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CONTENTS

B. MOLESworTH ALLEN: Dr. R. E. Ho_truM: An Appreciation - - :

H. M. BurKILL: Richard Eric Holttum, Croix de Guerre. A note - -

A. C, JeErmy: Dr. R. E. Holttum — Eightieth Celebration - - - -

K. C. CHEANG & A. G. ALPHONSO: Holttum’s contribution to horticulture in ae

the Malaysia — Singapore region - - - - - 9-12-39

BARBARA JOE HOSHIZAKI: Staghorn Ferns Today and Tomorrow - - - 13-15 4

SOEJATMI SOENARKO: A New Species of Nastus from Sumba - - - 17-19

Davip W. LEE: On Iridescent Plants - - - - - - 21-29 =

A. G. Piccotr: The Ferns of Gunong Ulu Kali - - - - 31-43

G. J. de JONCHEERE: Specific Concept in Humata pectinata - or) STG gpa 45-58

B. L. Burtr: Curcuma zedoaria - - - - - - - 59-62

K. IwaTSuKI: Studies in the Systematics of Filmy Ferns II. A note on Meringium 4

and the taxa allied to this - - - - - - 63-74

J. DRANSFIELD: Calamus caesius and Calamus trachycoleus Compared - - 75-78 a

K. U. KRAMER: Synaptospory: a hypothesis. A possible function of spore =

sculpture in pteridophytes - - - - - - - ito 4

E. SOEPADMO & E. E. ona ee Motpheioey oe Sage oe Dennsiaciia ae

sensu Holttum~ - 85.95

W. R. Pui_ipson: An Enumeration of the Malesian Species of Aralia - - 97-100, ‘2

LIEW FAH SEONG: Scanning Electron Microscopical Studies on the Spores of aa

Pteridophytes XI. Oleandraceae - - - - = : 101-110 Bh

i 122

R. D. HOoGLAND: Saurauiae Gerontogeae I. Notes on Malayan species

Tetsuo KoyaMa: The Cyperaceae Tribe Cypereae of Ceylon - - 123-164

S. C. CHIN: The Limestone Hill Flora of Malaya I - - - ‘ = 165-219

J. A. CRABBE: Holttum’s New Taxa and Name-changes in Ferns - - - 221- 238

M. G. Price: Philippine Dryopteris : ; F So eee ©

WARREN HERB WAGNER JR. & FLORENCE S. WAGNER: Fertile-sterile Leaf

Dimorphy in Ferns - - - - : a : .

GUNNAR SEIDENFADEN: A note on Dendrobium serpens - - - -

BENJAMIN C. STONE: Notes on the Systematy of Malayan Phanerogams XXV.

Araliaceae - - - - - - - - -

A. C. JermMy & T. G. WALKER: A note on the cytology ba Batrychiags

lanuginosum and the occurence of genus in Malesia - 2

W. R. PHILIPSON: A revision of the Malesian species of Arthrophyllum - -

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" i Atoohe i

A SPECIAL ISSUE TO MARK

THE EIGHTIETH BIRTHDAY

OF R. E. HOLTTUM, 1975

powyp uig jwwsy <q ydoisojoyd

siodesulg ‘uspieH suRjog “WINTIegiox{ 94} 32

SL61 ‘WOLLIOW OMNW AAVHOIYy

THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XXX 10th October, 1977

Dr. R. E. HOLTTUM: An Appreciation

B. MoLeswortH ALLEN

Los Barrios, Spain

Dr. Holttum when writing about the classification of the Leptosporangiate

ferns, most of which in the past were united in the Polypodiaceae, stated that

“these ‘modern’ ferns are so extremely varied in form and in the details of all

their parts, that no one man could pretend to a critical knowledge of them all,

even as to their assignation to genera.”

It is strange that he should write this, for his amazing faculty for critical

taxonomic interpretation has enabled him to understand so much about these

living ferns, both in the field and herbaria. Thus he may well have come as near

to having just this critical knowledge about which he writes, as any one man may

possibly ever have. Future pteridologists will have not only the advantage of our

present-day work, but computers will be used to store much of the knowledge

that people of Dr Holttum’s age have had to keep in their heads. So although

there will possibly be a more complete taxonomic and cytological understanding

of the world’s ferns in the future, this will surely be a combination of man and

his aids, and not held by one man alone owing to the limit of our brain power.

The steps that Holttum stands on today were made firm by Carl Christensen

who brought fern classification into the 20th century, and by the important

morphological work of Bower and others. In turn each pteridologist was fortunate

in having the published works of great thinkers of the past who were responsible

for these higher steps from which Christensen and Copeland worked.

No doubt Holttum was fortunate in that fate pointed the way at the beginning

of his career, which started as a palaeobotanist under the guidance of no less a

scientist than Dr Seward. This gave him an excellent background for the phylogeny

of the living ferns when he arrived in Singapore and commenced his taxonomic

work, which he was to continue for the next 40-50 years. Again, the area was a

fortunate choice; its rich fern flora with so many distinct and unrelated genera

must have given him much scope for his enquiring mind. The many relic ferns

growing in their natural environment, gave living examples of evolutionary evidence

towards a new understanding of the ancestry of different genera and their relation-

ship. The hot, wet climate of Singapore and the Malay Peninsula, without such

natural calamities as typhoons and long droughts, was conducive to the continua-

tion of these primitive types of ferns, many of which were not greatly changed

from their ancestors. Matonia is one such example of a family once widespread

in Mezozoic times and some fossils have been found with almost unchanged frond

shape. The distribution of Matonia pectinata is now extremely limited and is

2 Gardens’ Bulletin, Singapore — XXX (1977)

confined to a relatively few hill and mountain ridges in the Malay Peninsula and

Borneo, growing where the soil is leached and poor, and where it is less likely to

suffer competition from the surrounding lush tropical plantlife.

All this was at his doorstep so to speak, and he made the most of it which

many of us have not done, and no doubt he was helped and guided by Copeland’s

work on Philippine ferns. One assumes that during these years his fertile mind

was gathering and storing information on the classification and evolution of the

vast array of the ferns before him. He also assimilated much knowledge of the

flowering plants as well.

Then in Singapore between 1942-46 he was able to correlate this knowledge,

for being cut off from the outside world he was able to work in the herbarium

of the Singapore Botanic Gardens, almost without interruption. This resulted in

the impressive taxonomic publication on the ferns of the Malay Peninsula and

Singapore. In the course of this work, significant facts came to light which altered

much of the previously accepted concept of the fern classification of Christensen

and others. He also pointed out that there were, at least, several families which

needed far more study especially in field observation, one of these being the

Thelypteridaceae. He kept Thelypteris more or less together thus apposing Cope-

land (Genera Filicum 1947) who contended that this generic name was invalid,

Lastrea being the legimate one. Whilst not actually disagreeing with this, Holttum

maintained that many changes would have to be made in this group of ferns,

and until this was done, Thelypteris should stand. As we know, it is a revision

of this vast and complex family that Holttum himself has undertaken. and from

his active brain we are seeing these ferns in a new light, and currently a series

of new combinations and genera are emerging. He has also proposed the conserva-

tion of the name Thelypteris.

So the publication of the Ferns of Malaya in 1954 with its appendix con-

taining cytological notes of a 100 Malaysian ferns by Professor Manton, began a

new era towards the knowledge and understanding of these tropical ferns. At last

there was a critical study of this area with up to date name combinations basinyms

and sufficient synonyms from which one could work.

As those of us who had followed Christensen, Bower and Copeland (and for

me, Dr Holloway) in our formative years, this book as well as his papers on

classification, meant yet another pteridological step upwards. Living in Malaya at

the time, this publication began a new era for me also. I arrived with a copy of

Beddome’s “Ferns of British India, Ceylon and the Malay Peninsula” (1892),

and a separate of Holttum’s ‘““The Common Ferns of Singapore” which had been

written for the Singapore Naturalist in 1924. (It was almost impossible to obtain

copies of van Alderwerelt van Rosenburgh’s handbook on the ferns of the Malayan

Islands).

The diversity of the ferns with so many unfamiliar genera, not to mention the

richness of the species of these tropical rain forests, was enough to daunt anyone

without constant access to herbaria and a botanical library. Ridley’s “‘Ferns of the

Malay Peninsula” (1926) was very limited in its list of species, whilst many of the

names used were difficult to trace, so that it helped only in a small way. I found

that Holttum’s key in his paper together with the notes on the ferns, gave a good |

introduction to the genera one encountered every day, but the vast number of other

ferns one saw in the jungle was over-powering. It was a lengthy business, often

empirical. to try to determine species, usually ending in a deep depression of doubt

which hardly encouraged intelligent collecting, unless one was a collector by

instinct So Ferns of Malaya made an enormous difference to those interested in

the S. E. Asian ferns.

My association with Dr. Holttum, who was a friend of my husband’s parents,

has spanned many years, but it differs from most fern botanists in as much as I

was never long enough in any botanical department to undertake fern research, so

An appreciation 3

reluctantly became a field botanist, observing and collecting. Holttum never failed

to give me encouragement in this, especially when it became tedious. It was

stimulating to be in the jungle with him, for his field knowledge, not only of

ferns, but also of the angiosperms, was immense, as was his enthusiasm. An

interesting plant would be heralded by “a-ha, a-ha”, as he bent to look at it and

use his lens. On these periodic field trips I learned about tropical botany; he

pointed out the strange stamens of Lauraceae, and the amount of caulifery in

these rain forests. He seemed untroubled by the continual heat and high humidity,

and the rough jungle floor affected him far less than me, even although he had

suffered an attack of polio after the war. He was an easy houseguest having

continual good spirits, and only once do I remember that these deserted him. It

was when he brought the sad news of the death of our mutual friend, A.H.G.

Alston in 1958. The passing of this gentle person cast a shadow over our trip that

time, which was to see the extent of Cyathea polypoda on Mt. Kledang near Ipoh,

where it had been locally common.

Holttum encouraged _ selective

collecting, stressing the importance

of ecological factors, and although

not, I think, a patient man especially

with what he considered stupidity,

he would give painstaking help to

the student of any branch of botany.

He also had the gift of being able

to impact his knowledge easily, cy. Me WWHfP

which is reflected in many of his ee RS?

students during his professorship : 12 oN

at the University of Malaya in 1s 3 wie

Singapore, who have since become | Aza eZ

competant botanists. He wrote in x ie ea

simple language to encourage the \ 3 yp

beginner and his “Plant Life in Oa ae:

Malaya” (1954) is an excellent Wie hem

example of simplicity and erudition. eS

The output of work is staggering a

when one realises that apart from “ff WS

the Ferns of Malaya he published Grammitis

a companion volume on the orchids; holttumii

a horticultural textbook “‘Gardening

in the Lowlands of Malaya” (1953) as well as the monographs on gingers and

bamboos and the current one on the Thelypteridaceae. Any one of these could be

considered the culmination of a life study.

Mention must be made of the enormous help given to the field botanist by

publications of the vegetation of specialised localities. The Ferns of Mt. Kinabalu

(1934) by Christensen and Holttum in which many species were described, is one

of these. I speak from experience, having climbed this exciting mountain, for the

abundant fern flora, although having close affinities with the Malay Peninsula,

has interesting endemics and a temperate element from the southern hemisphere.

So much of this could be appreciated by having read this publication first.

Although much more is now known of the ferns there, this paper remains important,

not only in the enumeration of the species, but also for its field notes.

_ Dr. Holttum’s energy seems unabated and he still travels widely, not only

in the pursuit of ferns and to receive well deserved honours, but to visit his

daughter and grandchildren in Australia, which he and his wife have just done.

So we wish them well with many happy years ahead.

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Richard Eric Holttum, Croix de Guerre

A Note

H. M. BurKILL

Royal Botanic Gardens, Kew

Holttum’s autobiographical note in Flora Malesiana Bulletin 28 (1975): 2477-

500 contains an omission, in part at least self-effacing, but I am sure its honourable

nature is of sufficient interest to bring it to the attention of his friends and

colleagues. He refers very briefly to his service in the Friends’ Ambulance Unit

during the Great War of 1914-18. He went up to St. John’s College, Cambridge, in

October 1914 and took Class I Honours in the Natural Sciences Tripos, part 1, in

June, 1916. A normal university curriculum would lead on immediately to the

completion of the degree course in the following year. But conditions were not

normal and instead he broke the university timetable to join the Friends’ Ambulance

Unit for service in France, his university career being completed only after peace

had been restored. His service records, preserved in the Library of the Religious

Society of Friends in London run: date of joining 12/7/1916; training at Jordans

12/7-1/8/1916; ward orderly, St. George’s Hospital, London, 3/8-16/12/1916; in

France: posted to Headquarters Instructional School 2/7/1917; cook Section

Sanitaire Anglaise 19 27/1/1917; steward, SS.A. 19 25/9/1918; left unit

7/1/1919. This service with the French Army was recognised by the French

Government by decoration with La Croix de Guerre, the citation for which reads:

ECKERSLEY, STANLEY. B.; HEALD, ALFRED; HOLTTUM, R. ERIC;

JONES, JAMES; WILSON, ANTHONY L.; WILMOT, HERBERT R.;

WRIGHT, HERBERT

Ordre du Régiment 20 Janvier 1919

‘Convoyeurs a la Section Sanitaire Anglaise 19, ont fait preuve en toutes

circonstances sous le feu de l’ennemi de courage et de dévouement aux

blessés aux cours des opérations auxquelles cette section a pris part. (Flandres,

1917; Somme — Mars-Avril, 1918; Marne — Mai, Juin et Juillet, 1918;

Champagne — Octobre, Novembre, 1918.)’

[as in H. Wright: Two Years with the French Army: Section Sanitaire Anglaise 19,

Pelican Press, London, 1919, p. 117. Also in Meaburn Tatham & James E. Miles

(Editors): The Friends’ Ambulance Unit, 1914-1919, a record, Swarthmore Press,

London, 1920, p. 215.]

The bibliographic list in Flora Malesiana Bulletin 28 gives 1921 as the year of

his earliest publication. In fact, the period of his service in France in the Friends’

Ambulance Unit was the scene for his first published article: Social Life of the

Convoy, pp. 91-95 in Herbert Wright’s book cited above, predating all others by

two years. It is an interesting note on how he and his colleagues lived in France,

and makes light of time that must have been to a very large extent one of tedium,

squalor, fear and fortitude. The article shows an amateur theatrical group called

‘The Duds’, but unfortunately he can not be recognised amongst them.

_ _ Iam grateful to Mr. Malcolm Thomas of the Society of Friends for his help

in finding this information.

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Dr. R. E. Holttum — Eightieth Celebration

A. C. JERMY

British Museum (Natural History)

London

On 30 July 1975, some 75 friends and colleagues gathered in the Jodrell

Laboratory, Royal Botanic Gardens, Kew, England, at the kind invitation of the

Director, Professor J. H. Heslop-Harisson, to celebrate the eightieth birthday of

Dr. Richard Eric Holttum.

The occasion was organised by a group representing the Linnean Society of

London (from which Dr, Holttum was presented with the Linnean Medal in

1964), the British Pteridological Society (of which he was a Past President) and

the Royal Botanic Gardens, Kew, at which Holttum spent most of his retired, but

hard-working, hours.

After sherry in the Staff Common Room guests assembled in the Lecture

Theatre to applaud a number of presentations to Eric Holttum made by

distinguished guests on behalf of well-wishers both in England and abroad.

Mr. P. M. Brenan, Deputy Director of R. B. G. Kew, representing the Director

who was unfortunately abroad, presided over the proceedings. He welcomed the

guests and presented Dr. Holttum, on behalf of the Director and staff of Kew,

with a picture of Holttumara ‘Cochineal’ painted by Rosemary Lowe from

material specially sent from Singapore.

Professor Irene Manton, President of the Linnean Society of London, then

presented Dr. Holttum with a portrait of himseif drawn by Mrs. Julia Pannett and

subscribed to by friends and colleagues in the United Kingdom, Singapore, New

Guinea, New Zealand, Holland, Denmark, Switzerland, United States of America,

Jamaica, South Africa and India. Professor Manton went on to present him with

a special issue of the Fern Gazette, the journal of the British Pteridological Society,

published on that day in Holttum’s honour. It was then the turn of Dr. G. Seiden-

faden, formerly Danish Ambassador to Thailand and a well-known orchidologist,

who rose to read a statement of good wishes from the President, Mr. Quek Kiah

Huat, and Members of the Orchid Society of South-East Asia, and presented

Holttum with a gold medal struck by the Society to commemorate his eightieth

birthday. Telegrams and letters of good wishes were then read from Mr. A. G.

Alphonso, Deputy Commissioner i/c Singapore Botanic Gardens; the Singapore

Branch of the Malayan Nature Society; Mr. John Ede, President of the Singapore

Gardening Society; and Dr. R. M. Tryon, President of the American Fern Society.

At this point, Mrs. Jean Brenan, presented Mrs. Ursula Holttum with a

bouquet of flowers, whereupon Professor C.G.J.J. van Steenis, formerly Director

of the Rijksherbarium, Leiden, on behalf of colleagues at Leiden and the Flora

Malesiana Foundation, presented Holttum with a live young plant of Platycerium

holttumii Hennipm. and Joncheere.

The guests had a pleasant walk through the Gardens to the Restaurant where

they enjoyed a dinner given in Dr. Holttum’s honour. The guest speakers,

Mr. Henry Schollick, President of the British Pteridological Society and Professor

C.G.J.J. van Steenis gave short but amusing speeches, befitting the informality

8 Gardens’ Bulletin, Singapore — XXX (1977)

of this delightful and memorable occasion, Eric Holttum, dressed in a colourful

Singapore shirt, replied both to the presentation before dinner and to the saluta-

tions of the after-dinner speeches.

Even the English weather celebrated the occasion: a screen temperature of

30°C was recorded that day at Kew.

Guests at

Professor Holttum’s Birthday Celebrations

30 July 1975

Dr C. D. Adams

Mr G. Black

Dr D. F. Blaxell

Mr & Mrs J. P. M. Brenan

Mr F. H. Brightman

Mr H. J. Bruty

Mr & Mrs J. A. Crabbe

Dr & Mrs J. Croxall

Dr D. F. Cutler

Mr J. W. Dyce

Mr & Mrs P. J. Edwards

Mr L. Forman

Dr & Mrs G. W. Gillett

Dr A. D. Greenwood

Miss Mary Gregory

Mr & Mrs J. W. Grimes

Mr E. Hennipman

Dr & Mrs G. Herklots

Miss D. Holttum

Dr C. Hubbard

Dr M. Jacobs

Dr Frances Jarrett

Mr & Mrs A. C. Jermy

Mr G. J. de Joncheere

Mrs R. Lowe

Dr J. D. Lovis

Mrs MacDougal

Prof. Irene Manton

Mr P. McKenzie Black

Mr W. Marais

Dr & Mrs R. Melville

Dr & Mrs Panigrahi

Mrs J. Pannett

Mrs A. P. Passey

Miss T. K, Power

Dr & Mrs D. A. Reid

Prof. & Mrs P. W. Richards

Dr & Mrs N. Robson

Mr H. Schollick

Dr G. Seidenfaden

Mrs P. Simmonds

Dr Anne Sleep

Dr & Mrs W. T. Stearn

Prof. & Mrs C. G. G. J. van Steenis

Mr P. Taylor

Mr P. Temple

Dr & Mrs B. A. Thomas

Dr J. Vaughan

Dr & Mrs T. G. Walker

Mr P. J. Wanstall

Mr S. L. Williams

Dr & Mrs T. C. Whitmore

Mr & Mrs J. S. Womersley

Mr & Mrs J. Woodhams

Holttum’s contribution to horticulture in the

Malaysia-Singapore region

K. C. CHEANG* and A. G. ALPHONSO**

Horticulture both in Malaysiat and Singapore has taken on a very significant

role to-day. With rapid urbanisation the importance of planting trees and garden

plants whether for beautification or for other aesthetic reasons, has become not

only a very important undertaking, but a way of life with the peoples of the

region. Many, both amateurs and professionals, who are involved with gardening

and other horticultural pursuits in this region, make constant references to two

famous publications — “‘Gardening in the lowlands of Malaya,” and ‘Flora of

Malaya, Vol I — Orchids.’’ The author of these publications is none other than

Professor R.E. Holttum. To the amateur who tends his house garden, to the

professional gardener who runs a commercial orchid nursery or ornamental

garden, to the horticulturists of horticultural establishments, Holttum is a house-

hold name, That this should be so is not surprising as Professor Holttum has

done more for the promotion of horticulture in this region than anyone else.

