374, ed. 3, 2 (1879) 327.

Camellia quiscosaura (Korth.) Seem., Trans. Linn. Soc. Lond. 22 (1859) 345.

Pyrenaria sp. Vidal, Rev. Pl. Vasc. Filip. (1886) 57.

The Genus Camellia

Pig. ft.

Camellia lanceolata (Bl.) Seem. Java, Herb. Lugd. Bat. No. 918, 190-2] (habit), Junghuhn

10 (flower bud and floral parts), Murata et al. 880 (dehiscent fruit and seeds); Sumatra, Soegen

625 (indehiscent fruit).

68 Gard. Bull. Sing. 42(2) (1989)

Thea lanceolata (Bl\.) Pierre, Fl. Coch. 2 (1887) sub t.119; Merri. En. Philip. 3 (1923) 69 (incl. vars.

mollis, microphylla and stenophylla).

Thea quiscosaura (Korth.) Pierre, l.c. sub t.119.

Camellia minahassae Koord., Med. Lands Pl. Tuin 19 (1898) 350, 643.

Thea lasiostyla Warb. ex Kochs, Bot. Jahrb. 27 (1900) 579, 582.

Thea montana (Blanco) Merr., Philip. Bur. Govern. Lab. Pub. 27 (1905) 21.

Thea megacarpa Elm., Leafl. Philip. Bot. 5 (1913) 1842.

Thea minahassae (Koord.) Koord.-Schum., Syst. Verz. 3 (1914) 86.

Camellia megacarpa (Elm.) Coh.-Stuart, Med. Proefst. Thee 40 (1916) 68.

Shrub or small tree, 3-6 m, rarely to 9 m tall; young branches hirsute, older ones

glabrescent or glabrous. Leaf-blades lanceolate, oblong-elliptic to elliptic, apex

obtuse or acute, attenuate or nearly rounded at base, 5-10 (-14) cm long, 2-3.5 (-S)

cm wide, thin leathery, the margin bluntly serrate or serrulate, glabrous above, the

lower surface usually villose on young leaves and nearly glabrous on mature ones

except sometimes on the elevated midribs; side veins 6-7 pairs, usually less distinct;

petioles short, 3-7 (-10) mm long, pubescent and then glabrous. Flowers solitary,

or sometimes 2, rarely few together, in upper leaf-axils, sessile or nearly so; sepals

4-6 (-8), unequal, ovate or suborbicular, 2-7 mm long, apex rounded or retuse,

pubescent externally. Corolla white or yellowish green, 1.5—2 cm across; petals 4—5

(-8) free, imbricate, oblong-lanceolate to elliptic, 7-12 mm long, glabrous, some-

times sericeous on the tips of some petals. Stamens 25-40; filaments of the outer

ones entirely united into a fleshy tube, 6-8 mm long, most anthers adherent to the

inside of the filamental tube, sessile or on tip of the free portion of filaments; 5-8

inner stamens free from the tube. Gynoecium 6-7 mm long; style solitary, nearly

as long as the ovary, 3-branches above; ovary 3-loculate, densely tomentose. Cap-

sule green to pale white grey, mostly flattened, 3-lobed, 1.5-2 cm long, and 2-2.5

(-3) cm wide, dehiscing loculicidally from the above into 3 thick woody valves.

Seeds 2-4 (-7) in each locule, obconical or semi-spherical, 1-1.5 cm long, the upper

(or outer) surface nearly rounded, the lower (or inner) surfaces usually in 2-3 flattened

planes due to mutual compression. Embryo with 2 large thick cotyledons; endosperm

absent.

Distribution: Malesia (Sumatra, Java, Borneo, The Philippines, and Sulawesi).

Specimens cited below are representative ones only, all from Leiden (L).

Sumatra: Sine loc. Forbes 2692; Bancoolen, Soegeng 625.

Java: Sine loc. Collectors unknown. Herb. Lugd. Bat. nos. 952, 54-077, 952, 54-943,

908, 190-21; G. Salak, Blume (?) s.n. Herb. Lugd. Bat. nos. 908, 189-180, 908,

190-195 (lectotype of Calpandria lanceolata Bl.); G. Geulis, Preanger, Bakh. v/d Brink

5972; Sine loc. Junghuhn 84, 90, 93, 141, 179 & 429; Sine loc. Koorders 8127, 8128,

8143, 1844 & 8145; Sine loc. Korthals s.n. Herb. Lugd. Bat. Nos. 908, 191-1201, & 908,

191-1202 (type materials of Calpandria quiscosaura Korth.) G. Slamet, C. Java,

G. Murata et al. J. 880; Sine loc. Zollinger 1623.

Borneo. Sabah: Mesilau River, Chew & Corner RSNB 7099; Marai Parai, Clemens

32483; Ranau, Aban Gibot SAN 68580; Sandakan, F.E. & Aban SAN 65455. Kali-

mantan, N. of Balikpapem, Kostermans 4274; Mt. Medadem, Sangkulirang Dist.