Holttum first came out to Singapore in 1922, at a time when there were a few

Chinese commercial gardens growing some orchids and a few flowering and

foliage plants. Most of these gardens were in Orchard Road, Thomson Road and

the Newton and Bukit Timah districts of Singapore, and a few in Penang. Interest

in gardening among the people at this time was keen, but there was no proper

guidance by an authority on the subject. Holttum befriended the Chinese gardeners,

and spent considerable time discussing with them horticultural matters, and at the

same time observing the traditional Chinese methods of growing plants, one of

which was the use of burnt clay as a potting medium. There was here an inter-

change of knowledge between the botanist and the practical gardener. A few of

these gardeners who are alive to-day and whose children now carry on the

management of their nurseries, speak very highly of Holttum, both as a horti-

culturist and as a kind man. Holttum’s knowledge of horticulture was not obtained

from books alone. His was a practical approach to the subject as a result of long

experience in the growing of tropical plants in the region. He did not as a rule,

resort wholly to scientific methods, or put forward scientific suggestions. In the

pursuit of gardening, he always made it easy for the ordinary man.

In 1928 Holttum together with John Laycock and Emile Galistan, two very

keen orchid growers founded the Malayan Orchid Society. At this time there was

a great interest shown by the public in the growing of orchids, and the three must

have realised that such a Society would play an important role in the promotion

of orchid growing in the country. A few years later the Singapore Gardening

Society was formed, and Holttum was mainly responsible for its founding.

Meetings were regularly held for members, and the two Societies under the expert

guidance and advice of Holttum flourished, and played a great role in the pro-

motion of orchid growing and gardening in the region. Annual shows were held,

the first being an orchid show organised by the Malayan Orchid Society (now

the Orchid Society of S.E. Asia) on the 27th and 28th of March 1931 at the

* lately Head, Botanic Gardens, Penang.

** Botanic Gardens, Singapore.

ft here denotes Malaya, Sarawak and Sabah.

10 Gardens’ Bulletin, Singapore — XXX (1977)

YMCA Building in Stamford Road, Singapore. Holttum was mainly responsible

for putting up the exhibits of the many species of orchids and the newly raised

hybrids of the Singapore Botanic Gardens. It was Holttum who for the first time

at this exhibition demonstrated to the Singaporeans and Malaysians alike, the

Knudson method of asymbiotic flask culture of growing orchids, showing flasks

of orchid seeds at the germination stage, others with seedlings at different stages

of growth, and finally established seedlings planted on pieces of wood. This

technique in orchid propagation, which Holttum started in 1928 was the turning

point in the growing of orchids in the Malaysia-Singapore region, and in fact the

whole of S. E. Asia. Seeds could now be successfully germinated and as such, was

a tremendous asset in the hybridisation and breeding of orchids in the region.

Holttum himself produced a large number of hybrids. The orchid shows continued

annually until 1934, when the first flower show including both ornamental and

orchid plants was held at the New World Stadium from the 6th-8th April, and

jointly organised by members of both the Malayan Orchid Society and the

Singapore Gardening Society. These shows were to continue annually to the present

time. Holttum must have known that horticultural shows are of immense value

to both professional and amateur growers, for they serve to raise the standard of

horticulture in the country by giving the growers a chance to compare their

exhibits and to show what can be achieved. They also help to encourage the non-

gardener to take an interest in gardening. Holttum retired as Director of the

Singapore Botanic Gardens in 1949 and was appointed Professor of Botany at the

newly formed University of Malaya. He held this post until 1954 when he returned

to England. For over 25 years he had devoted his time energetically to the

activities of the Malayan Orchid Society and the Singapore Gardening Society.

Although the flora of the Malaysia-Singapore area is one of tropical rain

forest, it is surprising to note that there are few indigenous plants of horticultural

value. Native ornamental plants and flowering trees are few compared to the

abundance of species in the flora of the area. The flowering trees and ornamental

plants grown in one’s garden, in parks and open spaces, by roads and streets and

even in the Botanic Gardens, have mostly been introduced especially from Central

and South America, and other parts of Tropical Asia. The introduction of new

plants into Malaysia and Singapore has therefore played an important role in the

horticultural beautification of the area. Although private individuals, business

houses and other establishments have to some extent been responsible in the

introduction of plants into this region, it was Holttum, during his 27 years of office

as Assistant Director and Director of the Gardens Department, Straits Settlements

(Singapore, Malacca and Penang), who was mainly responsible for the countless

numbers of flowering trees and shrubs introduced to the two Botanic Gardens in

Singapore and Penang from tropical regions of North and South America, Africa,

Asia and Australia. Many of these plants to-day are extensively grown in parks,

by roadsides, and private gardens and commercial nurseries throughout Malaysia

and Singapore.

Holttum is a widely travelled person, having attended many international

horticultural conferences. His lectures and papers presented at these conferences

included topics on various aspects of horticulture in this region, particularly relating

to orchids and ferns,

Some thirty odd plants, both species and hybrids, have been named after

Holttum. Perhaps the trigeneric orchid hybrid Holttumara cochineal (Arachnis x

Renanthera x Vanda) is the most outstanding as far as horticultural plants are

concerned. A painting of this hybrid was made by a staff member of the Royal

Botanic Gardens, Kew, and presented to Holttum on his 80th birthday. Some

years back the authors on a plant collecting trip to Kaki Bukit in Perlis, Peninsula ~

Holttum’s contribution 11

Malaysia, discovered a staghorn fern which was appropriately named Platycerium

holttumii, for no one in this region has done more towards popularising the

growing of ferns as a decorative house plant than Holttum.

In 1940 for the first time, three local officers were recruited by Holttum for

horticultural training. K. C. Cheang was assigned to the Botanic Gardens in Penang,

A. G. Alphonso to the Botanic Gardens in Singapore and N. V. Lange to the Parks

Department in Singapore and subsequently to the Parks Department in Ipoh,

Perak. Holttum must have realised at this time that the future of horticulture in

the region rested with qualified local horticulturists. Up to such time horticulturists

were expatriate officers recruited from the Royal Botanic Gardens, Kew. Apart

from the training received directly under Holttum, the officers were sent to the

College of Agriculture in Serdang, Malaysia, and later K. C. Cheang proceeded to

Massey University, New Zealand, N. V. Lange to the Institute of Parks. Administra-

tion, England, and A. G. Alphonso to the Royal Botanic Gardens, Kew. On their

return they were appointed Curators, and were subsequently to head their

respective departments. It was during the period of training under Holttum that

the authors had come to appreciate him as a truly outstanding horticulturist. His

immense knowledge of local plants and his long standing experience in practical

gardening, won him the admiration of all who came to know and learn under him.

Above all, he was a very kind person, and was always willing to help his fellow-

men. We are proud and honoured to have served under Holttum.

Holttum’s contribution in the field of horticulture has won him many honours

and he has been the recipient of a number of awards from horticultural societies

in the U.S.A., Great Britain and Singapore. In recognition of his work on orchids

in Malaysia and Singapore, Holttum, at the 4th World Orchid Conference held in

Singapore in 1963, was awarded the Gold Medal of the Malayan Orchid Society.

In addition, the Orchid Society of S.E. Asia awards annually a Gold Medal known

as the Eric Holttum Gold Medal to the originator of the best locally produced

hybrid. On the occasion of his 80th birthday the Orchid Society of S. E. Asia,

at a ceremony at the Royal Botanic Gardens, Kew, presented him with a Gold

Medal suitably inscribed, and through the generosity of the Singapore Gardening

Society, the Botany Department of the University of Singapore presents annually

a Holttum Silver Medal to an honours student who has performed well in his

examinations.

Perhaps Holttum’s biggest contribution to horticulture in the Malaysia-

Singapore region lies in his published works. He is a prolific writer, and his many

articles on gardening in the Malayan Agri-Horticultural Association magazine for

well over 20 years, are not only instructive but of great benefit to both the amateur

and professional gardener. The Malayan Orchid Review, the official organ of the

Malayan Orchid Society was first published in 1931. Holttum was the main contri-

butor. He was the expert, and his many and varied topics in the Orchid Review

have helped orchid growing in this region. By far, his best works in horticulture

and by which the Malaysian and Singaporean will best remember him, are his

“Gardening in the Lowlands of Malaya” and his ‘Flora of Malaya, Vol I —

Orchids”, mentioned earlier. The former is an excellent book on_ practical

gardening in this region. It is as popular to-day as when it was first published in

1953. The latter, the most outstanding publication on orchids of this area, has

been a constant companion of the orchid growers. Holttum is also the author

of many other publications on local plants of horticultural interest. His ‘‘Flora of

Malaya, Vol Il — Ferns” and articles on Bamboos, Gingers, Maranta, Grasses

and others, published in the Gardens’ Bulletin, although taxonomic or

botanical in nature, have in many ways helped the horticulturist in his work,

especially in the identification of plants he deals with.

12 Gardens’ Bulletin, Singapore — XXX (1977)

We in this region owe a debt of gratitude to Holttum for his outstanding

contribution to horticulture. To-day, in both Malaysia and Singapore, there is a

thriving orchid industry. The value of present exports of cut orchid blooms from

both these countries has reached nearly ten million local dollars. Orchid nurseries

have come up in large numbers, and there is no doubt that the orchid industry will

continue to expand. This situation would not have come about had it not been

for the pioneer work of Holttum in orchid growing. His introduction of asymbiotic

flask culture of orchids n 1928, the breeding and hybridisation of orchids which he

started, and the encouragement he gave to professionals and amateurs in the field

of horticulture have borne fruit.

The role he played as adviser to the Parks Department and in the planting

of roadside trees in the early years, his able administration of the two Botanic

Gardens of Singapore and Penang, and the valuable knowledge he has imparted

to the students as Professor of Botany at the University of Malaya, bear testimony

to his foresight and vast horticultural knowledge. And above all, his many horti-

cultural publications will be constant reminders to us all of his dedicated service

in this region.

This number of the Gardens’ Bulletin is specially brought out to commemorate

the 80th birthday of Professor Holttum, and in writing this article the authors join

the many Malaysians and Singaporeans, the orchid growers, the amateur and

professional gardeners, and the members of the Orchid Society of S. E. Asia and

the Singapore Gardening Society, in wishing him good health and happiness in

the years ahead.

Staghorn Ferns Today and Tomorrow

BARBARA JOE HOSHIZAKI

Botanical Gardens-Herbarium

University of California

Los Angeles

The unique appearance of staghorn ferns (Platycerium) has attracted the

attention of enthusiasts for well over a hundred years. Modern transportation and

communication have brought all 18 species into cultivation. So great is interest in

this fascinating fern group that it seems appropriate to note recent horticultural and

botanical works that have solved many puzzles and to look ahead to problems yet

to be answered.

HorRTICULTURAL CONTRIBUTIONS. Horticultural interest in staghorn ferns has

greatly benefited botanical work. The perseverence and tenacity of Platycerium

fanciers who seek out new plants, grow them, and develop new varieties is amazing.

Through their efforts I have been able to study every one of the known Platycerium

species as live plants. The opportunity to watch plants grow from spore to maturity

contributed greatly to the botanical knowledge of the genus.

Platycerium enthusiasts in United States cultivated a new species from

Malaysia for several years before it was recognized and named by botanists

(Joncheere and Hennipman 1970). This handsome plant, now called Platycerium

holttumii, was well known enough by amateurs to have been photographed for two

popular books prior to being formally named (Rie et all 1957, Franks 1969). A

second new species, the P. grande of the Philippines, has been recognized in the

last few years (Joncheere and Hennipman 1970), another exciting event to

platycerium fanciers! These relatively recent discoveries kindle the imagination of

collectors who hope that different species may still be discovered. Collectors continue

their quest for new and unusual platyceriums. Shipments of P. wallichii have

reappeared in the United States, and these plants do show some frond variation.

The significance of these variations has yet to be investigated. These plants were

reported to have been collected along the India-Burma border. Native plants of

P. stemaria have an interesting range of frond variation which may interest botanist

as well (Joe 1964). But besides seeking variations among wild plants, horticulturist

are developing new garden varieties. The multititude of new garden varieties is due

to the increased number of people raising plants from spores. A hybrid platycerium,

P. mentelosii (P. stemaria x superbum) has been developed and other hybrid

combinations will most likely follow (Hoshizaki 1975). Growers tell us that

platyceriums are quite plastic in their leaf form. Slight changes in the environment

may cause base fronds to become partly foliaceous, normally entire fronds to

become forked and so forth.

Staghorn fern propagation by meristem or tissue culture is being actively

investigated by commercial nurseries. They claim that plants may grow twice as

fast through meristem culture as by spores. Meristem plants tend to produce

mutations more frequently than spore grown ones and this interest horticulturist

and geneticist.

__ The Malaysian plant, Platycerium ridleyi, thought extinct, has been found and

introduced into United States cultivation. It grows very nicely in southern California

14 Gardens’ Bulletin, Singapore — XXX (1977)

greenhouses if adequately protected from slugs and snails. Spores from these

introduced plants have produced mature plants, thus insuring the species’ survival

and reducing the need to collect these rare plants from native habitats, a happy

circumstance for conservationists.

Much more information is needed on the cultural conditions for Platycerium

wallichii of Southeast Asia, and P, madagascariense and P. quadridichotomum both

of Malagasy. These species, being difficult to cultivate, create demands for replace-

ment, a situation which is not encouraging for conservationists. Hopefully more

work along the lines of Boyer will help in the successful growth and propagation of

these plants. Boyer closely examined the ecology, physiology, and mineral needs

of two African species, P. angolense and P. stemaria (Boyer 1964).

The type of foregoing activities and results generated by horticulturists hold

promise of supporting and even giving direction to scientific work and increasing

our understanding of the genus. To help science, horticulturist should be encouraged

to maintain reliable records as to where native plants were collected, or if new

plants were developed in cultivation, the names of the parent or parents involved.

Conditions which may have caused unusual growth patterns should also be recorded.

Horticulturists on the other hand should protect native plants and prevent their

extinction from indiscriminate collecting.

BOTANICAL CONTRIBUTIONS. The name changes among platycerium species

confuse and frustrate growers and most botanists. If it is any comfort to know,

botanist specializing in nomenclature have for years been trying to unravel the

technical complexities in determining the proper name for certain species. A series

of papers and rebuttals dealing with this problem has been published in the last

few years by G. J. Joncheere and the late C. V. Morton (Morton 1964, 1970,

Joncheere 1967, 1974). The main issues first center around whether Platycerium

vassei as conceived by botanist, not horticulturist should be called P. alcicorne and

second, whether P. angolense should be called P. elephantosis, A few botanist are

following this intricate and complicated problem and their views will be forth-

coming. Morton’s argument, that the correct spelling of P. stemaria is not P.

stemmaria, seems to be upheld. The acceptance of P. wandae as the legitimate name

over P. wilhelminae-reginae- seems without complication (Joncheere 1968). How-

ever, among plants introduced from New Guinea and now growing at Longwood

Garden, Pennsylvania are two kinds of P. wandae, One produces long fertile fronds

fitting the original description of P. wilhelminae-reginae and the other produces

short fronds closely fitting the description of P. wandae. In all likelihood both

plants are one species. Short and long fertile frond forms have also been noted in

P. coronarium, However, further study and field observations on long and short

frond plants might be of interest.

It is indeed unfortunate that the name Platycerium grande must now be

applied to the Philippine plant instead of the Australian plant. The Australian

plant must now be called P. superbum in accord with Joncheere and Hennipman

(1970). I examined the rhizome scales of the Philippine plant and they differ from

the Australian plant, though both are closely related. Rhizome scales of the

Philippine plant are very similar to P. holttumii except for having slightly fewer

cells in the marginal hairs (highest number of cells in the longest hairs were mostly

7 in P. grande and 8 in P. holttumii). The scale morphology is consistent with other

morphological data which relates P. grande, P. superbum, and P. holttumii

(Hoshizaki 1970, 1972).

The relationships of the Australian-Javan species (Platycerium bifurcatum,

P. veitchii, P. willinckii, and P. hillii) and their distribution still needs to be clarified

with more data. Some of the features separating these supposedly different species

are not stable from observations on plants in cultivation. Mr. Ernest Todd of New

South Wales, Australia has been investigating the distribution of the Australian-

Javan species. In a personal communication Mr, Todd reports that the collections of

Staghorn Ferns 15

P. bifurcatum by Bambler and others in New Guinea were probably taken from

cultivated or escaped plants most likely introduced by German missionaries in the

early part of the century. Distributions of some of the Malayan-Asiatic species

also need further clarification.

Platycerium coronarium and P. ridleyi are very closely related species, yet

there is a gap between these two species and their closest relatives. It doesn’t seem

too far-fetched to speculate that some yet undiscovered species may be found in

the Malaysian area that bridges this gap and others as well.

With the relatively wide-spread use of the scanning microscope studies should

be encouraged on the ontogeny of stellate hairs in this genus and its proposed

relatives (Pyrrosia and Drymoglossum). It might be worthwhile to also study and

compare these genera as to sporangial structure. Within the platyceriums are species

with complete to incomplete annular rings, laterally and apically located lip cells,

and stalk structure variations.

Certainly there are many topics I have not touched upon that should be

considered in future studies. However, I want to stress that botanist and horti-

culturist have much to look forward to in knowing more about these unusual

plants.

Literature cited

Boyer, Yvette née Menoux. 1964. Contribution a l’estude de l’ecophysiologie de

deux fougeres epiphytes: Platycerium stemaria (Beauv.) Desv. et P. angolense

Welw. Théses présentées a la Faculté des Sciences de l'Universite de Paris.

Série A, No. 4191, No. d’ordre: 5042,

Franks, Wendy. 1969. Platycerium Fern Facts. Privately published. 119 p.

Hoshizaki, Barbara Joe. 1970. The Rhizome Scales of Platycerium. Am. Fern J.

60: 144-160.

Hoshizaki, Barbara Joe. 1972. Morphology and Phylogeny of Platycerium Species.

Biotropica 4: 93-117.

Hoshizaki, Barbara Joe. 1975. A Staghorn Fern (Platycerium) Hybrid. Am.

Fern J. 65: 99-101.

Joe, Barbara. 1964. A Review of the Species of Platycerium (Polypodiaceae).

Baileya 12: 69-124.

Joncheere, G. J. de. 1967. Notes on Platycerium Desv. I. Nomenclature and

typification of the genus and species in Desvaux’s original publication of 1827.

Blumea 15: 441-451.

Joncheere, G. J. de. 1968. Notes on Platycerium Desv. II. P. wilhelminae-reginae

v.A.v.R. reduced to P. wandae Rac. Blumea 16: 109-114.

oe G. J. de. 1974, Nomenclatural notes on Platycerium (Filices) Blumea

2: 53-55.

Joncheere, G. J. de and E, Hennipman. 1970. Two new species of Platycerium and

the identification of P. grande (Fee) Presl. Br. Fern Gazette 10: 113-118.

en. es V. 1964. The Nomenclature of a Madagascarian Platycerium. Baileya

12: 36-38.

perion. C. V. 1970. A further note on the type of Platycerium alcicorne. Am. Fern

. 60: 7-12.

Riie, Edgar Aubert de la, Francois Bourliére, Jean Paul Harroy, 1957. The Tropics.

Knopf. 208p.

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A new species of Nastus Nees (Gramineae) from Sumba

SOEJATMI SOENARKO

Herbarium Bogoriense

Indonesia

Summary

An illustrated description of Nastus reholttumianus S. Soenarko is presented and this new

species is compared with N. rudimentifer Holtt. and N. obtosus Holtt.

When Holttum (1956) transferred the Malesian species of Oreiostachys

Gamble to Nastus Nees, three species of Nastus were known from New Guinea. In

1967 he described four more species and Bor (1972) added one to these; thus

there are now eight species endemic to New Guinea. Outside New Guinea there

is One species; it occurs in Java.

In most of the Malesian Nastus the spikelets are more or less cylindrical, thus

the fourth and the fifth glumes, lemma and palea are rounded at the back (except

those in N. schlechteri (Pilger) Holtt., in which they are more or less 1-keeled),

with usually prominent nerves; moreover the palea is bifid. A plant collected by

Mr. Iboet from the island of Sumba (Indonesia) has characteristically small and

laterally compressed spikelets, lightly 1-keeled glumes, lemma and palea, with

prominent middle nerve and several (usually 6) faintly lateral nerves, and

acuminate palea, These structures, together with the leaf size (up to 8 cm long

and 14 mm wide) differentiate the Sumba plant from the other Malesian Nastus,

and it is regarded here as a new species, N. reholttumianus. The near relatives of

this new Nastus are N. rudimentifer Holtt. and N. obtusus Holtt. which have similar

panicle with spreading branches and short-tipped lower glumes. However, this new

species differs in several respects from the two latter; their characters are compared

in the table below:

characters reholttumianus rudimentifer obtusus

leaf-blade of flowering | 4.5-8 cm long (8) 10-12 cm long | 12-15 cm long

branch 7-14 mm wide 15-27 mm wide 12-15 mm wide

spikelet + laterally + cylindrical + cylindrical

compressed

5-7 mm long 12 mm long 13-20 mm long

rachilla extension absent present, sometimes | usually absent

absent

lemma 6 mm long, 10 mm long, 14-17 mm long, with

glabrous glabrous appressed hairs on the

back

palea

acuminate bifid bifid

Nastus reholttumianus S. Soenarko, sp. nov., N. rudimentifer Holtt. affinis

sed spiculis 5-7 mm longis, latere compressis, lemmate 6 mm longo, lamina 8

longa differt. Typus: Sumba, Iboet 443 (holotypus, BO).

Fig.

Gardens’ Bulletin, Singapore — XXX (1977)

1. Nastus reholttumianus sp. nov. J, flowering branches; 2, spikelet; 3-7, glumes I —

V respectively; 8, lemma; 9, palea; 10, ovary; 11, lodicule; 12, spikelet after removing

the glumes and lemma.

Nastus 19

Culm not seen, upper part bearing flowering branches 3.5 mm diam., either

solid or hollow, with annular woody ring below each node; branches glabrous,

3-8 in each node. Culm-sheath not seen. Flowering branch 11-22 cm long; leaf-

blades 4.5-8 cm long, 7-14 mm wide, oblong-lanceolate, glabrous, acuminate,

rounded at the base; leaf-sheath glabrous; ligule less than 1 mm long; auricle up

to 1 mm long, fringed (fringe 1-2 mm long). Inflorescence open panicle with

spreading branches, main axis and branches with minute appressed hairs. Spikelet

more or less laterally compressed, up to 7 mm long, glabrous; glume I: 1.5 mm

long, narrowly ovate, 1-keeled on the back, with mucronate apex; glume II: 2.5 mm

ovate, slightly 1-keeled on the back, mucronate; glume III: 3.5 mm long, slightly

1-keeled on the back, ovate, faintly 6-nerved with conspicuous median nerve (and

so are the median nerves of glumes I and II); glume IV: 5.5 mm long, ovate to

ovate-oblong, boat-shaped, acuminate, 3 mm wide, faintly 7-nerved; glume V:

more or less similar to glume IV, but slightly longer; lemma and palea more or

less equal, 6 mm long, ovate-oblong, acuminate, boat-shaped to slightly com-

pressed; lodicule 1 mm long with fringes; ovary stalked with conical fleshy apex,

glabrous, the stalk often elongates up to 2 mm long; rachilla extension absent;

stamens 6, anthers 2.5 mm long. (Fig. 1).

Sumba. Mangiliwari near Maomaru, 9. V. 1925, Iboet 443 (BO, holo).

The author dedicates this new species to Prof. R. E. Holttum who has

contributed so much to the knowledge of S. E. Asian bamboos, especially of

Malaya and New Guinea, and has encouraged the author in studying Malesian

bamboos. Thanks are due to Sdr. Damhuri (Herbarium Bogoriense) for preparing

the figures.

References

Bor, N. L. (1972). A new species of Nastus from New Guinea. Ost. Bot. Z.

120: 87-91.

Holttum, R. E. (1956). The bamboo-genera Nastus and Chloothamnus. Kew. Bull.

10: 591-594.

(1967). The bamboos of New Guinea. Kew Bull. 21: 263-292.

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On Iridescent Plants

Davi W. LEE

Botany Department

University of Malaya, Kuala Lumpur

Abstract

Plants with leaves of iridescent blue and green colour are common in the deep shade

of Malaysian rain-forests. Simple anatomical observations have revealed that the green

iridescence is due to the refraction of diffuse light onto specially-oriented chloroplasts by

lense-shaped cells. Blue iridescence colour is due to the operation of thin film interference

filters in or on the epidermis. The advantage of such a filter in forest shade plants is the

effective absorption of red wavelengths of light at the expense of the reflection of less

important blue wavelengths. This report documents iridescence in the leaves of many

unstudied taxa (mainly pteridophytes), common in Malaysian rain-forests.

Life in a tropical rain-forest is not quite as easy for green plants as popular

accounts would indicate. First by, the forest soils are generally rather sterile.

Secondly, plants are under intense pressure of predation by insects and other

animals, Finally, the light conditions are frequently less than optimum—from very

bright conditions in the canopy to deep shade on the rain-forest floor. In this

discussion we are concerned with the deficiency of light in the shade at the bottom

of the rain-forest; in this environment can be seen a number of plants whose leaves

display brilliant blue and green colours. These colours can be described as

iridescent because of their intensity and metallic quality. The association of these

iridescent plants with the shady and moist environments has compelled) many

naturalists to reason that the colours must be associated with adaptations which

help the plants to survive in the dark habitats. The rain-forest floor is one of the

darkest environments in which plants grow. Not only is the total light quantity

a small fraction of that above (less than one percent) but also the light quality is

diminished; more of the wavelengths vital to photosynthesis, with the exception of

some red light, have been absorbed by the foliage overhead (Evans, 1966; Federer

and Tanner, 1966; Bjorkman, 1972). The purpose of this discussion is to explore

how the iridescence of some Malaysian plants is associated with adaptations to

increase the absorbance of diffuse and weak light for maximum photosynthetic

efficiency.

LEAF ANATOMY AND FUNCTION

Leaves possess many structures which are significant in the efficient absorption

of light in shady environments (see Ray, 1972; and Esau, 1965). Figure 1 sum-

marizes these structures.

The Cuticle. The function of the eit is two-fold (see Martin and J uniper

1970 for further details). First, it repels water and prevents the rapid dessication

of the leaf. Secondly, it can act as a selective filter in reflecting harmful ultraviolet

light, and allowing the absorption of visible light essential for photosynthesis.

The Epidermis. This superficial cell layer, particularly on the upper leaf

surface, may possess special adaptations for efficient light absorption. It may have

bumps which remain dry when the lower areas are covered by rainwater; these

allow more efficient light absorption in wet conditions (Haberlandt, 1914).

22 Gardens’ Bulletin, Singapore — XXX (1977)

Epidermal cells may also have regularly convexly curving upper surfaces; these

focus diffuse sunlight onto specially orientated chloroplasts. Finally, the epidermal

cells of extremely shade-tolerant plants (especially pteridophytes) may contain

ee which can more directly receive light before it is scattered by the

cell walls.

Photosynthetic Parenchyma. Both the palisade and spongy mesophyll layers

may have special adaptations for low light environments. They may be specially

oriented beneath the lens-like epidermal cells. In shade-tolerant plants these cells

would have fewer and larger chloroplasts. The protein/chlorophyll ratio, which

reflects the relative plant investment in light reception versus biosynthetic

apparatus, would be low. Finally, the cells may be closely packed together to

minimize the reflection (and loss) of light between the cells.

Stomata. In low light environments the stomata will not be a limiting factor,

and we might expect various arrangements on the upper and, especially, lower

leaf surfaces.

Leaf Size and Shape. In low light environments leaves tend to be large and

thin, with a high surface/volume ratio. This ratio is increased even more in plants,

as ferns, with highly divided leaves.

Thus, the task in understanding the significance of iridescence in shade-tolerant

plants is to explain how some attributes of leaf structure confer an advantage on

the plants and, at the same time, physically account for the iridescent colour.

IRIDESCENCE

Many plants from diverse taxonomic groups, from humid rain-forests

throughout the tropics, possess iridescent leaves (Richards, 1952). Iridescent plants

can be observed in Malaysian rain-forests (see Table 1), and some are extremely

common. Selaginella willdenowii is a scandent pteridophyte frequently encountered

in rain-forests, plantations and belukar throughout Southeast Asia. Another

pteridophyte, Athyrium crenatoserratum, can be frequently observed in local rain-

forest, as is Phyllagathis rotundifolia (Melastomataceae). Blue-leaved sedges of

the genus Mapania commonly grow in Sarawak rain-forests, Others are less com-

mon. The two kinds of iridescence observed in these plants, green and blue, are

discussed separately below.

Green Iridescence. The first observations on green iridescence were made

on the cave moss, Schistostega, which grows in caves and other dark and wet

environments in Europe and North America, Richards (1932) reported that the

brilliant green colour encountered at particular angles was due to the functioning

of certain protonemal cells as refractive lenses, focussing the diffuse light onto the

specially oriented chloroplasts. Analysis of Malaysian green iridescent plants

indicates that the same principles operate (see Figure 2 and Plate 1). These plants

all share cells (epidermal or palisade) whose outer wails have a uniform convex

curve. The chloroplasts are always oriented at the distal end of these cells. When

we look at these plants the intense green colour can be observed only at an angle

perpendicular to the surface of the plant. At other angles the colour is dull green

or even whitish. This effect is due to the focusing of the diffuse light into a more

intense beam directly onto the chloroplasts. These organelles, with their parallel

stromal organization, absorb the light useful in photosynthesis and reflect the

unused green light directly out of the long cells. Thus when we look at these plants,

as at the entrance to Batu Caves near Kuala Lumpur or on rocks in especially shady

forest habitats, we see the green light reflected in an intense and narrow beam. At

other angles the light appearance of the leaves is due to reflection of whole light

off of the cell surface acute angles and also the scattering of light within the cells.

The birefringence of the cellulosic walls may also be a factor affecting the quality

of the light reflected at acute angles.

Iridescent Plants 23

CUTICLE

UPPER

i a EPIDERMIS

PALISADE

AND MESOPHYLL

PARENCHYMA

, Oe cone ed

AA e-\

J pel psn YT Ly eens VASCULAR

+ Oee BUNDLE

QO \

LOWER

say —— EPIDERMIS

Fig. 1. A diagram of the typical structure of a leaf.

BRILLIANT GREEN COLOUR

DULL GREEN COLOUR

WHITISH YELLOW COLOUR

LEAF SURFACE

INTENSE GREEN

LIGHT REFLECTED

[TENSE GREEN

DIFFUSE LIGHT FEIGHT REFLECTED

BACK SCATTERING

OF WHITE LIGHT

DIFFUSE LIGHTK

FROM EPIDERMAL |

CELLS

Fig. 2. The mechanism for the production of green light in iridescent green plants.

24 Gardens’ Bulletin, Singapore — XXX (1977)

Blue Iridescence. Men have been intrigued by the startling blue plants ever

since they were first encountered. In the Malaysian rain-forest Selaginella willdenowii

is the most striking example. In traditional languages the name ‘“‘peacock plant”

is frequently used to describe these pteridophytes (as the Malay “‘paku merak’’).

One aboriginal Senoi name for these plants translates roughly as “‘the hair on a

tiger’s rump” (Gerard Diffloth, personal communication). These plants have been

included in various medicinal remedies probably simply because of their colour.

Selaginella willdenowii has been used in preparations for aching back as well as

fever (Burkill, 1935). European scientists made frequent mention of the colour but

gave no explanation. Stahl (1896) studied §. willdenowii while working in the

famous Treub Laboratory at Bogor. He observed that the iridescent colour was

due to pigmentation granules in the cuticle, Fox and Wells (1971) and Lee and

Lowry (1975) recently published more adequate explanations for colour in

Selaginella willdenowii, likely also to be the mechanism of colour formation for the

other blue plants listed in Table 1. Stahl’s original observations can be discounted

for three reasons. There are no presently known pigments in pteridophytes with

the spectral properties to account for the blue colour. Secondly, in re-examining

the epidermis of these plants we observed no pigment granules. Thirdly, the blue

colour of the leaves disappears when they are covered with water. Thus, the colour

must be due to some physical optical effect of the leaf surface, and not a pigment.

There are three physical phenomena which can account for iridescent colour as

seen in insects (see Michelson, 1911; Fox, 1959; Neville and Caveney, 1969).

These could also be invoked to explain the blue leaf colour, Diffraction of light on

thin gratings disperses different colours at different angles to the object’s surface.

In S. willdenowii and the other plants, only blue colour can be observed. Tyndall

scattering (which explains blue sky colour) polarizes light; no polarizing effects

have yet been observed in any iridescent plants. Light interference, due to thin

films, can produce uniform colours over relatively wide angles of incidence, and is

the likely physical explanation of the phenomenon.

The principle of a thin film (or quarter wavelength) interference filter is not

familiar to most biologists, and requires some explanation. As a specific example,

if the conditions described in Figure 3 are met, optical interference will result in

the differential absorbance and reflection of different wavelengths of light. First,

the filter must have a slightly different refractive index (r) than the surrounding

medium, the air above or the cell below. Secondly, the filter must be exceedingly

thin, a fraction of the wavelength of visible light. Thus, in the filter described in

Figure 3, the thickness (142, ) is fraction of red visible light at 600 yp. If we trace

the path of blue light (at 405 ») through this filter, we see that light of this wave-

length will be reflected by the filter. The distance travelled from A and back to C

equals the wavelength of that light in the filter. Retarding the light by one wave-

length puts the light (visualized as a transverse wave) in phase, and the energy

at this wavelength moves out of the filter. However, light with four times the wave-

length of the filter thickness (or red light of 795 ») would be retarded half a wave-

length at the same point, and its energy would be nullified. Thus such a filter can

selectively enhance the reflection of certain wavelengths and allow the absorbance

of other wavelengths. Two common examples are the rainbow colours seen from

the thin oil films on rain puddles, and also the special anti-reflection coatings on

quality cameras and binoculars. For a more technical explanation see Vasicek

(1960).

To test the above hypothesis Lee and Lowry (1975) analyzed the reflectance

of light from both the iridescent and the older green leaves of S. willdenowii by

using a slightly modified spectrophotometer. The analysis gave a peak of reflectance

by the blue leaves at 405 p». They then constructed a filter model fitting this

observation, as in Fgure 3. They assumed a filter refractive index slightly in excess

of cell contents (of about r= 1.35), or about r= 1.4 as in Figure 3. In such a

filter a concellation of the reflection effects would be observed at about 530 p,

Iridescent Plants 25

/ : AIR (r=1.00)

FILTER (r=1-40)

CELL (r=1-35)

Fig. 3. Operation of a simple thin film interference filter.

1. REFLECTS BLUE LIGHT.

IN THE FILTER, AT AN EFFECTIVE WAVELENGTH OF 284, THE LIGHT IS

DELAYED 4 WAVELENGTH AT POINT B AND IS NULLIFIED— CANCELLED AS

OVERLAPPING TRANSVERSE WAVES WOULD SHOW. AT POINT C LIGHT IS

DELAYED A FULL WAVELENGTH AND IS REFLECTED OUT — WAVES COINCIDE.

AT THIS POINT THE REFLECTED LIGHT HAS A WAVELENGTH OF 4051

(1.4/1.0 x 284) AND IS BLUE.

2. NO EFFECT ON LIGHT AT 530 pu

AT 2} THE FILTER THICKNESS THE WAVES WOULD CANCEL.

3. ABSORBS RED LIGHT.

IN THE FILTER LIGHT AT A WAVELENGTH of 568, IS DELAYED + WAVE-

LENGTH AT C AND IS NULLIFIED— CANCELLED AS OVERLAPPING TRANS-

VERSE WAVES WOULD SHOW. THUS MORE OF THIS LIGHT PASSES THROUGH

THE FILTER AT B. HERE THE EFFECTIVE WAVELENGTH IN AIR IS 795 u

(568 x 1.4/1.0) AND IS RED.

with increased absorbance at higher wavelengths. In the difference spectrum

obtained by subtracting values of the green leaves from the blue leaves; a curve

similar to that predicted was observed (see Figure 4). Light above 700 » would

be reflected by all leaves (Gaussman and Allan, 1973; Gates et al., 1965) and

would obscure the filter effect at greater wavelengths. Thus, we have good physical

evidence that such filters operate in the blue plants.

The operation of such a filter has obvious adaptive significance for these

shade-tolerant plants. Blue light (of which there is little in the dim light of the

forest floor) is reflected, and the red light, which is more significant in the total

photosynthesis by these plants, is absorbed. Thus, these filters can operate as

“anti-reflection”? ccatings which increase the absorption of photosynthetically active

light on the shady forest floor.

Selaginella willdenowii plants possess additional adaptations to increase photo-

synthetic efficiency. Firstly, the epidermal cells contain single large chloroplasts at

the distal ends of the cells (see Plate 2). Thus, most of the photosynthesis occurs

26 Gardens’ Bulletin, Singapore — XXX (1977)

just below the leaf surface. Secondly, the epidermal cells have regularly convexly

curved surfaces which appear to focus light onto the specially oriented chloroplasts,

as previously described for the iridescent green plants, Finally, the iridescent leaves

have protein/chlorophyll ratios significantly lower than those exposed to the sun

(Krishnan, 1975). These plants possess a number of remarkable adaptations for

the improvement of photosynthetic efficiency. The analogy of a camera with coated

lenses and high speed film seems apt in describing the function of these plants.