Kostermans 13386.

The Philippines: Dist. of Lepanto, Luzon, Curran 10952; Bayninau, Mt. Prov.

Luzon, Conklin & Buwaya 80654; Mt. Pico de Loro [lols Prov., Luzon, Edano 17346,

The Genus Camellia 69

17946; Baguio, Benquet, Luzon, Elmer 8349, 14282; Puerto Princesa (Mt. Pulgar),

Palawan, Elmer 12822 (iso-type of Thea megacarpa Elm.); Mt. Tabayoc, Luzon,

Jacobs 7516; Guimaras Island, Sulit 1/801.

Sulawesi G. Bonthaim, Bunnemeyer 12]45; Lombasang, Bunnemeyer 11130, 11321,

11475; Mandao, Minahassa, Koorders 18100 B, 18992 B, 19286 B, 19287 B (type

materials of Camellia minahassae Koord.); Gowa, Garentang, Neth. Ind. For. Serv.

no. bb. 20438; Manado, Neth. Ind. For. Serv. no. bb. 18081; Palopo, Melenjong,

Neth. Ind. For. Serv. no. bb. 24132.

Ecology: In primary and secondary forests from lowlands near sea level to high

altitudes (2,000-2,500 m). It also occurs on limestone hills in Kalimantan (Koster-

mans 13386). Flowering and fruiting almost year round.

Note: This species, as noted by several earlier workers, is highly variable in degree

of pubescence of branchlets and leaves, in size and number of sepals (perules) and

petals, and in size and shape of fruits and seeds. Based on the leaf characters of the

Philippine materials alone, Merrill (1923) recognized 3 varieties (namely, var. mollis,

microphylla and stenophylla). Sealy (1958, p. 144) emphasized the shape and number

of the perianth parts between the Javanese and Philippino specimens. Furthermore,

attempts to establish separate species on some local populations, e.g. Calpandria

quiscosaura Korthals from Java, Thea megacarpa Elmer and Thea montana (Blan-

co) Merrill from the Philippines, and Camellia minahassae Koorders from Sulawesi,

have been proved futile. A species widely distributed from central southern Sumatra

to the northern and southern tips of Sulawesi, and from sea level to nearly 2,500

m, is understandably unlikely to be a homogeneous taxon. Yet the androecial fea-

tures are so unique, and the gynoecial, fruit and seed characters so generally uniform,

that only exhaustive field examination can perhaps determine and discrimate the

variants.

Acknowledgements

I am grateful to the Director and staff of the Singapore Botanic Gardens, for the

use of the herbarium and library facilities, and to the Director and staff of the

Rijksherbarium, Leiden for the loan of the entire collection of Camellia specimens.

I would like also to thank Mrs Ng Siew Yin for her kind assistance in several ways

and my wife, Mrs Ro-siu L. Keng for preparing the illustration of this paper and

for her encouragement.

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A New Species of Licuala (Palmae) from Peninsular Malaysia

SAW LENG GUAN! and JOHN DRANSFIELD?

'Forest Research Institute Malaysia, Karung Berkunci 201, 52109 Kuala Lumpur, Malaysia.

?Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K.

EFFECTIVE PUBLICATION DATE: 15 MAR 1990

Abstract

A new species of understorey palm, Licuala thoana L.G. Saw & J. Dransf. is described from Johor,

Peninsular Malaysia.

Introduction

Centred in the Malesian region, there are about 25 species of Licuala native to

Peninsular Malaysia (Furtado 1940). This genus of mostly understorey fan-leaved

palms is one of the most conspicuous and common components of the forest under-

growth. Most species are rather local in distribution. In fact in Peninsular Malaysia

alone, about 19 species are said to be endemic. This rather diverse genus is greatly

in need of a general revision.

Five species of Licuala were recorded in the Ulu Endau area (Dransfield & Kiew

1987) during the Malaysian Heritage and Scientific Expedition to the Endau-Rompin

area (Kiew et al. 1987). Two of these are endemic to Johor (L. kiahii Furt. and L.

lanuginosa). With the discovery of this new species, another rare but locally com-

mon species is added to the area of the proposed National Park. Although the genus

needs a general revision, we feel the need to describe this new species in order to

draw attention to the uniqueness of this area from a conservation standpoint.

All the native peninsular Malaysian species of Licuala have fronds divided into

segments and generally circular in outline. This new species is distinct in having

fronds that are usually undivided and paddle-shaped. The species epithet honours

our friend and colleague, Dr. Tho Yow Pong of the Forest Research Institute

Malaysia, for his enthusiasm and diligence for conserving the Endau-Rompin area.

Licuala thoana L.G. Saw & J. Dransf. sp. nov.

A ceteris speciebus Peninsulae Malayanae folio plerumque indiviso statim distinguibilis; floribus dense

ferrugineo-lanuginosis et fructibus immaturis roseis L. ferrugineae Griff. affinis sed folio et inflorescen-

tia minima bene distincta. Typus: Johor, Saw Leng Guan FRI 36354 (holotypus KEP: isotypus K).

Solitary acaulescent undergrowth palm. Stem subterranean, c. 30 mm diam. Leaves

c. 8 in crown, marcescent; leaf base sheathing in the basal 10 cm, c. 5 cm wide at

the insertion, the sheath lacking a conspicuous persistent ligule and disintegrating

into coarse, rather fragile fibres, the abaxial sheath surface bearing scattered caducous

dot-like scales; petiole 21-40 cm long, triangular in cross section, c. 7 mm wide, c.

5 mm thick, armed through much of its length with rather regularly arranged, marginal

spines to 2.5 x 1 mm, the spines shorter or lacking in the distal portion, petiole

glabrous adaxially, abaxially bearing scattered dot-like scales; leaf-blade usually en-

tire, broadly paddle-shaped, rarely divided into 3 segments, the two lateral narrower

than the centre, the entire blade 34-51 x 25-30 cm, lower margins smooth, distal

W2 . Gard. Bull. Sing. 42(2) (1989)

Fig. 1. Licuala thoana. A sheathed stem and one leaf; B inflorescence; C flower; D flower, one petal

removed to show androecium; E petal, interior view; F petal exterior view; G gynoecium. All from FRI

36354. Drawn by Saw Leng Guan.

A New Species of Licuala 73

margin shallowly induplicately lobed, with shorter indentations associated with abaxial

ribs, and deeper indentations associated with adaxial ribs, in all the blade with c.

17 folds on each side of the costa, blade glabrous adaxially, abaxially with scattered

caducous brown indumentum, especially along the folds. Inflorescence interfoliar

to 30 cm long, bearing 2-3 spicate partial inflorescences; peduncle to 21 cm long,

semi-circular in cross section, c. 4 mm wide at the base; prophyll strongly 2-keeled,

strictly tubular, c. 8 x 1.2 cm, apically becoming somewhat fibrous, abaxially covered

in abundant rusty-brown indumentum; bracts subtending partial inflorescences 3,

strictly tubular, 25-50 x 7-11 mm, distally disintegrating into soft fibres 8-25 mm

long, abaxially the bracts covered with rusty-brown indumentum; rachillae to 45 x

2.5 mm, very densely covered in rusty-brown tomentum composed of hairs to 1.5

mm long, rachilla bracts minute, obscured by the hairs. Flowers irregularly triangular-

ovoid, c. 5 x 5 mm, borne singly on short protuberances c. 0.5 mm high, 1 mm

diam.; calyx very thick, coriaceous, tubular in basal 1 mm, with 3 gibbous, rounded-

triangular lobes to3 x 3 mm, the margins entire, abaxially covered in shaggy brown

hairs to 0.3 mm long; corolla very thick, coriaceous, explanate at anthesis, only

slightly longer than the calyx, tubular in basal 2 mm, with 3 triangular lobes 2 x

3 mm, abaxially covered in densely adpressed brown hairs, adaxially lobes marked

with impressions of anthers; staminal ring borne at the mouth of the corolla tube,

0.7 mm high, the 6 free filaments equal, c. 0.6 mm long, anthers somewhat api-

culate, 1.2 x 0.8 mm; ovary turbinate, c. 2 x 1.5 mm, covered with a ring of dense

adpressed brown hairs at the widest point, style slender, | x 0.1 mm. Mature fruit

not known; immature fruit with 1-3 carpels developing, 5 x 2 mm in available

material, bright pink.