The other blue plants (see Table 1 and Plate 2) are different from Selaginella

willdenowii in several details, although the same colour-producing principle

probably operates for all of the plants. In S. willdenowii the blue colour can be

removed by dipping the plants in water (blueness reappearing after drying). Other

plants do not lose their blueness upon immersion. In the first case, the filter must

be in some contact with the surface. In the second case the filter is likely to be

found within the cell wall, and not directly exposed to the air. Scans of leaf

reflectance for Athyrium crenatoserratum and Lindsaea scandens are similar to

Selaginella willdenowii, except that the blue peak is at 410 p in Lindsaea and

415 » in Athyrium. In both of the flowering species observed, Phyllagathis rotundi-

folia and Begonia pavonina, I could not ascertain the wavelength of the blue colour

because the colour was effectively produced when the incoming light was at a

smaller angle to the surface than the reflected light. The method of measurement

required a fixed incident angle of 60° for both. The physical constitution of the

filters is not presently known, but in all cases the colour is due to a physical optical

effect and not to a pigment.

Other blue plants possess distinct anatomical features in addition to Selaginella

willdenowit. In Athyrium crenatoserratum (Plate 2), the epidermal cells also

contain chloroplasts, and the outer wall has the convex curve described above. The

blue colour is particularly apparent in the older leaves of this common fern, Perhaps

the most spectacular iridescent blue plant of all is the rare begonia, Begonia

pavonina. When looking directly at these plants one sees an intense dark blue

colour. However, if the angle of observation is changed so that light is reflected

at a more acute angle, the colour changes from blue to dull green and then to gold.

An interference filter would account for the blue colour, and the gold colour is

probably due to the refractive properties of the epidermal cells. The plant is now

under further study. The chloroplasts of Begonia pavonina are at the bottom of the

palisade parenchyma cells. Both light refraction and interference appear to enhance

light absorption in this plant, as in Selaginella. The palisade cells are packed very

closely together to minimize light loss due to scattering effects. Immediately beneath

is a compact mesophyll layer, the cytoplasms of which is dark red with antho-

cyanins. This layer probably reflects red light (comparatively more abundant on

the rain-forest floor) right back up to the chloroplasts in the palisade cells. Thus

the Begonia pavonina plants appear to have leaves with lenses, filters and special

mirrors—all to increase conversion of precious light energy to sugars in the shady

forest environment.

FURTHER RESEARCH

Rain-forest floor plants, as in Malaysia, have some spectacular adaptations for

the efficient utilization of light. Further studies may uncover new principles, and

will help illuminate the functional significance of anatomical details of leaf

structure. I see some specific areas of research potential.

First, although there is now some evidence for the operation of thin film

interference filters in plants, no one has yet been able to elucidate the structural

basis of the filter in the different plants. Lee and Lowry (1975) discounted the

‘

Iridescent Plants 27

1S)

ST]

500

Wavelength /

Fig. 4. The relative reflectance of light by green (— — — — — — ant ie

Selaginella leaves as recorded by a spectrophotometer. The bottom curve (——:————)

is the difference between the two curves above and gives a plot similar to that expected for

a thin film interference filter.

cuticle as being the molecular basis of the filter in S. willdenowii; non-polar

solvents, which dissolve cuticle waxes, did not remove the interference. Fox and

Wells (1971) speculated that the structural basis for the optical effect may be the

trapping of thin layers of air by the epidermis, as is well known for some insects.

Critical experimental support for the optical phenomenon of interference should

include physical information on the nature—including the thickness and refractive

index—of the filter. The small dimensions will necessitate the use of sophisticated

electron microscope techniques. If the thickness and refractive index of the filters

were known, then we could precisely predict the nature of the interference effect.

The basis for the filter may well lie in the orientation of cellulose microfibrils in

the cell wall; further research will solve this problem.

Rather esoteric studies on the leaves of shade-tolerant plants may eventually

be of some practical importance, In designing more efficient plantation agriculture

systems for the tropics, biologists will have to consider the kinds of plants to grow

in the understoreys of the plantations. Knowledge of leaf structure may help us

select plants which could thrive in such environments. Who knows? maybe

someday we will spray interference filters onto the leaf surface of understorey

plants, making them blue—and more productive.

Acknowledgements

The following individuals were extremely helpful in the various stages of the

preparation of this manuscript. Prof. R. E. Holttum gave encouragement and

additions to Table 1; Mr. Willis Littke also gave additions. Drs. B. C. Stone,

J. B. Lowry and Peter Ashton, and Prof. Paul Richards gave advice and encourage-

ment. Prof. Anne Johnson identified the liverwort. Encik Jayamohan Krishnan and

Encik Ho Kok Wai provided photographs; Cik Tan Geok San gave technical help

and Cik Wan Shahrizah typed the manuscript.

28 Gardens’ Bulletin, Singapore — XXX (1977)

References

Bjorkman, O. 1972. Photosynthetic adaptation to contrasting light climates.

Carnegie Institution Year Book 71: 82-135.

Burkill, I. H. 1935. A dictionary of the economic products of the Malay Peninsula,

two vols. London, Crown Agents for the Colonies.

Esau, Katherine. 1965, Plant anatomy, second edition. John Wiley and Sons,

New York.

Evans, G. C. 1966. Model and measurement in the study of woodland light

climates. In R. Bainbridge, G. C. Evans and O. Rackham, eds. Light as an

ecological factor. Blackwell Scientific, Oxford.

Federer, C. A. and L. B. Tanner, 1966. Spectral distribution of light in the forest.

Ecol. 47: 555-560.

Fox, D. L. 1953. Animal biochromes and structural colours, Cambridge University

Press, London.

—_——-—— and James R. Wells. 1971. Schemochromic blue-leaf surfaces of

Selaginella. Amer. Fern. Jour. 61: 137-139.

Gates, D. W., H. J. Keegan, J. C. Schleter and V. R. Weidner. 1965. Spectral

properties of plants. Applied optics 4: 11-20.

Gausman, H. W. and W. A. Allen. 1973. Optical parameters of leaves of 50 plant

species. Plant Physiol. 52: 57-62.

Haberlandt, G. 1914, Physiological plant anatomy. MacMillan and Company,

London.

Krishnan, Jayamohan. 1975. Physiological leaf anatomy of Selaginella. B.Sc.

Honours Thesis in Botany, Faculty of Science, University of Malaya, Kuala

Lumpur,

Lee, D. W. and J. B. Lowry. 1975. Physical basis and ecological significance: of

iridescence in blue plants. Nature 254: 50-51.

Martin, J. T, and B. E. Juniper. 1970. The cuticles of plants. Edward Arnold,

London.

Michelson, A. A. 1911. On metallic. colouring in birds and insects. Phil. ae

21: 554-567.

Neville, A. C. and S. Caveney. 1969, Scarabaeoid beetle exocuticle as an oma

analogue of cholesteric liquid crystals. Biol. Rev. 44: 531-562.

Ray, Peter, M. 1972. The living plant. Holt, Rinehart and Winston, New York. R

Richards, P. W. 1932. The ecology of bryophytes. In Fr. Verdoorn, ed:: Manual

of bryology. Martinus Nijhoff, The Hague.

1952. The tropical rainforest. Cambridge University Press, London.

Stahl, E. 1896. Uber Bunte Labblatter. Ein Beitrag zur Pilanzenbiolage II, Ann.

Jard. Bot. Buitenzorg 13: 137-216. ‘“

Vasicek, A. 1960. Optics of thin films. North Holland por pte: Company,

Amsterdam.

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Plate 1. Photographic details of the leaf anatomy of green iridescent plants. A — Transverse

section of a Cyathodium foetidissimum thallus showing the transversely curved epidermal

cells and the distally oriented chloroplasts. B— Upper leaf surface of Monophyllaea patens

showing the long and thick-walled palisade cells which help deflect the green light in a

narrow path out of the leaf.

Plate 2. Photographic details of the leaf anatomy of blue iridescent plants. A — Transverse

section of Selaginella willdenowii leaf. Note the peculiar shape of the epidermal cells and the

large single chloroplasts. B— Scanning electron micrograph of the surface of the S. will-

denowii leaf; the convex curvature of the epidermal cells is easily seen. C — Transverse

section of leaf of Athyrium crenatoserratum. Note the convexly curving epidermal cells. The

two transverse-section photographs are courtesy of Encik Jayamohan Krishnan; SEM micro-

graph was supplied by Encik Ho Kok Wai.

The Ferns of Gunong Ulu Kali

A. G. PiccoTT *

Summary

The various habitats on the peak of Ulu Kali and in the surrounding area, the Genting

Highlands, are described. Some one hundred species and varieties of ferns found between

5,000 feet and the summit are listed.

Gunong Ulu Kali is the southernmost high mountain peak in the Main Range

of Malaya, and is only 20 miles NNE of Kuala Lumpur. The environment and

vegetation of the mountain and surrounding area, the Genting Highlands, have

been described briefly by Burgess (1969). Before the Hotel and Casino Complex

was opened in 1971, this part of the Main Range could only be reached by walking

and climbing through the jungle. Now, at Genting Simpah, a road branches off the

main Kuala Lumpur/Kuantan trunk road and winds its way to the top. At about

5,000 feet, with one mile still to go, the character of the vegetation changes. The

trees are smaller, gnarled and mossy. There are fewer climbers, abundant lower

down, and fewer tree ferns. Higher still, the vegetation changes again and becomes

the dwarf and scrub forest of the Cloud Zone. The trees here are stunted; there is

little or no grass and epiphytes are abundant.

Though few people venture beyond the Complex, the road continues. Your

vehicle can take you within yards of the summit of Gunong Ulu Kali (5,814 feet),

and along the whole length of the ridge to Bukit Genting Chin Chin. The slopes

are steep or precipitous, and treacherous. On a clear morning the views from the

ridge are spectacular: neighbouring mountain peaks covered with jungle, the City

of Kuala Lumpur, the tin mines, and, in the far distance, the sea. It is cool,

refreshing and still. If cloud descends, it becomes chilly, visibility is limited to a few

yards and strong winds may blow.

The area studied includes Gunong Ulu Kali and the associated ridge, and also

some lower slopes towards the two pumps supplying the Complex with water. The

ferns listed were all found between the summit and about 5,000 feet. It was not

possible to record every species in all places, but the list does give an indication of

their frequency and distribution. For convenience, the area was divided into a

number of locations, each the centre of forays into the various habitats nearby.

Ferns of Malaya (Holttum 1966) was used as an aid to identification,

but since the publication of the second edition, the author has reappraised the family

Thelypteridaceae (1971) in which sense the generic names are used here, Specimens

were deposited at the Herbarium of the Royal Botanic Gardens at Kew, and an

incomplete set also at the Botanic Gardens, Singapore.

DESCRIPTION OF LOCATIONS

A is situated at the end of the road along the ridge, 1.75 miles from the car

park. The Radar Station occupies the small peak just beyond. There are still

remnants of scrub forest nearby, and lower down the steep slopes is montane

ericaceous forest, with conifers and Pandanus. Earthmoving has been carried out

on a large scale, leaving cleared areas and fresh earth banks. Drainage is channelled

*21, Jalan Dato Klana, Seremban, Malaysia.

ae Gardens’ Bulletin, Singapore — XXX (1977)

into a steep mossy valley on the eastern side, and this was where Habenaria

angustata, a terrestrial orchid, was found. It was growing in mossy peat and on

rotting tree-trunks.

B is 0.15 mile along the road, back towards the Hotel Complex. The ground

drops away sharply to the east. On the opposite side of the road are wet mossy

hollows with Pandanus klossii, and above them dwarf forest. The trees support an

abundant flora of mosses and lichens, and epiphytic ferns and orchids. In September

and October some trees were almost covered with pinkish purple orchid flowers. A

large crested lizard was first seen here.

Equisetum debile was growing in sand near the roadside drain at 0.25 mile.

Over a period of one and a half years it developed from a small tuft to a large

straggling mass.

C is 0.3 mile along, at the point where the road changes from one side of the

ridge to the other, Strong winds blow through the gap. Large granite boulders form

pinnacles with tree-roots growing over the weathering surfaces. Above the road is

dwarf mossy forest; at road level there are cleared areas and earth banks; and then

the ground drops away on either side. Pholidota parviflora grew here, and Coelogyne

sp. with fleshy red scapes and drooping racemes of flowers on long pedicels.

Further on, the road has been blasted out of the rock of the ridge and there is

a precipitous drop of about 1,000 feet on the western side. Here, an almost

horizontal vein — apparently decomposing felspar — has been exposed. The road

then crosses back to the east side of the ridge.

D, at 0.4 mile, is rather open. In the valley are larger trees, raising their crowns

to the level of the ridge, and tall tree ferns. Earth banks border the road. Fragrant

Coelogyne longibracteata and Dendrochilum simile were common and flowered

towards the end of the year.

E is at 0.5 mile, with large trees in the valley below. The stunted forest above

contains a high proportion of conifers which have the habit of dropping the ends

of their branches. Bulbophyllum angustifolium was in flower in February. There

are wet mossy hollows where epiphytic ferns are abundant and the fruiting bodies

of macroscopic fungi can be found.

F is very exposed at 0.8 mile, the road running almost along the crest of the

ridge. There are fresh earth banks, patches of secondary growth, fallen trees, dwarf

forest and the heads of small steep valleys. An established path goes into the forest

behind but care should be taken: it often becomes a running stream, and there

are scorpions.

G, 0.95 mile, is somewhat sheltered. The trees are larger and some have fallen,

making identification of epiphytes growing in the crown much easier. Coelogyne

longibracteata was abundant and flowered late in the year, Dendrobium cornutum

and D. atrorubens produced flowers in July. Wild bananas, ginger and a variety of

creepers can be seen. Terrestrial ferns with large fronds grow on the edge of the

clearing.

H is 1.1 miles along the ridge and there is a large clearing, an abandoned

building site, to the east. Beyond this a path leads into scrub forest on a short

northerly ridge. The stunted trees, which included Dacrydium beccarii and Rhodo-

dendron orion, are barely 6 feet tall and grow in thick peat and sphagnum root

mat. An arborescent grass was growing near the end of this ridge. Dischidia

astephana and some Nepenthes spp. were also here. In July many flowers of

Habernaria angustata were found. Dwarf forest grows on the other side of the road

and mossy hollows drain into streams which pass under the road.

1: Gunong Ulu Kali, 1.25 miles along the ridge and 0.5 mile from the Complex.

The Television Station has been built on the summit, but just below and across the

site of the construction workers’ hut is an area of nearly undisturbed scrub forest.

Ferns of Ulu Kali 33

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The twisted trees are covered with mosses, lichens and epiphytes. From here the

road winds downhill, with steep debris-covered slopes to the east. Further still the

road has been cut into the side of the mountain and the natural environment has

been much disturbed. Little remains of the natural vegetation: pioneer species are

becoming established.

34 Gardens’ Bulletin, Singapore — XXX (1977)

Locations J, K and L cover larger areas, down to an altitude of about 5,000

feet:

J is the land adjacent to the track which leads to the eastern dam and pump

house, and begins north of the main hotel and beyond the boating lake. It descends

some 1,000 feet, running round the mountain-sides. Deep road cuttings have been

made in the steep slopes. The incinerator and main refuse dump are situated along

here. This area is much more sheltered than the ridge and some very large trees

grow in the main valleys. Near the dam there are small swamps which drain into

the streams.

K begins just beyond the incinerator, where a rough track bears off to the

right towards Gunong Lari Tembakau. For a while a pack of ‘wild’ dogs inhabited

those parts of it where the ground has caved in. Dwarf and scrub forest clothes

the peaks, but the slopes are shaded by taller trees. There are some large mossy

boulders in one of the wetter patches, and amongst these Cryptostylis arachnites

and Ceratostylis ampullacea were flowering in January.

L is along a track which turns off the main road, one mile down and just below

the residence ‘Sri Genting’. It more or less follows the 5,000 foot contour and leads

to the western dam and pump-house, directly under the Television Station and the

Mushroom Farm, The track has been cut into the side of the mountain. It passes

through fairly tall sheltered forest, crossing several streams. Wet rock faces are

found near the streams, in places covered with mosses and liverworts, including

Marchantia sp., but few Hymenophyllaceae.

LIST OF GENERA

Ferns occurring on Gunong Ulu Kali, above 5,000 feet, arranged in families.

Schizaeaceae Grammitidaceae Oleandroideae subfam,

Schizaea Grammitis Nephrolepis

Matoniaceae Bae ey Oleandra

; alymmodon nae

Matonia M : Pteridioideae subfam.

Ctenopteris hs

Gleicheniaceae Scleroglossum Pieris |

Gleichenia Histiopteris

Dicranopteris Thelypteridaceae Pteris

Hymenophyllaceae ee 3 bide Asplenioideae subfam.

ingia

Hymenophyllum ! ; Asplenium

Cory phopteris

Trichomanes S phaeroste phanos Blechnoideae subfam.

Plagiogyriaceae Pneumatopteris Blechnum

Flagiogyria Christella Lomariopsidoideae subfam.

Cyatheaceae Dennstaedtiaceae Elaphoglossum

Cyathea Dennstaedtioideae subfam. Teratophyllum

Polypodiaceae Microlepia Dryopteridoideae subfam.

Dipteris Hy polepis Acrophorus

Belvisia

Li ‘ Ms airy

Gaines indesepidene subfam. Athyrioideae subfam.

; - Lindsaea Diplazium

silica § phenomeris

Cry psinus ss Adiantaceae

Goniophlebium Davallioideae subfam. Pityrogramma

Lecanopteris Davallia Vittaria

Ferns of Ulu Kali 35

List OF SPECIES AND VARIETIES

Ferns occurring on Gunong Ulu Kali, above 5,000 feet, arranged alphabetically.

with locations and brief notes.

1. Acrophorus blumei Ching apud C. Chr. Plate 1.

Becations:,:AlbEe-G, K:. L.

In shady hollows and valleys.

2. Asplenium caudatum Forst.

Locations: J, K, L.

On rocks by streams.

3. Asplenium nidus Linn.

Locations: J, L.

Epiphytic on larger trees in the deeper valleys.

4. Asplenium scortechinii Bedd.

Location: J.

Epiphytes on mossy trees near the dam.

5. Asplenium unilaterale Lam.

Location: L.

In rocky stream-bed.

6. Belvisia revoluta (Bl.) Copel.

Locations: J, L.

Epiphyte on mossy trees in wet hollows.

7. Blechnum orientale L.

Locations: H, and between I and Hotel Complex.

On earth banks and exposed rock faces, often stunted.

8. Blechnum vestitum (Bl.) Kuhn

Locations: A, B, C, E, I J, K.

In dwarf forest with Plagiogyria tuberculata.

9. Calymmodon cucullatus (Nees & Bl.) Pres]

Location: J.

Small epiphyte on mossy trees near dam.

10. Chingia pseudoferox Holtt.

Locations: I, L.

In the open by roadside drains.

11. Christella arida (Don) Holtt.

Location: I,

In open by roadside drain.

12. Coryphopteris badia (v.A.v.R.) Holtt.

Location: G.,

First record for Malaya.

In light shade in mossy hollows.

13. Coryphopteris gymnopoda (Bak.) Holtt.

Locations: A, B, E, I, L.

In light shade in dwarf forest.

14.

iT.

18.

20.

ZA;

pHs

23s

24.

25.

26.

ar.

Gardens’ Bulletin, Singapore — XXX (1977)

Coryphopteris gymnopoda (Bak.) Holtt. var. bintangensis Holtt.

Location: B.

On forest edge.

Crypsinus enervis (Cav.) Copel.

Locations: A, D, E, G, H, J, K.

Epiphyte in moderately exposed places.

Crypsinus laciniatus (Presl) Holtt. Plate 2.