Peninsular Malaysia. Johor: Labis Forest Reserve, Sungei Kinchin, logged over

lowland dipterocarp forest, alt. 30 m, 26 Aug 1988, Saw Leng Guan FRI 36354

(Holotype KEP; isotype K).

Ecology: An understorey palm of lowland dipterocarp forest, very common along

the Sungei Kinchin flood plain. The palm persists in forest that has been logged

although showing signs of frond yellowing when exposed. L. thoana is known from

this basin and surrounding areas only; it has not been observed at higher elevations,

for example the Gunung Beremban massif or the upper elevations of Gunung Keriong.

Both mountains are adjacent to the Sungei Kinchin basin. Neither was the palm

observed further downstream along the Sungei Endau or the Sungei Jasin.

This is an unusual and easily identified species, immediately distinguishable in

Peninsular Malaysia by its usually entire, paddle-shaped leaves, the short inflorescence

with unbranched partial inflorescences with axes and flowers all densely covered in

rusty-brown hairs and the pink young fruit. In the indumentum of the inflorescence

and the fruit coloration it approaches L. ferruginea Griff., but the latter is a robust

acaulescent species with large leaves divided into many segments and with highly

branched inflorescences.

References

Dransfield, J. and Kiew, R. (1987). An annotated checklist of palms at Ulu Endau,

Johore, Malaysia. Mal. Nat. J. 41: 257-265.

Furtado, C.X. (1940). The genus Licuala in the Malay Peninsula. Gard. Bull. Str.

Settlements 11: 31-73.

Kiew, B.H. et al. (1987). The Malaysian Heritage and Scientific Expedition: Endau-

Rompin, 1985-1986. Mal. Nat. J. 41: 83-92.

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Book Review

Tropical Woody Rubiaceae

By E. Robbrecht

Opera Botanica Belgica Vol. 1, Nationale Plantentuin van Belgie, Meise, 22 December 1988. 272 pages;

61 illustrations, 5 tables. ISBN 90- 72619-02-1. ISSN 0775-9592. 29 cm, softcover, threadbound.

For those who still want to know, the Rubiaceae (with 659 genera enumerated

by Robbrecht and an estimated 10,700 species) are the fourth largest angiosperm

family, after Asteraceae, Orchidaceae and Fabaceae. This family with 29 of its 38

currently accepted tribues being predominantly woody, has a mainly tropical distri-

bution, and is represented in temperate areas by a comparatively small number of

herbaceous taxa. How else to better understand such a family than through a proper

understanding of its woody representatives?

Robbrecht’s book aims at a synthesis of available information from the literature

(and there are 325 listed references in this book, which does not cover all publica-

tions on the family) and his own studies, mainly of tropical African representatives.

There is a useful brief description of methods used in studying domatia, crystals,

hairs and excrescences, microcharacters of the corolla, pyrenes, seed-coats, endo-

sperm and pollen, before the author launches into a detailed discussion (exceeding

a hundred pages) of ‘‘characteristic features of tropical woody Rubiaceae’’, and a

short general discussion of subfamilial classification. There are four appendices which

are very useful: (1) a survey of the classification proposed in the book; (2) additions

and corrections for the Rubiaceae in the Index Nominum Genericorum since 1979

and until October 1988; (3) an index to taxa in the present work and to the treat-

ments by Schumann (1891, 1897, 1900), Krause (1908, 1915), Verdcourt (1958) and

Bremekamp (1966); (4) a list of genera accepted by Robbrecht and their synonymy

and tribal or subtribal position.

The most interesting chapter (for reading) is that which surveys different characters

in the family. This includes habit and architecture; peculiarities in lifeform; the

morphology and anatomy of the vegetative axis (roots, stems, nodes and wood); leaf

characters; stipules; exudates, crystal types, trichomes and colleters; inflorescences;

flowers; fruits; seeds; pollen; chromosomes; biology; distribution. The illustrations

(line-drawings and photographs) are used to good effect. Mention of a few more

characteristics may be interesting additions to an already amazing treatment by Rob-

brecht. There are also epiphytes in Argostemma (see Robbrecht’s Table 3) and Aidiop-

sis forbesii in Malesia apparently has a strangling habit unique in the family. There

is a discussion of geofrutices (plants with massive woody subterranean stems which

sometimes ramily, notable in drier parts of Africa) and in this connection it is in-

teresting to note that Euclinia longiflora (Africa), Ixora coccinea and I. finlaysoniana

(Asia) can produce root suckers and also originate from somewhat monsoonal

climates. The Malayan Ixora scortechinii var. scortechinii is unusual in that its rosette-

like leaf arrangement traps organic litter and its internodes put out fine roots that

penetrate the organic matter accumulated; it is a free-standing terrestrial plant.