Locations: B, E, G, H, J, K, L.

Epiphyte in open places.

Crypsinus wrayi (Bak.) Copel.

Locations: A-C, E, G-L.

Small epiphyte on very mossy trees.

Ctenopteris contigua (Forst.) Holtt. Plate 2.

Location: H.

Epiphyte in mossy hollow.

Ctenopteris fuscata (Bl.) Kze

Locations: E, J.

Small epiphyte in dwarf forest.

Ctenopteris khasyana (Hk.) Holtt.

Location: E,

Epiphyte in mossy hollow.

Ctenopteris mollicoma (Nees & Bl.) Kze

Locations: A, J.

Epiphyte in exposed situations.

Ctenopteris moultoniit (Copel.) C. Chr. & Tard.

Location: J.

Epiphyte in sheltered valley.

Ctenopteris obliquata Copel.

Location: J.

Epiphyte in sheltered valley.

Ctenopteris tenuisecta (Bl.) J. Sm. Plate 1.

Locations: H, J, K.

Larger epiphyte in deep valleys.

Cyathea contaminans (Wall. ex Hook.) Copel.

Locations: E, G, I, J, K.

Large tree fern, on edge of forest and in valleys.

Cyathea hymenodes Mett.

Locations: F, G, H, J.

Tree fern on edge of dwarf forest.

Cyathea hymenodes Mett. (variety). Plate 1.

Location: H.

Tree fern on edge of clearing. I consider this sufficiently different from the

species in its much-reduced pinnae on the base of the stipe, to call it a variety

for the time being.

Collections. 21.06.75: 1334 (SING), 1335, & 1337 (K), 1338 (SING).

Ferns of Ulu Kali 37

28.

29.

30.

31,

ad.

a3,

34.

a0.

36.

af.

38.

39.

40.

Cyathea lurida (Bl.) Copel.

Locations: B, C, E, F, G.

Tree fern in dwarf forest along ridge.

Davallia trichomanoides BI.

Location: J.

Epiphyte in sheltered valley.

Davallia trichomanoides Bl. var. lorrainii (Hance) Holtt.

Location: L.

Epiphyte, in light shade.

Dicranopteris curranii Copel.

Locations: F, G, J, K.

On fairly exposed earth banks.

Dicranopteris linearis (Burm.) Underwood var. linearis Holtt.

Locations: B, H, I.

Colonising exposed clearings.

Dicranopteris linearis (Burm.) Underwood var. montana Holtt.

Locations: F, H, J.

On side of road cuttings.

Dicranopteris pubigera (Bl.) Nakai

Locations: G, I.

On roadside banks in exposed situations.

Diplazium accedens (B1.) Milde

Location: L.

In small sheltered valley.

Diplazium asperum Bl.

Locations: E, G, H, J, L.

On edge of forest.

Diplazium speciosum Bl.

Locations: B, C, D.

In fairly exposed situations near end of ridge.

Dipteris conjugata Reinw.

Locations: G, H, I, J, K.

Colonising steep earth banks.

Elaphoglossum sp. Plate 2.

Locations: A, B, C, D, F, G, H, J, K.

Epiphyte in less exposed places.

A new species which will be published and probably as E. robinsonii.

Differs from E. callifolium and E. malayense in having pale brown, thin, flat,

entire scales and very broadly pointed lamina. It matches exactly an

incomplete specimen collected by H. C. Robinson in 1913 on _ nearby

Gunong Mengkuang.

Collections. 12.10.74: 1093 (K); 9.11.74: 1095 (SING); 10.11.74: 1094 (K);

10.08.75: 1339 & 1340, 1344-1347, 1358 & 1359 (all K).

Gleichenia hirta Bl, var. amoena (v.A.v.R.) Holtt.

Locations: F, G, I.

In fairly exposed grassy clearings.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

>i.

54.

Gardens’ Bulletin, Singapore — XXX (1977)

Gleichenia longissima Bl.

Locations: F, G, H, I, J.

Scrambling on edge of forest.

Gleichenia microphylla R. Br.

Location: I.

In scrub forest and grassy clearing.

Gleichenia truncata (Willd.) Spr. var. plumaeformis (Presl) Holtt.

Locations: B, G, 1, .L.

Scrambling on edge of forest.

Gleichenia vulcanica Bl.

Locations: F, G, H, I.

In and on edges of dwarf and scrub forest.

Goniophlebium persicifolium (Desv.) Presl

Locations: J, K. L.

Epiphyte in sheltered places.

Goniophlebium prainii (Bedd.) C. Chr.

Locations: E, G, L.

Epiphyte on larger trees in valleys.

Grammitis hirtella (Bl.) Tuyama

Locations: “HAI 9:

Epiphyte in dwarf and scrub forest.

Grammitis hirtella (B\l.) Tuyama var. major Holtt.

Locations: A, H, I.

Epiphyte in very mossy dwarf forest.

Grammitis reinwardtii BI.

Locations: A, B.

Epiphyte on mossy trees.

Histiopteris incisa (Thunb.) J. Sm.

Locations: D, I, Fes.

Scrambling, on edge of forest.

Histiopteris stipulacea (Hk.) Copel.

Location: K.

On edge of small clearing.

Hypolepis punctata (Thunb.) Mett.

Location: E.

Few patches of small fronds in sandy clearing.

Hymenophyllum acanthoides (v.d.B.) Rosenst.

Locations: A, E, G.

Abundant on some mossy tree-trunks.

Hymenophyllum blandum Racib.

Locations: B, H.

Small epiphyte in dwarf forest.

Ferns of Ulu Kali 39

55.

56.

awe

58.

59.

60.

61.

62.

63.

64.

65.

66.

67.

68.

Hymenophyllum exsertum Wall. ex Hook.

iLocauoes:) A. B, C, H, LK. L.

Epiphyte on mossy trees.

Hymenophyllum javanicum Spr.

Location: J.

Epiphyte in valley near dam.

Hymenophyllum serrulatum (Presl) C. Chr.

Locations: I, K, L.

Epiphyte in less-exposed forest.

Lecanopteris carnosa (Reinw.) BI. Plate 1.

Locations: H, K, L.

Epiphyte in crowns of trees. Easily accessible in scrub forest of small ridge.

Lindsaea malayensis Holtt.

Locations: E, G, H, J, K.

In wet mossy rocky hollows, and sometimes on rotting fallen trees.

Lindsaea oblanceolata v.A.v.R.

Locations: H, K, L.

Climbing in wet hollows; L. pectinata of ‘Ferns of Malaya’.

Lindsaea rigida J, Sm.

Location: I.

In sheltered parts of mossy forest, climbing.

Loxogramme avenia (Bl.) Presl

Location: J.

Epiphyte and on rocks in wet valley near dam.

Macrothelypteris torresiana (Gaud.) Ching

Locations: A, I.

In the open by roadside drains.

Matonia pectinata R. Br.

Location: I.

On the top of roadside bank, very exposed, near summit.

Microsorium sarawakense (Bak.) Holtt.

Location: L.

Epiphyte near dam.

Nephrolepis davallioides (Sw.) Kze

Locations: E, J.

Epiphyte on few large trees in deeper valleys.

Nephrolepis tuberosa (Bory) Presl

Locations: I, J.

Fronds small, in peat at edges of clearings.

Oleandra pistillaris (Sw.) C. Chr.

Locations: G, H, J, K, L.

Straggly shrub in open sheltered places, Plants of a different habit, creeping

and producing fronds singly, found on the middle part of the ridge.

69.

70.

Th.

Fa.

73.

74.

75.

76.

77.

78.

79.

80.

81.

Gardens’ Bulletin, Singapore — XXX (1977)

Pityrogramma calomelanos (L.) Link

Locations: B, E, I, J, K.

Usually a small plant, rapidly becoming established on earth banks and in

clearings.

Plagiogyria tuberculata Copel.

Locations: A, B, C, E, F, J, K, L.

Abundant in wet dwarf forest, with Blechnum vestitum.

Pneumatopteris ecallosa (Holtt.) Holtt.

Location: G.

In shade on side of valley.

Pteridium aquilinum (L.) Kuhn var. wightianum (Ag.) Tryon

Locations: D H, I.

Colonising earth banks and small clearings, and rapidly becoming more

common.

Pteridium caudatum (L.) Maxon var. yarrabense Domin

Location: H.

One patch on edge of clearing.

Pteris longipinnula Wall.

Location: 1...

In shelter of forest.

Pteris tripartita Sw.

Location: I.

Single well-developed fertile plant on steep slope of debris.

Pteris vittata L.

Locations: B, E, I.

Common by side of road, but fronds often small.

Schizaea malaccana Bak.

Locations: H, I.

In moss on dwarf trees in sheltered hollows.

Scleroglossum minus (Fée) C. Chr.

Locations: A, B, G, H, I, J, L.

Epiphyte in mossy forest.

Scleroglossum pusillum (Bl.) v.A.v.R.

Location: H.

Epiphyte in scrub forest on small ridge.

Sphaerostephanos polycarpus (Bl.) Holtt.

Locations: A, B, I.

In exposed situations by roadside drains, Differs from lowland specimens in

having a more hairy lower surface and lacking glands on the upper surface,

but matches one collected on Taiping Hills by Day in the 1880s.

Sphenomeris chinensis (L.) Maxon var. divaricata (Chr.) Kramer

Locations: G, J, L.

On steep earth banks and cuttings.

Ferns of Ulu Kali 41

82. Teratophyllum aculeatum (BI.) Mett. var. montanum Holtt.

Location: L. 1}

High-climbing, clothing trunks of tall trees in valleys; upper limit of species

about 5,000 feet.

83. Trichomanes meifolium Bory

Locations: A, B, E, F, H, I.

Epiphyte in mossy dwarf forest.

84. Trichomanes obscurum Bl.

Locations: C, J.

In wet peaty hollows.

85. Trichomanes pallidum BI.

Locations: G, H.

Epiphyte on shadier side of mossy trees; fronds almost white.

86. Trichomanes palmatifidum C. Muell.

Location: C.

Epiphyte in moss on dwarf trees. Probably more abundant but difficult to

detect.

87. Vittaria angustifolia Bl.

Location: F.

Epiphyte in mossy forest.

88. Vittaria elongata Sw.

Location: L.

Epiphyte in shady forest.

89. Vittaria elongata Sw. var. angustifolia Holtt.

Location: L.

Epiphyte in forest.

90. Xiphopteris hieronymusii (C. Chr.) Holtt.

Locations: A, C, F, H, I, J, K.

Epiphyte in mossy forest.

91. Xiphopteris sparsipilosa (Holtt.) Holtt.

Locations: B, J.

Epiphyte in mossy forest.

LisT OF UNIDENTIFIABLE NUMBERS

92. Asplenium sp., possibly A. pellucidum Lam.

Location: J.

Epiphyte in small wet valley near dam.

Collection. 10.11.74: 1092 (K).

93. Belvisia sp.

Location: L.

Specimen from small valley near dam.

Collection. 10.11.74: 717] (K).

94. Ctenopteris sp.

Location: A.

Small epiphyte on mossy tree.

Collection. 12.10.74: 1055 (K).

42 Gardens’ Bulletin, Singapore — XXX (1977)

95. Cyathea sp., probably C. obscura (Scort.) Copel.

Location: J.

Small sterile plant growing near track.

Collection. 10.11.74: 7277 (K).

96. Diplazium sp., near D. pallidum BI.

Location: J.

Fertile fronds from side of track.

Collection. 10.11.74: 7011 (K).

97. Diplazium sp.

Location: L.

Incomplete fertile frond from edge of forest.

Collection. 19.07.74: 1085 (K).

98. Hymenophyllum sp., possibly H. polyanthos Sw.

Location: E.

Epiphyte on tree in valley.

Collections. 12.10.74: 1146 & 1147 (K).

99. Microlepia sp.

Location: G.

Single plant with sterile fronds only, much more hairy than M. puberula;

under observation but lost during road improvements.

Collections. 9.11.74: 7175 (K) & 1176 (SING).

100. Trichomanes sp., probably T. maximum BI,

Locations: H, I.

Fertile fronds from wet mossy hollows.

Collections. 2.02.75; 1144 (K) & 1145 (SING).

101. Trichomanes sp.

Location: L.

Sterile frond from mossy forest.

Collection. 19.07.74; 1248 (K).

The list of fern species was compiled after numerous expeditions to Gunong

Ulu Kali over a period of about four years. Records of previous plant collections

of that place are few. Burkill (1927) indicates that Burn-Murdoch obtained speci-

mens from the mountain in 1910, and that Robinson sent an expedition to nearby

Gunong Mengkuang Lebah in 1913. Ridley (1922-25) mentions the conifer

Dacrydium elatum growing at 2,000 feet on Gunong Ulu Kali, and the Singapore

Herbarium has a record of one of his collections there in 1914. More recently

Mrs. Allen (1963) collected ferns at Genting Simpah, a few miles away and at. a

much lower altitude. And in 1973 and 1975 the staff of the Herbarium,

Singapore Botanic Gardens, collected some ferns but mainly flowering plants from

the summit of Gunong Ulu Kali. Further visits could well produce records of more

and new fern species.

During recent years development has transformed the mountain peak into a —

new hill resort. This has resulted in changes in the composition and distribution of

the flora. In view of the proposed expansion and further development of the Genting

Highlands it would be interesting to follow these changes due to those in the habitat.

Ferns of Ulu Kali 43

Acknowledgements

I wish to express my gratitude to all who assisted in the preparation of this

article, and thank: Dr. R. E. Holttum, for examining and critically identifying the

numerous specimens sent to him, and for his most valuable comments; Mrs, N. P.

Wong of Genting Highlands Hotel Berhad, for the information she supplied;

Mr. John Piggott for the photographic records; Encik Othman bin Ali and

Mr. Philip Khoo, for preparing the art work.

References

Allen, B. M. (1963). Ferns of the Quartz Ridges. Mal. Nat. J., 17: 19-32.

Burgess, P. F. (1969). Ecological factors in hill and mountain forests of the States

of Malaya. Mal. Nat.. J., 22: 119-128.

Burkill, I. H. (1927). Botanical Collectors, Collections, and Collecting Places in

the Malay Peninsular. Gard. Bull. Sing., 4: 113-203.

Holttum, R. E. (1966). Ferns of Malaya (Revised Flora of Malaya, Vol. IL) 2 ed.

Govt. Print. Office, Singapore.

(1971). Studies in the Family Thelypteridaceae III. Blumea, 19.

17-52.

Ridley, H. N. (1922-25). Flora of the Malay Peninsula, Vols. I-V. Reeve, London.

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; é 7

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tiom, left: Cyathea hymenodes, variety with reduced sear ae sa

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Centre: Elaphoglossum sp., fertile (above) and sterile (below) fronds.

Bottom: Crypsinus laciniatus, fertile fronds, x 4.

Specific Concept in Humata pectinata (J. E. Smith) Desv.

G. J. DE JONCHEERE

Rijksherbarium

Leiden

Introduction

The genus Humata Cav., of Davalliaceous affinity, is a typically tropical East

Asian group of ferns, extending with a few outlying species to Madagascar in the

West, Japan in the North and far into the Pacific to the East, but having its centre

of distribution in Malesia. Thus, several species of Humata occur in Malaya-proper

and the reader may be referred to the account of the genus in Holttum’s well-known

book on the ferns of Malaya (1954).

Holttum remarks on the difficulty of specific delimitation in the genus gene-

rally, caused by plasticity and variability, also to be observed, by the way, in other

genera of the same relationship, like Davallia, Scyphularia, etc. In fact, Holttum

suggests in the elaborate observations he makes under the specific descriptions

that several species described from adjacent regions are doubtfully distinct from

the Malayan Humatae recognized by him. But in the case of Humata pectinata

(J. E. Smith) Desv., additional comment is limited to a short note on its ecology.

Still, a long history is attached to the name Humata pectinata, also as to how

it should be interpreted and it may therefore be interesting to follow the vicissitudes

of the specific concept that have been attached to this name.

Historical Account

It was perhaps ironical that J. E. Smith (1793), when first describing Davallia

pectinata, based his diagnosis on two specimens, collected in localities that could

hardly have been farther apart. He had received these plants from Banks and after

the diagnosis Smith added:

‘Habitat in India Orientali, D, Hurloch 1786, eandem forte in Otaheite

legit Nelson. H. Banks.”’

Alston (1933) commented on these types, still present in the Smith Herbarium

now at the Linnean Society in London, in a publication that will be more fully

discussed further on. It should be mentioned at this stage that Alston convincingly

explained that Hurloch’s plant came most probably from the Nicobar Islands, i.e.

the most westerly limit of the distributional area of the Humata pectinata alliance,

whereas the Nelson plant came from the most easterly (Otaheite = Tahiti).

The lamina of the two types are narrow-deltoid, the lowest segments being the

largest. However, the basiscopic side of the lowest segments in the Hurloch-collec-

tion has only one prominent lobe, whereas the Nelson-collection has several,

gradually passing distally into smaller crenations towards the segmental apex.

Gaudichaud (1827) described and illustrated Nephrodium gaimardianum,

typified by a specimen now at Paris from Lawak (= Rawak) Island near Waigeu

off West Irian, collected by his friend Gaimard. This must be considered an

entirely independent description: no mention is made of J. E. Smith’s publication

46 Gardens’ Bulletin, Singapore — XXX (1977)

and the likeness of N. gaimardianum to Smith’s plant. The fern Gaudichaud

illustrates is slightly narrower, lanceolate, and has one lobe on the lowest pair of

segments.

A year later Wallich published his list (1828) and gave the name Davallia

parallela to the Singapore collection No. 251. Wallich’s names are “‘nomina nuda’,

but D. parallela was validated by Hooker, as shown hereunder.

Wallich’s plant is hardly different from Gaudichaud’s type of Nephrodium

gaimardianum, with one small lobe on the lowest pair of segments.

Blume (1828) misconstrued Davallia pectinata J. E. Smith and gave that name

to a species of Prosaptia, at the same time describing Davallia intermarginalis from

Java, the type (at Leiden) being closely comparable to Gaudichaud’s and Wallich’s

types. Blume’s name was recognized as a synonym at an early stage in subsequent

literature.

A beautiful plate of Davallia pectinata J. E. Smith was issued by Hooker &

Greville (1831, pl. 139). The leaf is narrow-deltoid with many basiscopic lobes on

the lowest pair of segments (“‘pinnae’’). It was drawn from a specimen, collected by

Menzies in Tahiti and resembles closely J. E. Smith’s Nelson type of the same

origin.

It is significant that Hooker at that time had no doubts that Menzies’ plant is

conspecific with both the syntypes of J. E. Smith, as clearly indicated in the

elaborate text, accompanying the plate and in which Smith’s type-localities are

specially mentioned:

‘‘Hab. In insula Otaheite Menzies. In Malacca et in insulis Nicobar dictis.

Smith.”’

Amongst the above basic descriptions Hooker (1846) made a critical choice in

his monumental “‘Species Filicum’’. He retained two species, Davallia pectinata

J. E. Smith and Davallia parallela ‘“‘Wall’’. These taxa were, according to Hooker

“undoubtedly nearly allied, but distinct’, to be separated by the following

characters:

a. D. parallela has the shape of the frond less deltoid, also not so deeply

divided and therefore never really pinnate, as in D. pectinata.

b. Margins of segments (pinnae) are horizontally patent and entire, whereas

in D. pectinata they are spreading and crenate.

c. Lower margin of lowest pair of segments generally with one solitary lobe,

rarely more; in D. pectinata pinnatifid with several lobes.

d. Indusium opening to apex of segments; in D. pectinata opening obliquely

to the (crenate) margin.