Intrapetiolar rather than interpetiolar stipules, in a way quite out of character in a

rubiaceous plant, also occur in Mussaendopsis beccariana in Malesia. Shoot-tip and

bud waxes in Gardenia also occur for the Southeast Asian species. Yellow sap (in-

credible but true) exudes from the bruised bark and fruit of some Rothmannia

species in Malaya and Borneo. Some Naucleeae even produce a reddish sap! Mitriform

76 Gard. Bull. Sing. 42(2) (1989)

stigmas or types structurally close to this, important in Mitragyna and many Van-

guerieae, are not discussed. Some Uncaria species also consistently have anti-inhabited

twigs. Robbrecht has omitted mention of the claim (Tan and Rao 1981, Biotropica

13: 232) that vivipary exists in Ophiorrhiza; that is indeed not vivipary but rather

germination in situ, because the many tiny seeds are not always effectively expelled

from the purse-shaped capsules, where their germination occurs without any physio-

logical connections between parent and offspring tissues. The family is certainly not

only huge, but also rich in peculiarities. Chemotaxonomic characters are surveyed

in the chapter dealing with subfamilial classification, where it is perhaps more

appropriate.

Subfamilial classification and the problems of grouping genera into tribes is

clearly presented. Two subtitles on the title page of this book, ‘‘Characteristic features

and progressions’’ and ‘‘Contributions to a new subfamilial classification’’, indicate

how the survey of characters and trends are used here to evaluate and modify the

classifications proposed by Hooker (in 1873), Verdcourt (1958) and Bremekamp

(1966). Robbrecht makes good use of Venn diagrams displaying the various tribes

for indicating character distribution and comparing classification systems. Hooker

recognised 2 subfamilies, Verdcourt, 3 and Bremekamp, 8; Robbrecht suggests 4 —

Cinchonoideae, Ixoroideae, Antirheoideae and Rubioideae.

The assignment of ‘‘problematic’’ genera to tribes, and even tribes to subfamilies,

can remain difficult, often due to a lack of information. We may look at some

Southeast Asian genera in relation to this. Robbrecht remains uncertain where

Jackieae (represented by Jackiopsis, which has a 3-merous calyx and peculiar winged

fruit) will affiliate in the Rubioideae. He has synonymised the Coptosapelteae (with

contorted corolla lobes, unique T-shaped trichomes and a lack of raphides) with the

Cinchoninae (with imbricate or valvate corolla lobes and raphides present or not)

in the Cinchoneae. This demonstrates that tribal distinction is not easy, because

variable and constant character-states are used even for different subtribes within

a tribe. Bremekamp’s Pomazoteae is regarded by Robbrecht as a synonym of Hedyo-

tideae, with the exception of Klossia which is placed in the related Ophiorrhizeae;

no clear justification is made. Bremekamp credited the Pomazoteae with no raphides

although raphides have been subsequently discovered in Lerchea, Xanthophytum

and Pomazota; however there is little comparable information mentioned for K/ossia.

The laterally flattened capsules of the Ophiorrhizeae seem to me quite distinctive.

The odd genus Scyphiphora is put by Robbrecht in the Antirheoideae rather than

in the Gardenieae of the Ixoroideae, based on the opinion that the unique placenta-

tion in Scyphiphora (with one pendulous and one upwardly directed ovule inserted

on a mid-septal placenta) may be derived from the Retiniphyllum (with 2 pendulous

ovules attached to a mid-septal placenta) type (Retiniphylleae, Antirheoideae). Yet

Scyphiphora has mesophyll sclereids so far detected only in seven other genera, all

belonging to the Gardenieae and associated tribes.

It seems reasonable that Robbrecht has placed the Acranthereae and the ‘‘Mus-

saendeae’’ (illegitimate name) synonymous with the Isertieae (Cinchonoideae). But

Lecananthus to me seems to have greater affinities with the Schradereae (including

the Malesian Lucinaea and South American Schradera; Rubioideae) than the Iser-

tieae: Lecananthus has adhesive adventitious roots and congested flowers grouped

in heads, and lacks the capsules and winged seeds so often characteristic of the

Cinchonoideae. Similarly, Mycetia is more often associated with the Hedyotideae

(Rubioideae) than the Isertieae. The placement of Lasianthus in the Morindeae

(which seems well characterised by a strong tendency towards syncarpous fruits) rather

than the Psychotrieae is also controversial.

Such characters as presence or absence of raphides have been misinterpreted or

Book Review 77

even wrongly ascribed to taxa in the past, and definitions of tribes and subfamilies

have also shifted back and forth. In his work, Robbrecht has obviously attempted

to verify as much as possible the information reported earlier, and comes out with

what I consider the most important review and synthesis for the Rubiaceae. There

are few typos and inconsistencies, e.g., ‘“Anthocephalus’’ (which Bosser considers

a synonym of Breonia) is used on page 29, but is this intended to mean Breonia or

Neolamarckia, the latter name given by Bosser for Ridsdale’s intended genus? Rob-

brecht hopes his book has emphasized adequately that a concerted effort by all workers

is needed for a clearer understanding of such a complex family. His book is such

an excellent new chapter in our understanding of the Rubiaceae.

Reviewed by:

K.M. Wong

Forestry Department,

Bandar Seri Begawan 2067,

Brunei Darussalam

With Compliments

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GRASS-CUTTING, LANDSCAPING, TURFING

AND ALL HORTICULTURAL OPERATIONS.

CHYE THIAM ENTERPRISES CO

BLK 208 HOUGANG STREET 21

#02-205 SINGAPORE 1953

TEL: 2841938

LIP GUAN CONSTRUCTION PTE LTD

BLK 2002 BUKIT BATOK IND. PARK “A”

#03-38 SINGAPORE 2365

TEL: 5636633, 5636634 FAX: 5630626

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PARKS AND RECREATION DEPARTMENT

PUBLICATIONS FOR SALE

1. The Gardens’ Bulletin, Singapore (Series IV).

Price.

13(1) new impression: $12 33(1): $21.50, 33(2): $12.50, Index: $1.90

17(3): $12.50 34(1): $21.50, 34(2) with Index for 34: $11.50

18 & 19: $25 per vol. 35(1): $17.50, 35(2) with Index for 35: $21.50 |

20(1): $8 36(1): $21.50, 36(2): $14.00

25(1): $9, 25(2): $12 37(1) with Index for 36: $18.50

26(1): $18, 26(2): $18 37(2) with Index for 37: $16.50

27(1): $18, 27(2): $8.50 38(1): $20.00, 38(2): $13.00

28(1): $18, 28(2): $15 39(1) with Index for 38: $15.50-

29: $30 39(2): $15.50

30: $48 40(1) with Index for 39: $15.50

31(1): $10, 31(2): $12.50 40(2): $13.00

32: $15.50 41(1) with Index for 40: $6.50

The Freshwater Swamp-forest of S. Johore 41(2): $14.00

and Singapore by E.J.H. Corner (Gard. 42(1 & 2) with Index for 41: $15.00

Bull. Sing. Suppl. 1) $35

2. Selected Plants & Planting for a Garden City — Forty Shrubs, $1.20.

3. Selected Plants & Planting for a Garden City — Forty Climbers, $3.00.

4. A Guide to Tree Planting, $4.00.

5. Malayan Orchid Hybrids by M.R. Henderson and G.H. Addison, $15 (1969).

6. A Revised Flora of Malaya.

(a) Vol. 1, Orchids, by R.E. Holttum, $50 (3rd ed. 1980 Impr.).

(b) Vol. 2, Ferns, by R.E. Holttum, $20 (2nd ed. 1968).

7. Boletus in Malaysia by E.J.H. Corner, $50 (1972).

Items 1-4 can be purchased from the Commissioner, Parks & Recreation Department,

Botanic Gardens, Cluny Road, Singapore 1025; tel. no. 4709917.

For overseas orders, payment should be by bank draft or International Money Order and made

payable to the Commissioner of Parks & Recreation, Singapore.

Items 5-7 can be purchased from Singapore National Printers (Pte) Ltd, 303 Upper Serangoon

Road, Singapore 1334, tel. no. 2820611 ext. 240. 241 or 231.

All prices quoted are in Singapore. Dollars

Overseas postage is extra