Apart from these two species, Hooker described Davallia parallela var. f,

based on Cuming 61 from Luzon (BM, L), previously determined by J. Smith

(1842) as Humata pectinata (J. E. Smith). It has no lobing of the lowest pair of

segments, merely auricles.

As to the nomenclature adopted by Hooker: Wallich’s name was deemed

acceptable and thus “‘parallela’”’ was given priority over Gaudichaud’s epithet

“‘gaimardiana”’ which is mentioned in synonymy. Blume’s spurious conception of

““Davallia pectinata” was not understood (Hooker had not seen Blume’s types).

“Java, Blume”’ is included in the enumeration of collectors and localities mentioned

under D. pectinata and at the same time Davallia intermarginalis Blume is mentioned

as a (doubtfully) separate species, the description from the Enumeratio being merely

literally cited on the following page of the “‘Species Filicum”’.

Humata pectinata 47

That Hooker had his doubts on the origin of Smith’s Hurloch type — and

by inference on Blume’s “‘Davallia pectinata”” — is indicated by his remark under

D. pectinata:

“Sir Jas. Smith gives the East Indies as a locality, on the authority of Mr.

Hurloch, but perhaps erroneously, for I have never seen it from the Continent

of India, only from the Pacific’.

This interesting note also clearly shows that Hooker still considered the

syntypes of J. E. Smith to be conspecific; moreover, that an initial concept was

developing in Hooker’s thoughts as to a possible geographical separation of his two

species.

At this stage a short review of the generic assignment of the taxa under discus-

sion would be appropiate. Hooker had a very broad and mostly unnatural conception

of the genus Davallia, as originally conceived by J. E. Smith. However, in the case

of D. parallela and D. pectinata, being definitely Davalliaceous ferns, an inclusion

in Davallia-proper could be defended, especially as Hooker lists these taxa as

belonging to the subgenus Humata (Cav.). Still, in Hooker & Baur’s (1842) ‘Genera

Filicum’’, Humata Cav. is recognized as a proper genus, distinct from Davallia

J. E. Smith, having an indusium that is free at the sides. As this subject is not falling

within the strict scope of this article, it may suffice to say that in modern literature

the genus Humata is universally recognized. One must add, however, that it is clearly

and closely allied to Davallia s.str. and although only differing from Davallia

virtually in one character, is an easily definable group and perhaps is best regarded

as a “Genus of convenience’’, to speak with Copeland.

Continuing the historical review on the subject: in the Synopsis of Hooker

& Baker (1867) the two species were maintained, but the differentiating characters

were for the greater part eliminated. In fact, the only clearly definable feature left

was the lobing of the lowest segments: pinnatifid “with lobes sometimes 4 inch

long” in Davallia pectinata, whereas in D. parallela “‘the lowest pair (is) sometimes

auricled”’. Gone are the differences in frond-form (described as ovate-lanceolate in

both species); the orientation of the sori (oblique in both species); the pinnate or

pinnatifid condition (being cut down nearly or quite to the rachis in both species).

Hooker commented under D. pectinata: ‘“‘Quite similar to the preceding

[D. parallela] in size and texture’.

D. intermarginalis Blume was no longer mentioned. “‘D. pectinata Blume non

Smith’ was included in the synonymy of D. contigua var. D. blumei Mett.

[= Prosaptia] in the Appendix. D. gaimardiana (Gaud.) was mentioned in the

synonymy of D. parallela as oldest name, but not (surprisingly) adopted.

The geographical details were given as: D. parallela in Malayan Peninsula and

Polynesian Islands; D. pectinata in Tropical Polynesian Islands only.

Hooker’s views had a deciding influence generally in the last century and in this

particular case maybe up till now. To mention a few examples, Brackenridge (1854)

recognized Humata pectinata “‘J. Smith” from Tahiti, H. parallela “(Wall)” from

Samoa, enthusiastically referring to Hooker’s (later discarded) differentiation in the

“Species Filicum” regarding the orientation of the sori. Presl. (1849) conceived a

new genus Pachypleuria, differing from Humata Cav. mainly in non-dimorphic

fronds, and made the new combinations Pachypleura parallela (Hooker) Presl and

P. pectinata (J. E. Smith) Presl, the former being even assigned by Fée (1852) as

sole species to yet another genus, Pteroneuron, differing by (pseudo) dorsal sori,

but neither gave any new ideas on the specific delimitation of the species in

question.

The situation seemed to have been consolidated by Christensen (1906) in his

Index, where he recognized Humata gaimardiana (Gaud.) J. Smith (Syn. Davallia

48 Gardens’ Bulletin, Singapore — XXX (1977)

parallela [Wall.] Hooker) and Humata pectinata (J. E. Smith) Desv., the former

occurring from Burma through Malesia to Polynesia, the latter in Polynesia only.

There is in this period, however, one notable exception. It was Luerssen (1871)

who, in his description of the ferns in the Fiji and Samoan Islands, strongly com-

mented on the weak differentiation given by Hooker between Davallia parallela

and D. pectinata:

‘“‘Alle angegebenen, sogar von Hooker und Baker noch festgehaltenen

Merkmale, welche die Davallia parallela Wall von Davallia pectinata trennen

sollen, taugen nicht’’.

which is to say that, according to Luerssen, even the few means of differentiation,

maintained in Hooker & Baker’s Synopsis, most emphatically do not hold good.

His elaborate comments clearly point to the many transitions, even on the same

plant, that exist in the characters, supposed to serve as differential, all being variable

within the traditional specific delimitation of both taxa.

Luerssen gave a full and complete list of the synonymy, in which Davallia

pectinata J. E. Smith is included, but nevertheless adopted the later name Davallia

gaimardianum (Gaud) ‘‘Presl’’, for which reason is obscure. Presl’s combination is

even illegitimate according to present rulings, having been published in the Tentamen

(1836) with a query-mark and as a synonym in the Epimelia (1849).

Apart from Diels (1899) nobody ever took notice of Luerssen’s view and it

remained a lone cry in the — taxonomic — wilderness, one might say.

Christensen’s influence on and significance for modern taxonomic fern-studies

has been reemphasized quite recently by Holttum (1975). It is therefore no wonder

that the former’s specific concept on the subject taxa in the Index (1906) and the

modified version in the Third Supplement (1933) — further discussed below — can

be retraced in all modern regional floras in the Far East, to mention only: Van

Alderwerelt’s (1908) Malayan Ferns, Backer & Posthumus’ (1939) Varenflora van

Java, Tardieu & Christensen’s (1939) Flore Générale de IlIndochine Vol 7,

Holttum’s (1954) Ferns of Malaya and Copeland’s (1958) Fern Flora of the Philip-

pines; and as far as the Pacific is concerned: Copeland’s (1929 and 1932) treatises

on the ferns of Fiji- and Society Islands, Christensen’s (1943) revision of the

Pteridophyta of Samoa and Brownlie’s (1969) Pteridophyta of New Caledonia.

In the Third Supplement Christensen also recognized and confirmed several new

species that had been described in the meantime as belonging to the same alliance.

Van Alderwerelt van Rosenburgh (1920) proposed Humata lanuginosa from

Sumatra, syntypes Lérzing 4567, 4764 and Biinnemeyer 3881 (BO, L). In his

description there is no character that is in any way new to the taxa discussed, apart

from the profuse scaliness and the presence of hairs on the lower surface of the

lamina.

There is another point in Van Alderwerelt’s publication that deserves special

comment. His types are all from Central Sumatra near Toba Lake, from

1100-1800 m, from where several other collections have been made, also from Mt.

Dempo and Sibayak at higher altitudes. This is unusual, as elsewhere, also in

Sumatra, collections are from the coastal plain or foothills and also often from the

sea-shore.

The duplicate of Biinnemeyer’s type-specimen, apparently sent to Leiden some-

what later, has the epithet “‘lanuginosa’’ deleted and ‘“‘gaimardiana’”’ written instead

by Van Alderwerelt himself. This tends to show that the author did not believe

long in his new creation; as several lamina of the syntypes show multiple lobing —

and going by the criteria that Van Alderwerelt had used in his Malayan Ferns, a

rectification in Humata pectinata would have been more justified. ;

;

Humata pectinata 7 49

Copeland was more prolific. In his work on the Ferns of the Society Islands

(1932) he remarks (p. 63):

‘“‘Apparently in this group of [Humata pectinata] each island has developed

a peculiar strain which might be construed as a distinct species’.

but (fortunately) describes and illustrates only two, viz. Humata huahinensis, type

Grant 5295 from Huahine and H. melanophlebia, type Grant 5144 from Tahaa (not

seen). This self-imposed restriction is explained on p. 12 where, as on the other

islands like Moorea and Bora Bora, “‘the characteristic representatives are less fixed

in their pecularities and are accordingly left without distinctive names’’.

Humata huahinensis is described by Copeland as being very near to H. pecti-

nata, but having a black stipe and deviating towards H. gaimardiana by being

pinnatifid and position of sori, also the closely placed segments.

As to Humata melanophlebia, Copeland comments that it is like H. huahinensis,

but pinnate with more remote pinnae (or segments more distally) which are clearly

lobed, mainly basiscopically, the segments becoming inciso-serrate. Copeland (on

page 63) mentions Grant’s collection 5144 [= type of H. melanophlebia] again

under H. huahinensis. Be that as it may, the proposed elevation to “‘species” of

these Pacific representatives does not thereby become more convincing.

In 1940 Copeland described and illustrated another species, Humata tenuivenia,

type Brass 14082 from New Guinea:

“‘Humata pectinatae affinis, venis tenuibus inconspicuis, soris perlatis curvis,

indusiis brevibus destincta’’.

The (iso)types (L and BO) and the photograph show these specimens to be

robust plants from a shady wet habitat (“low epiphyte in coastal swamp-forest’’).

Copeland specially comments on the curved sori which can face the margin and

apex or, in extreme cases, is curved so far that it faces at an angle to the costa too.

A few years earlier Copeland had been the instigator of yet another name. He

had requested Alston to have a new look at J. E. Smith’s types of Davallia pecti-

nata. As a result, Alston (1933, l.c.) lectotypified Davallia pectinata J. E. Smith on

the Hurloch specimen, dispersing Hooker’s previous doubts as to the origin by

showing that it came most probably from the Nicobar Islands and in any case from

Western regions, not from the Pacific. Arguing further, Alston came to the con-

clusion that the name Humata gaimardiana (Gaud.) J. Smith (of Christensen’s

Index with a predominantly Western distribution) must in fact be replaced by

Humata pectinata (J. E. Smith) Desv. and that the other syntype of J. E. Smith

(Hooker’s and Christensen’s Humata pectinata with a Pacific distribution) had no

name. Alston thus reversed the ideas hitherto current on geographical distribution,

but maintained the traditional separation of two specific entities.

The new name Alston introduced was Humata banksii and the typification is

based on the Tahiti collection No. 1769 (BM) by Banks, ‘‘Nelson’s specimens being

poor, both in the herbaria Smith and Banks’’. Alston’s description emphasizes

“profundae pinnatis’”’ and “‘inferioribus margine inferiore pinnatifidis’’, otherwise

does not give any further characters of differentiation from his H. pectinata.

Actually Alston followed the description as included by Hooker & Baker in the

Synopsis for (their) Davallia pectinata. Completely absent are comments on notably

geographical distribution, or a comparison with e.g. Copeland’s previously described

species (1932) of which H. Auahinensis is practically identical with Banks’ plant.

It is, as if Alston was wary to incriminate himself further in sorting out the

antecedents of his new proposal.

50 Gardens’ Bulletin, Singapore — XXX (1977)

Nevertheless, Alston’s views, whether stated or only implied, were immediately

followed by Christensen (1933) in the third Supplement of the Index and con-

sequently in all the more local Flora’s mentioned previously: Western plants now

indiscriminately being called Humata pectinata (J. E. Smith) Desv., Eastern (Poly-

nesian) plants referred to Humata banksii Alston.

But, Copeland did not agree. His comment on the Humatae in the “‘Oleandrid

Ferns of New Guinea’”’ (1940) may be appropriately quoted in full at the end of this

historical survey:

“This is the largest and most difficult genus of the group. Typically

epiphytes, the individuals are subject to wide variations in exposure, and

some of them are very responsive to these differences. Independent of the

environment, some species seem to be notably variable ... Smith combined

a Malayan and a Tahitian plant in describing Davallia (Humata) pectinata.

H. parallela (Wall) Brack. and H. gaimardiana (Gaud.) J. Sm. were sub-

sequently described in the same group. With many specimens from the

Society Islands, it seemed to me that each island had its own, more or less

distinct form, none of these like the comparatively uniform Malayan plant.

It occurred to me that the Malayan plant might be the real H. pectinata,

so I invited comparison of types by Mr. Alston. The result was his finding

the Tahiti specimens [sic!] to be of two species, H. pectinata and H. banksii.

It is my conclusion, not his that the Malayan plant must be H. parallela, for

I have no Malayan specimen duplicated by any from Tahiti.

Some characters which usually serve as specifically diagnostic, serve so badly

in Humata. Size varies greatly as a matter of plasticity (response to environ-

ment). So with size does the dissection of the frond; and so probably do

texture and laxness. Paleae are likely to be deciduous. And dimorphism is

subject to some reversion.

In the light of the foregoing discussion it will be understood that considerable

work on this genus leaves me ill-satisfied. The presentation here given is the

best I can make with the present material’.

One could have full sympathy with this lament, were it not that in the ensuing

key, size, dissection of frond, degree of scaliness and dimorphism figure promi-

nently as differentiating characters. Of the 21 species, Copeland distinguishes in

New Guinea alone, not less than six are described as new, of which one, Humata

tenuivenia falls within the affinity of the taxa under review, as discussed above.

Copeland’s confused rejection of Alston’s interpretation of J. E. Smith’s types

was, for that matter, corrected later (1958) in the Fern Flora of the Philippines,

where he uses the name Humata pectinata (J. E. Smith) Desv. for Philippine

specimens, mentioning Davallia parallela ‘“‘Wall’”’ as a synonym.

Observations

When looking back on the features that have been used to distinguish between

the taxa, as discussed heretofore, a hard core of only two characteristics remain,

viz. the lobing of lowest pair of segments and the pinnatifid against the pinnate

condition.

As to the last feature, the wing alongside the rachis is always distally broader

than below and is, even in the most deeply incised leaves, never entirely absent.

One cannot talk therefore of a real pinnate condition, it is always pseudo-pinnate

and this variable point of distinction is in reality non-existent.

Humata pectinata 51

As to the (basiscopic) lobing of the lowest pair(s) of segments, transitions can

be found on the same rhizome sometimes, from non- one-, two- to multi-lobed. A

clinal pattern can be observed, whereby the multilobed basal segments become less

frequent from East to West (nearly absent in Malaya and the Philippines), whereas

the non-lobed or auricled/one lobed segments decrease from West to East, to being

virtually absent in the Eastern Pacific.

The same pattern of gradual transitions can be found in the length of the lowest

segments, being either longer, equal to, or shorter than the next pair, this making

the overall leaf-shape either narrow deltoid, lanceolate/linear or narrow-ovoid. The

sinuses vary in width from nearly equal to the width of the segments to mere slits,

variable even on the same leaf or plant.

The above observations make differences in leaf-morphology unfit for the

proper distinction of species and the recognition of Humata banksii Alston (and

Humata huahinensis Copel.) seems to be based on weak ground.

Humata melanophlebia Copel. is an extreme case, where the lobing is also

conspicuous on the next lower pair(s) of segments and more or less extended to the

acroscopic side of the segments as well.

Differences in scaliness have hardly been used in proposing separate taxa, the

variability in caducity and density having been generally understood. Van Alder-

werelt’s “‘hairs’’, described in Humata lanuginosa, are nothing but the highly

dissected, hyaline squamules generally found — but not so obvious — on the

underside of the lamina.

Actually, often the scales are not, or not well described, a glabrous or sub-

glabrous condition prevailing in mature leaves. But young leaves (and stipes) are

always scaly and the reader may be referred to Appendix I, where a full description

of the scales can be found.

The venation is always + flattened, conspicuous below, + parallel with none

or 1-2 forkings, 3 forkings only occurring in very large leaves, as in Humata

tenuivenia Copel. The ancillary veinlet round the (terminal) sorus, basis for Fée’s

genus Pteroneuron can also be absent, or the veinlet is forked far below the sorus,

thus becoming just a part of the normally forked venation, Transitions can be

found even on the same segments.

The position and orientation of the sorus is also inconstant. It is always

intramarginal and the indusium can be semi-circular to crescent-shaped, in extreme

cases becoming reniform (Humata tenuivenia Copel.). Distally the sorus mostly

faces the apex of the segment, lower down (obliquely) the margin.

Conclusion

The above review was made in the framework of a general study of the genus

Humata Cay. — and other davallioid genera — the results of which will eventually

be published in the Flora Malesiana. As such, Van Steenis’ forceful, but lucid

essay on “Specific Delimitation” is here considered an excellent guide to extricate

oneself from further confusion. In itself the present study is an illustration of

practically all the pitfalls, so convincingly summed up by Van Steenis, that can

beset taxonomy and although his work is mostly based on vast experience in

phanerogamic taxonomy, it is certainly also fully applicable to ferns.

Therefore, the author cannot but come to one conclusion, viz. that all the taxa

under discussion can best be regarded as one polymorphic species with a wide

distribution, for which the name Humata pectinata (J. E. Smith) Desv. should be

52 Gardens’ Bulletin, Singapore — XXX (1977)

used. Luerssen’s views of nearly a hundred years ago are thus herewith fully

confirmed.

Delimitation from allied species like Humata repens (Linn. f.) Diels, H. vestita

(BI.) Moore and H. heterophylla (J. E. Smith) Desv. gives no difficulty and shall

not be further elaborated on in this context.

Recognition of infra-specific taxa is omitted; the variable, transitional (and

clinal) nature of the characteristics, mentioned hitherto make them unsuitable for

the delimitation of recognisable subspecies or varieties.

The geographic distribution of Humata pectinata as here construed is a clear-

cut case of dependency on certain climatological circumstances. The area covers

practically exactly and totally the zone of small seasonal variations in temperature

(up to 35° F = 20° C) in the Eastern Tropics and Pacific, with the exception of

that part of the zone which is liable to an appreciable monsoon. This explains the

striking absence of the species from (East) Java, South-Sulawesi, Lesser Sunda Isles

and Southern Moluccas in an otherwise continuous distribution throughout Malesia

and Polynesia up to Tahiti. New Caledonia in the South and Taiwan in the North,

in both of which islands the species is rare, are only just outside the zone as above

referred to. Climatological maps covering the region can be found in Goode (1947).

A special note should be devoted to the occurrence of the species in Java.

Although Blume’s type of Davallia intermarginalis came ostensibly from Western

Java, no further collections — apart from an old Junghuhn specimen at BO — have

been made in that area for nearly 150 years. Nevertheless, potentially West Java

(as against the eastern part of that island) could harbour our species, having only

a weak monsoon. Could it have become extinct there by the vast destruction of

lowland forest of that island?

The high-altitude collections in Central Sumatra have already been commented

upon. This could be a special ecotype, but from New Guinea specimens from

medium elevations (1100 m) are also known.

It could be argued that research of a karyological nature, especially cyto-

genetical, could produce better substantiated and more refined results than is

possible by limiting the study to only herbarium material. Whilst this limitation

is recognised it is regretted that such an approach is precluded by the unavailablity

of live material. Whether it would really help in tackling the problem remains to

be demonstrated. The mere thought of the difficulties involved in obtaining live

plants from the whole distributional area and the time-consuming organization in

rigging up a hybridization programme suggests that the above reserve if not illusory

would, in practice be hard to tackle in a satisfactory way.

Finally this relatively uncomplicated, but nevertheless perforce lengthy account

on how the species Humata pectinata has to be construed can be equally repeated

for other taxa of the genus. As the publication of such detailed discussions would

lead too far and be a tedious repetition, this study may serve as an example as

to how the specific concept in the forthcoming treaties of the whole group will be

maintained. That a considerable reduction in names will result — as already

indicated by Holttum (1954) — is certain.

Acknowledgments

To the keepers of the Herbaria at Bogor, British Museum and Kew, I would

like to express my gratitude for cooperation and assistance extended. To my friend

E. Hennipman of Leiden Herbarium I am thankful for reading the script and his

sound advice.

Humata pectinata

Bibliographical References

v. Alderwerelt v. Rosenburgh, C. R. W. K. (1909) Malayan Ferns: 284-291.

. (1920) Bull. Jard. Bot. Bzg. 3/2: 155.

Alston, A. H. G. (1933) Phil. Journ. Sc. 50/2: 175-176, t. 1 fig. 1.

Backer, C. A. and Posthumus, 0. (1939) Varenflora Java: 96.

Blume, C. L. (1828) En. Pl. Javae, Fil.: 229, 230.

Brackenridge, W. D. in Wilkes, C. (1854) U.S. Expl. Exp. 16: 229.

Brownlie, G. (1969) Fl. Nouv. Cal. 3: 149.

Christensen, C. (1906) Index Fil.: 353, 354.

. (1934) Suppl. 3 (1917-1933): 112.

. (1943) Pter. Samoa, Bish. Mus. Bull. 177: 37.

Copeland, E. B. (1929) Ferns Fiji, Bish. Mus. Bull. 59: 89.

. (1932) Pter. Soc. Isl., Bish. Mus. Bull. 93: 11-12, 62-63, t. 12.

—— . (1940) Phil. Journ. Sc. 73/3: 350, t. 3.

. (1953) Fern. Fl. Phil.: 178.

Diels, L. (1899) Nat. Pfl. 1/4: 209.

Fée, A. L. A. (1852) Gen. Fil.: 320, 321, t. 25B fig. 1.

Gaudichaud-Beaupré, C. (1827) Freyc. Voy. Bot.: 335, t. 12 fig. 1.

Good, R. (1947), Geography FI. Plants: t. 21, t. 24.

Holttum, R. E. (1954) Ferns. Mal.: 364~371.

. (1975) Taxon 24/4: 499-500.

Hooker, W. J. (1846) Spec. Fil.: 153, 154, t. 42A.

Hooker, W. J. & Baker, J. G. (1868) Syn. Fil.: 89.

Hooker, W. J. & Bauer, F. (1842) Gen. Fil.: t. 114A.

Hooker, W. J. & Greville, R. K. (1831) Ic. Fil.: t. 139.

Luerssen, C. (1871) Fil. Graeff., Mitt. Gesamtg. Bot. 1: 206-207.

Presl, C. B. (1836) Tent. Pt.: 79, 128.

. (1849) Epim. Bot.: 98.

Smith, J. (1841) Journ. Bot. 3: 416.

. (1842) London Journ. Bot. 1: 425.

Smith, J. E. (1793) Mém. Ac. Tprin 5: 415.

v. Steenis, C. G. G. J. (1957) Fi. Males. ser. 1, 5/3: CLXVII-CCXXXIV.

Tardieu-Blot, M. L. & Christensen, C. (1939) Fl. Gén. Indo-Chine 7/2: 109.

Wallich, N. (1828) List: No. 251.

54 Gardens’ Bulletin, Singapore — XXX (1977)

APPENDIX I

NOMENCLATURE

Humata pectinata (J. E. Smith) Desv. (1827). Prodr.: 323. — Davallia pectinata J. E.

Smith (1793) Mém. Ac. Turin 5: 415; Trichomanes Poir. in Lam (1808) Encycl. 8: 78;

Pachypleuria Presl. (1849) Epim.: 98 — Type: Hurloch/Soc. Unit. Frat. s.n. from Ind. Or.

(probably Nicobar Isl.) as lectotypified by Alston (1933) Phil. Journ. Sc. 50: 175, t. 1 fig. 1.

Nephrodium gaimardianum Gaud. (1827) Freyc. Voy. Bat.: 335, t. 12 fig. 1; Nephrolepis

Pres]. (1836) Tent.: 79; Humata J. Smith (1842) London Journ. Bot. 1: 425; Davallia [Presl.

(1836) Tent.: 128 (comb. ill.)] Kuhn (1869) Zool. Bot. Ges. Wien: 580 — Type: Gaudichaud

(Gaimard) s.n. from P. Lawak = Rawak/Waigeu/W. Irian.

Davallia parallela [Wallich (1828) List No. 251 (nomen nudum)] Hooker (1846) Sp. Fil.:

153, t. 4248; Pachypleuria Presl. (1849) Epim.: 98; Pteroneuron Fée (1852) Gen. Fil.: 320,

t. 256; Humata Brack. in Wilkes (1854) U.S. Expl. Exp. 16 Bot.: 229; Oleandra Keys. (1873)

Pol. Cyath. Herb. Bung.: 41 — Type: Wallich 251 Singapore.

Davallia parallela var. 8 Hook. (1846) Sp. Fil.: 153 — Type: Cuming 61 Luzon.

Davallia intermarginalis Blume (1828) Enum.; 230; Pachypleuria Presl. (1849) Epim.: 98;

Humata Moore (1861) Index: 296 — Type: Blume s.n. Java.

Humata lanuginosa v.A.v.R. (1920) Bull. Jard. Bot. Bzg. 3/2: 155 — Syntypes; Lérzing

4567, 4764, B. Baroe/Sumatra; Biinnemeyer 3881 G. Malintang/Sumatra.

Humata huahinensis Copel. (1932) Bish. Mus. Bull. 93: 11. t. 126 — Type: Grant 5295

Huahine/Soc. Isl.

Humata melanophlebia Copel. (1932) Bish. Mus. Bull. 93: 11, t. 128 — Type: Grant

5144 Tahaa/Soc. Isl.

Humata banksii Alston (1933) Phil. Journ. Sc. 50: 176 — Type: Banks 1769 Tahiti/Soc.

Isl.

Humata tenuivenia Copel. (1940) Phil. Journ. Sc. 73: 350, t. 3 — Type: Brass 14082

Idenburg R./W. Irian.

DESCRIPTION

Rhizome upto several metres long-creeping, slender (14-24 mm), blackish with white-

chalky patches, dictyostelic and dorsiventral, vascular tissue much dissected with two main

bundles; bearing articulated fronds, on short phyllopodia, 2-5 cm or more apart; thickly

set with appressed. imbricated scales, 4 to 5 mm long, + 1 mm wide at the oblong peltate

base, from there tapering to the acute apex, castaneous, the thin margin often hyaline /whitish,

when young bearing marginal crinkly hairs which are mostly soon deciduous, the edge becom-

ing entire at an early stage.

Stipe thin (1 mm g) but firm, as long as, or somewhat shorter than the lamina, green

to reddish brown when living, brown to blackish when dried, sulcate; vascular tissue with

mostly three bundless, becoming fused upwards; loosely set with caducous scales and often

entirely glabrous with age; scales like those of rhizome but not so appressed and more

rounded, becoming smaller, paler, the edge more dissected.

Lamina firm, coriaceous, (5) 7-18 cm long, 4-6 (8) cm wide, narrowly deltoid, lanceolate /

linear to narrowly ovate, dark green above, paler below, pinnatifid/pectinate with truncate/

cordate base and short coadunate apex; segments many (12-30) which are sessile and

confluent by a narrow wing along the rachis; wing occasionally inconspicuous between the

lowest pairs of segments (the lamina then becoming pseudo-pinnate) but always widening

upwards; sinuses deep and narrow, to about as wide as the pinnae but usually much

narrower, becoming distally shallower towards the crenate to entire apex; rachis prominent,

green to brown/blackish, sulcate except at the apex; scales loosely covering the young

unfolding leaves, equal to those on the stipe, however, especially on the lamina becoming

even more hyaline and dissolute, variously deciduous but mostly leaving a few traces on

+ See surface of rachis and costa, as well as on the margins of the often (sub) glabrous

amina.

Segments to 6 (8) mm wide, straight and horizontally patent or a little curved upwards,

the apex bluntly acute to rounded; edge in barren leaves rarely quite entire, commonly

shallowly crenate/sinuate at apex only, less often the incisions becoming more pronounced

and continuing to bottom of sinus; in the very slightly and inconspicuously contracted fertile

leaves incisions more prominent; the lowest pair of segments variable, either somewhat

shorter or equal (ovate-lanceolate form), or if more developed, becoming longer than the next

pair of segments (narrow-deltoid form), the basiscopic edge being either entire, auricled,

often prominently, or two/multi lobed, the lobes grading to the crenated apex, the lowest

segments then becoming unilaterally pinnatifid; venation, apart from the prominent costa,

Humata pectinata 55

distinct on lower surface only, the veins coarse, brown and parallel, not or 1, 2, (3) times

forked, both upper and lower basal primary veins springing from very base of costa or in

larger fronds the lower directly from the rachis.

Sori terminal on the swollen acroscopic (when forked) vein-ending, forming an intra-

marginal, often crowded row; away from the apex often with an accessory posterior veinlet

running round the sorus to just within the leaf-edge; indusium brown, + | mm wide, finely

striated, firm and permanent, semi-circular or crescent-shaped to reniform, the lower straight

to convex side attached in the middle for + half its length, otherwise the indusium free,

opening to the apex or (especially lower down) obliquely to the leaf-edge; sporangia with

12-14 indurated cells; spores monolite, verrucate-rugulate, 20 x 13 4 with crenulated margin.

GEOGRAPHICAL DISTRIBUTION

From Southern Burma (Mergui), Lower Thailand, South Viet Nam, and Taiwan,

throughout Malesia with the exception of those regions with an appreciable monsoon (Middle

and Eastern Java, Lesser Sunda Islands, South-Sulawesi and South Moluccas), extending

further into Tropical Polynesia to Society Islands in the East and New Caledonia in the South.

ECOLOGY

Epiphyte, high and low, on trees; or terrestrial on boles and rocks, in often exposed

habitats near seashore and in light forest, steep banks, even padangs on bare sand, but also

known from swamp-forest in shade. Can apparently stand a lot of exposure, the leaves

curling up in dry periods [Holttum (1954)]. From sea-level to 800 m, going up to 2000 m

in Sumatra and to medium altitudes in New Guinea.

APPENDIX II

ILLUSTRATIONS

Various facies of Humata pectinata (J. E. Smith) Desv.

1. Sinclair, S. F. 40582 (1955) North side of Cape Rochado-Malacca- Malaya.

“Rocky wooded seashore. Creeping rhizome on sea cliffs. Fruiting. Fronds curled

up with the drought. Flattened by immersing in water”.

Two leaves from same rhizome.

2. Lérzing 15533 (1929), East Mt. Sibayak-Sumatra-Indonesia, 1300-1440 m.

“Primary forest, epiphytic, and, in lighter places, on prostrate trunks and rocks.

Not rare”.

Two fronds from same rhizome.

3. Liitjeharms 4674 (1936), Enggano off West Sumatra-Indonesia.

“Epiphytisch op klapperstammen. Strand bij Kiojoh”.

Small leaves on rhizome.

4. Van Niel 3383 (1964), Tutong-Brunei-North Borneo.

“Under clumps of trees in an open vegetation (sand)”.

Two leaves on same (branched) rhizome.

5. Cuming 61 (1836), Luzon-Philippines.

No further details — Type Davallia parallela var. § Hooker.

One leaf from isotype at L.

6. Brass 14082 (1939), Bernhard Camp-Idenburg River-W. Irian-Indonesia.

“Low epiphyte in flooded rain forest of rive-plain at 50 m.” — Type Humata

tenuivenia Copel.

One leaf from isotype at L.

7. Van Balgooy 1983 (1971), W. slope Pahia-Bora2-Society Islands, 300 m.

“Epiphyte, rootstock creeping”.

One large, one small leaf from same collection.

Gardens’ Bulletin, Singapore — XXX (1977)

Humata pectinata

Gardens’ Bulletin, Singapore — XXX (1977)

Curcuma zedoaria

B. L. BuRTT

Royal Botanic Garden, Edinburgh

My previous notes on the nomenclature of Curcuma zedoaria (Notes Roy.

Bot. Gard. Edinburgh, 31: 226. 1972) were inaccurate. At that time I pointed out,

correctly, that ‘“‘Amiomum zedoaria Berg.”’, which is always cited as the basionym

of Curcuma zedoaria, has no existence as a binomial, I then said that the first

valid publication of the name Amomum zedoaria was by Plenck in 1789, by which

time the plant had already been named A. Jatifolium by Lamarck (1783). Where

I, and others, have erred till now is in failing to notice that Amomum zedoaria was

validly published in the German version of Houttuyn’s Natuurlijke Historie by

Christmann & Panzer (1779). Details of this work and its relation to Houttuyn’s

original publication are given by Merrill (in Journ. Arn. Arb. 19: 291. 1938).

Briefly, the important point for our present purpose is that the work by Christmann

& Panzer is not a literal German translation from Houttuyn: it incorporates both

changes and new matter. Merrill listed all the new names he observed, but

unfortunately Amomum zedoaria in the German version escaped him, and, not

having the original work at hand, this helped to lull me into a sense of security.

Although the work as a whole may be attributed to Christmann & Panzer, Merrill

showed that new names appearing in volumes 1-7 should be attributed solely to

Christmann.

Christmann cites a number of authors under Amomum zedoaria: Bergius,

Rheede, Rumphius, Petiver and others. It is the description by Rheede under the

name Kua which gives the most detailed and reliable account of this plant: this

was cited by Bergius at the head of his references and formed the basis of

Lamarck’s Amomum latifolium and it is Rheede’s illustration of it that was re-

drawn for Plenck’s plate. There is every reason for taking this reference as the

“lectotype” of the name Amomum zedoaria Christm., and this I accordingly

propose. —

Amomum zedoaria was published by Christmann in 1779 and it thus provides

the earliest valid post-Linnaean epithet for a Curcuma. Its transfer to that genus

may still be attributed to Roscoe, for Roscoe cites Willdenow and Willdenow gives

the reference “‘Houtt. Linn. Syst. 5, p. 12” which proves to refer to the German

edition of Christmann & Panzer. It may be noted that in Houttuyn’s original

Dutch edition (Handl. Pl. Kruidk. 7: 10. 1777) zedoary is referred to Amomum

zerumbeth, which is Zingiber zerumbet (L.) Roscoe.

As the basis for the name Curcuma zedoaria is Rheede’s description and plate

it has seemed not without interest to supply a translation and reproduction.

Kua

Plate VII

Kua, in Brahmi Acua, is the special and common name for all those plants

that ought to be referred to species of ginger; they are eight in number. Kua,

Tsjana Kua, Mallan Kua, Manga Kua, Mangella Kua, Intsyi Kua, Katou-Intsyi

Kua. The first species likes all soils, especially the sandy ones in which it grows

60 Gardens’ Bulletin, Singapore — XXX (1977)

spontaneously and spreads. The rhizome runs cut horizontally from a single centre,

white, thick, tuberous, with slender fibrous rootlets here and there, firm, run

through with many whitish threads [i.e. vascular bundles], with glabrous skin; it

is divided along its length into tuberous segments about two fingers thick and a

span [c. 19 cm] long, branched, and filled with a clear juice. The taste is sharp,

pinching the tongue, irritant, aromatic, The scent is strong and pleasant as if many

aromas were mixed together. The leaves arise from the ends of the rhizome, up

to a cubit [c. 46 cm] long and two spans [c. 38 cm] wide in the middle; acuminate

at the tip, narrowed at the base, with a prominent midrib; this sends out many

lateral veins, running transversely forward, equally spaced; they disappear at the

extreme edge where they embrace the margin of the leaf with finer veinlets; the

inner part of the leaf is somewhat folded and reflexed, marked with grooves. The

colour on the inner part is dark, on the outer paler. For the rest the leaves are

glabrous, shining, clear. The stems are a foot and a half high, full of greenish

pith, but no shoots are to be seen on them; however, from the rhizome a special

flowering stem arises, covered by a number of long leafy scales very closely

enfolding it; and these leaves are more than a digit long, broadened upwards and

then again contracted to a small point, with the tips bent back from the scape;

they are glabrous with slender longitudinal veinlets which, unless they be broken,

are nearly invisible; the Jower ones are green, the middle ones yellow, the upper

red purple or light blue, the uppermost becoming almost white, pretty, sweet-smell-

ing, and tasting like the root. The flowers are produced in the gaps between the

scales [i.e. bracts], 2 or 3 at a time, bell-shaped, the lower part and inside whitish,

thin, enclosed in a transparent membrane [i.e. calyx], 6-petalled [i.e. with 3 petals

and 3 petaloid staminodes] the upper one pellucid, shaped like a helmet, erect;

they are yellow ana white, folded, and scarcely to be seen unless the whole flower

is separated from the stem and in turn taken apart itself, when one is seen to be

bent down to a fimbriate margin; in the middle the flower produces a thread

[i.e. the style] provided with a spur like a dragon’s head, and recalling a little

winged bird with erect tail [this refers to the spurred anther between the lobes of

which the upper part of the style is held]. The scent is pleasant, but when the

flowers are broken from the pouch and bruised between the fingers they have a

similar smell to the leaves, and a rather bitter flavour. The seed capsules are small

and round, depressed, and contain grey seeds like those of Tsjana Kua, but they are

rarely found. The rhizome is long persistent; the leaves die down and shoot forth

again in July and August. The grated roots, washed several times, leave a flour

much valued by the indigenous peoples who make a porrage from it. A potion of

bruised root and fresh juice arrests all inflammation of the intestines, purges the

kidneys, stops white flux, cures gonorrhea and purges the blood. The juice of the

leaves is drunk by people with dropsy, it takes away the swelling of the abdomen,

is a moderate laxative, and expels the viscid slime of the intestines.

[Footnote by Commelin.] Kua, according to the opinion of the illustrious Paul

Hermann is the garden Zerumbeth of Garcia [da Orta], whose description like that

of the Arabic authors is rather obscure; by the Singhalese it is called Walinghura,

that is wild ginger, the Malabars call it Kua. The opinion of Hermann agrees very

closely with that of our author.

The other plants mentioned by their local names, of which the spelling varies

slightly in different parts of the work, are: —

Tsjana Kua — Costus speciosus (Koenig) Sm.

Mallan Kua — Kaempferia rotunda L.

Manga Kua — Boesenbergia rotunda (L.) Mansf.

Mangella Kua — Curcuma longa L.

Intsyi Kua — Zingiber officinale (L.) Rose.

Katou-Intsyi Kua — Zingiber zerumbet (L.) Rose.

Mala-Intsyi Kua — Alpinia allughas (Retz.) Rosc.

4 uy . §

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eek A eee ie

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- < vies 1 ee gh wet

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; ee

Curcuma zedoaria | 61

Rheede’s description does not, of course, give the precise technical detail that

we look for to-day; for that we turn to Holttum’s account of the Zingiberaceae

of the Malay Peninsula (this Bulletin, 13: 71. 1950). Rheede, however, supplies a

remarkably vivid account of a plant whose whole structure was a novelty at that

time. It has not seemed justifiable to reprint the original latin, but those interested

in the development of botanical terminology may be interested to note that in

it calyx refers to the cup or pouch formed by the bract, filamentum to the thread-

like style, and pollen to the flour obtained from the root.

I have referred throughout to Rheede. Hendryk van Rheede tot Draakestein

was appointed Governor of the Dutch possession in Malabar in 1667. He organized

the bringing together of the materials that were sent back to Holland and published

as the Hortus Malabaricus. Burkill (Chap. Hist. Bot. India, pp. 6-7, 1965) tells

us that the drawings were made by an artist-missionary Matthaeus, that the accounts

of the plants were rendered into Portuguese by an interpreter and thence into latin

by the secretary of the local government Hermann van Doner. Then drawings and

descriptions were assembled by Johannes Casearius and were sent to Holland for

publication, where they were edited partly by A. Seyn but largely, after Seyn’s

death, by J. Commelin who was in charge of the Hortus Medicus at Amsterdam.

Curcuma zedoaria is a species that has been in cultivation for a very long

time and its natural origin has never been precisely established. This is a parti-

cularly difficult problem as it very easily becomes naturalized. In fact it is on

record that although C. zedoaria very rarely flowers under conditions of cultivation,

it does so freely where it runs wild. It is now found in most parts of India and

south-east Asia. Like other members of the genus it dies down completely after

flowering and has a resting phase: which at least suggests that its origin was in

the monsoon areas rather than in Malaya. If, as often stated, it originated in N. E.

India, this would be a good focal point for its spread south into peninsular India,

east to China and SE to Malaya.

Synonymy and useful references are detailed below; those appearing in square

brackets are not nomenclaturally valid names.

Curcuma zedoaria (Christm.) Roscoe in Trans. Linn. Soc. 8: 354 (1807) et

Monandr. Pl. Scitam. t. 109 (1825); Horan., Monogr, Scit. 23 (1862); Baker in

Hook, f., Fl. Brit. Ind. 6: 210 (1890); Trimen, Handb. Fl. Ceylon, 4: 241 (1898);

K. Schum., Pflanzenr. Zingiber. 110 (1904); Gagnepain in Lecomte, Fl. Gen. Ind.

Chin. 6: 67 (1908); Merrill, Enum. Phil. Fl. Pl. 1: 243 (1924) et in Trans. Amer.

Phil. Soc. N.S. 24 pt. 2: 119 (1938); Burkill, Dict. Econ. Prod. Mal. Pen. 714

(1935); Holttum in Gard. Bull. Singapore 13: 71, fig. 5 (1950); Wealth of India

2: 405 (1950); Backer & Bakh. f., Fl. Java, 3: 71, 72 (1968).

Syn: [Kua Rheede, Hort. Malab. 11: 13, t. 7 (1692)]

[Zedoaria Camellus, Herb. Stirp. Luzon, Syll. 23 No. 9 in Ray, Hist. Pl. 3

App. (1704).]

[Zedoaria officinarum Petiver, Gagophyl. Nat. dec. 3, 5, tab. 23 f. 1

(1704-67).]

[Zerumbed vel Tommon [primum] Rumph., Herb. Amboin. 5: 169 excl. t.

68 (1747) — fide Valeton in Merrill, Interp. Herb. Amboin, 164 (1917).]

[Amomum scapo nudo, spica laxa truncata Bergius, Mat. Med. 4 (1778),

ed. 2, 4 (1782).]

Amomum zedoaria Christm. in Christm, & Panzer, Linn. Pflanzensyst.

5: 12 (1779); Plenck, Ic. Pl. Med. 2: 12, t. 11 (1789); Willd., Sp. PI.

1: 7 (1797).

Amomum latifolium Lam., Encycl. 1: 134 (1783). Type: Kua Rheede.

Gardens’ Bulletin, Singapore — XXX (1977)

Curcuma pallida Lour., Fl. Coch. 9 (1790) et ed. Willd. 12 (1793). Type:

‘“‘in agrestis in Cochinchina et Cantone Sinarum’”’ (no specimen known).

Curcuma zerumbet Roxb, in Asiat. Research, 11: 332 (1810) et Pl. Coro-

mand. 3, t. 201 (1819); Alston in Trimen, Handb. Fl. Ceylon Suppl. 281

(1931); Burtt & Smith in Notes R.B.G. Edin, 31: 203 (1972) sub Curcuma

(Erndlia subpersonata Giseke); nom. illegit.

Curcuma speciosa Link, Enum. Pl. Hort. Berol. 1: 5 (1821); nom. illegit.

Studies in the Systematics of Filmy Ferns

II. A note on Meringium and the taxa allied to this

K. IWATSUKI

Department of Botany, Faculty of Science

Kyoto University, Kyoto, 606

Japan

Summary

The evaluated taxonomic features of Meringium and the ‘genera’ allied to this are revised

from the standpoint of comparative morphology. The features observed in detail are denticu-

lation, hairs, internal celi walls, and sorus. Based on the observation of these features, the

systems given by Copeland and Morton are critically discussed by the author who proposes

several amendments for their systems, such as: Hymenophyllum s.str. is distinguished from

Meringium only by the soral construction and Mecodium by the hairs and sorus; Hemicya-

theon is identical with Meringium; and H. levingei is better segregated from Hymenophyllum

s.str.

The species belonging to Hymenophyllum s.1. were classified into 13 genera by

Copeland (1938, 1947). Among them the ‘genera’ centering to Meringium are

treated here to revise the system of the filmy ferns, According to the definition

by Copeland, Meringium is the genus of some 60 species in the tropics and

southern hemisphere, having a combination of such features as: 1) the segments

and wings of axes denticulate at margin, 2) the involucre closed at the lower

portion with bivalvate upper portion, 3) the receptacles growing indefinitely and

extruded from the lips of involucre, 4) cell walls thicker and coarsely pitted.

These characteristics are not completely represented by some species referred to

Meringium. In this part of this series, several taxonomic characteristics of

Meringium are revised, comparing with those of Hymenophyllum s.1., especially

with the ‘genera’ allied to Meringium, e.g. Amphipterum, Buesia, Hemicyatheon,

Leptocionium, Myriodon, and Hymenophyllum s.str. including H. levingei.

Morton (1968) placed Meringium in Hymenophyllum subgen. Hymenophyllum

and classified it as a distinct section named Ptychophyllum arranged next to sect.

Hymenophyllum. In this system, he evaluated the denticulate margin of the

ultimate segments, or of the wings as well, and separated Amphipterum and

Leptocionium, placing them in subgen. Mecodium and Sphaerocionium, respec-

tively. Copeland took much value, on the contrary, to the structure of sorus.

In the following discussion all the species will be named under Hymenophyllum

s.l., pending the nomenclatural discussion until the system of the filmy ferns will

be revised as a whole. The names of subdivisions of the family will be adopted

in accordance with Copeland’s system, or in some cases with Morton’s.

Continued from the Fern Gazette 11 (2 & 3): 124.

This study is partly supported by a grant in aid of scientific research of the Ministry

of Education, Japan, no. 034047.

64 Gardens’ Bulletin, Singapore — XXX (1977)

General features of Meringium sensu Copeland

Before going further, the general features of Meringium will be summarized

comparing with those found in the ‘genera’ allied to this, although some of them

will be treated in detail in the next section.

Rhizome. As usual in the case of Hymenophyllum s.1., the rhizome is long

creeping, irregularly branched, wiry, nearly the same in thickness as or a little thicker

than stipes. The apex is variously hairy with usually caducous hairs, the older

portion being glabrous or very sparsely hairy. Roots are rather irregular in

arrangement and bear hairs on all the surfaces.

Fronds. The size of fronds is variable according to the species, or even within

a species to some extent. The smallest is found in H. lobbii bearing fertile fronds

less than 1 cm in length including stipes. On the contrary, the largest frond of

H. penangianum at hand is nearly 35 cm in length. The fronds seem to elongate

to some extent in humid mossy forest forming slender outlines. H. armstrongii 1s

referred to Meringium and has only a few lobes, and H. lobbii, H. blandum and

several others are pinnate in plan or at most with the pinnae a few times forked.

The larger fronds are pinnate several times with simple or forked ultimate segments.

The outline of the ultimate segments is long and slender, or comparatively short

and broader downwards,

Stipes are terete or winged nearly to the base. The hairs are present or absent

according to the species; the presence on stipes correlates with that on other

axes. The rachis is similar to the stipe, winged throughout in many cases.

The wings on rachis and stipes are similar to the ultimate segments, denticulate

or entire, plane or crisped, glabrous, and (2—) 4-12 (-25) cells broad.

The ultimate segments which, as in the filmy ferns in general, consist of one

cell layer except for the costae, are round to obtuse at apex, denticulate, or entire

in some species, plane to distinctly crisped.

Hairs. The hairs on the lower surface of axes, if any, are the same as those

on rhizome. The coloration of the hairs is pale brown in the apical part of fronds

and rhizome but dark brown in the older portion. The hairs are setose, pointed

at apex, consisting of two to seven cells, and 0.5—2 mm in length.

In H. johorense, H. reductum, and H. armstrongii, multicellular setae are

found at margin of segments, and in Amphipterum the multicellular hairs are at

margin of accessory wing. Except in these cases, no hairs are found on the laminar

portion of the fronds in Meringium and its allies.

Cell walls. The internal cell walls are thick and coarsely pitted in many species

but not so thick and slightly waved in the others. This feature is evaluated by

Copeland to distinguish Meringium from Hymenophyllum s.str.

Sorus. The involucre is obconic at base with distinctly bivalvate lips. The

size of involucre is rather variable and the lips are entire or denticulate. In most

cases, it has been observed that the accessories on the involucre are like wings or

bundles of hairs. The receptacles are clavate, grow indefinitely and extrude from

the lips of involucre.

Character phylogeny of four evaluated features

Among the diagnostic features for defining Meringium and the ‘genera’ allied

to this, four important characteristics will be treated here to revise the system of

the species in question; They are denticulation, cell walls, hairs, and sorus. The

evolutionary trends within each of these features will be discussed to elucidate

the phylogenetic importance of them. |

Meringium and allies | 65

1. DENTICULATION

The dentation of lobes is usually correlated with the veins for the megaphyllous

leaves, though that of the filmy ferns in question is like that found in the lobes of

the bryophytes without any actual relation to the veins. The marginal growth of

the ultimate segments and wings as well is unequal, resulting in the irregular

margin or the occurrence of denticulation, although the arrangement of it is regular

in most cases (plate I). The denticulation here under consideration is, therefore,

morphologically not homologous to that of vascular plants in general, though it

will be described here under the term denticulation.

The denticulation occurs in various forms in the species of Meringium. The

wings of rachis and costae are entire or denticulate, and the denticulate segments

and wings are either plane or crisped. The lips of involucre are toothed in many

species and entire in the others. The occurrence of denticulation is also various

according to the species. In H. edentulum the denticulation is rather rare, while

H, meyenianum and allied species have regularly denticulate margin of segments,

and the segments and wings of H. denticulatum are copiously denticulate and

crisped to some extent. The last tendency proceeds comprehensively in H.

acanthoides.

The denticulate margin of ultimate segments is one of the key characters of

Meringium, sharing it with Hymenophyllum s.str. Among the species of Meringium

regarded by Copeland, however, there are several which have ultimate segments

with entire margin: they are H. macroglossum, H. pachydermicum, H. penan-

gianum, H. pollenianum, H. pulchrum, and H. ricciaefolium. Except for this

characteristic feature of the entire margin of ultimate segments, they are close to

the species of Meringium, or to H. meyenianum, in having the same morphology

in hairs, cell walls, and sorus.

Contrary to this case, a few species referred to Mecodium by Copeland are

included by Morton in sect. Ptychophyllum, an equivalent of Meringium in the

system of the latter: they are H. reinwardtii, H. samoense, H. taiwanense, and

H. thuidium, In the diagnostic features other than the denticulate segments, these

four species are different from H. meyenianum as noted by Copeland, and by

Tagawa (1940) for the third species. The margin of ultimate segments is copiously

crisped for these species, and the denticulation seems to be more irregular (Fig. 2).

The arrangement of laminar cells at margin of segments is irregular in H. fimbria-

tum, appearing somewhat denticulate, though the denticulation is different from

that in Meringium species (Fig. 3). I prefer to exclude these species on the basis

of their morphology in sorus, hairs, and cell walls in addition to the irregular

denticulation at margin of segments, and would place them in Mecodium close to

M. javanicum.

Accessory wings and Amphipterum

There are four species belonging to Amphipterum which is distinct in having

accessory wings not in the plane of the lamina. Copeland (1938) distinguished

Amphipterum generically from Meringium, although he (1937) had correctly

suggested that ‘I would not consider it expedient to distinguish it generically or

otherwise if the wings on the veins were the only distinction’. I would follow

Copeland to consider that H. fuscum and three other species are close to each

Other but are different from Meringium in combination of various features,

admitting that the definition can be given solely upon the accessory wings on the

veins.

Accessory wings are prominent on veins, leaving no room to place any inter-

mediate between presence and absence of this feature (Fig. 10). The cells compos-

ing the accessory wings are quite the same as those of laminar surface in size,

form, arrangement, and in structure of cell walls (Figs. 11 and 23). From this

66 Gardens’ Bulletin, Singapore — XXX (1977)

fact the accessory wings are considered to be the extra-outgrowths of the laminar

surface in different levels from the usual ones. This kind of extra-expansion of

laminar surface is unique to the filmy ferns but is not found in any other group

of the megaphyllous plants.

Except in the case of Amphipterum among the filmy ferns, the same kind of

accessory wings is found in Dermatophlebium, or Hymenophyllum subsect.

Piumosa belonging to subgen. Sphaerocionium, although I know little of the

eight species be!onging to this and am not able to conclude here whether they form

a single taxon or not. In addition, the accessory wings are various in occurrence:

three New Guinean species have the accessory wings on the upper side of veins as

well, while the west Malesian H. fuscum has the wings only on the lower surface

of veins; or among eight species of Sphaerocionium two species have the accessory

wings only on the lower surface and six others have the wings on both the surfaces.

In all the cases the cellular construction of the wings is identical with that of the

ultimate segments of the species concerned. In the cases of both Dermatophlebium

and Amphipterum, I have no sufficient materials at hand to conclude whether each

of them forms a distinct taxon or not, though it will be safely said that the feature

is derived parallel to each other in the above two distinct groups.

‘Scales’ of H. levingei and Buesia

H. levingei has hardly been recognized for a long time in spite of appropriate

description and figures given by Clarke (1880). On the lower surface of midrib

of ultimate segments are the ‘scales’ attached to the axes longitudinally in two

rows and hardly continuous with the next ones on the same row but imbricating

to the opposite ones on the neighbouring row. The base of ‘scale’ is 3-7 cells

in breadth and upper half consists of several cells arranging in one row and

appearing as an articulated hair with larger apical cell of clavate outline (Fig. 12).

The ‘hairs’ on the upper surface are longer than the hair-like portion of the

‘scales’ on lower surface, and they are articulated though not typically in construc-

tion with thin side walls and thick septae (Fig. 13). The cellular construction of

the ‘scales’ is similar to that of laminar portion of ultimate segments (Figs. 14 and

15), so that the relationship between ‘scales’ and lamina is like that of accessory

wing and Jamina in Amphipterum and others. Moreover, it will be noted here that

the ‘scales’ and ‘hairs’ are in two rows on both the upper and lower surfaces of

midrib of ultimate segments, and this arrangement is comparable with that of the

accessory wings in Amphipterum. From these facts, it will be suggested that the

‘scales’ and ‘hairs’ of H. levingei are the denticulation of the accessory wing, the

laminar portion of such a wing being reduced in most cases.

The morphology of the scales is common between 1H. levingei and the species

belonging to Buesia. In the latter the ‘scales’ are often longer, with longer apical

cells. The arrangement is typically in two rows, imbricate, as easily seen by the

naked eye.

Multi-directed projections and Myriodon

I have once referred Bornean materials to Myriodon (Iwatsuki, 1968) based

solely upon the literature for the characteristics of the latter, but am doubtful at

present to regard them as Myriodon which has peculiar projections on various

lines of the axes of various orders. In H. acanthoides the marginal teeth are in

various directions owing to the comprehensive crispature of segments and the

reduction of laminar portion, though the midribs, or axes, are still continuously

winged in this species with the base of projections in one line. Contrary to this

the projections on axes are recorded to be not in one line in the case of H. odonto-

phyllum, differing in this respect from the Bornean materials. The reduction of

laminar surface is not particular among the filmy ferns as known in Macroglena,

Trichomanes setaceum and others. When I referred the Bornean materials to

Meringium and allies 67

Myriodon I considered that the materials are in the extreme form in the series of

H. denticulatum — H. acanthoides — H. brassii, represented by the reduction of

jaminar surface, though it is not probable if Copeland’s observation was correct

concerning H. odontophyllum.

The multi-directed projections of Myriodon may be speculated to have been

resulted by the marginal denticulation and the splitting of accessory wing as

schematically represented in Fig. 42. The teeth, or marginal projections, of segments

and wings of H. acanthoides are similar to the projections of Myriodon and are

also comparable with the ‘scales’ of H. levingei or of Buesia, which may be derived

from the splitting of the accessory wing. In case the accessory wings are in double

lines on both the surfaces, and the laminar parts as well as these accessory wings

are split or reduced to remain as marginal teeth, the multi-directed projections will

possibly be formed as in Myriodon, and the intermediate conditions are available

as noted in the above description, although these ‘conditions’ do not necessarily

represent the evolutionary intermediate but only the character phylogeny (Merkmals-

phylogenie) is treated in this speculation.

2. INTERNAL CELL WALLS (plate II)

The internal walls of the laminar cells of Meringium are thick and

coarsely pitted like those of Selenodesmium and some others belonging to Tricho-

manes s.1, Contrary to these cases, the internal cell walls of Mecodium and

Hymenophyllum s.str. are generally thin and straight or slightly waved at most,

though there are species which do not accord well with the definition of this

feature of the taxa concerned. Among the species which belong to Meringium,

those with thinner walls are: H. armstrongii, H. blandum, H. bontocense, H. bivale,

H., fejeense, H. macgillevirayi, H. multifidum, and H. viride. Even in these species,

the cell walls are not straight but more or less waved.

The thickness of the internal cell walls is also various among the species of

Meringium, From the above observation, the difference in the structure of the cell

walls is rather comparative, and not applicable to the difference in the taxa of

higher rank, although a general tendency can be recognized as is treated such by

various authors.

The cell walls of Hymenophyllum s.str. are usually described as thin and

straight, as illustrated by Copeland (1937) except for H. simonsianum. In my obser-

vation, however, the cell walls are waved or pitted for all the species examined con-

cerning Hymenophyllum s.str. The thickness is various according to the species,

or even within the species for H. barbatum in which the southern form has thicker

and more waved internal cell walls.

Most of the species of Mecodium have thin and straight internal cell walls,

although there are several species with thick and coarsely pitted cell walls, such

as: H. crispato-alatum, H. exsertum, H. fimbriatum, H. flabellatum, H. javanicum,

A. le ratii, H. montanum, H. oligosorum, H. opacum, H. riukiuense. The distribu-

tion of the various forms of cell walls is similar to the case of Meringium and

Hymenophyllum s.str., though the ratio of occurrence is different according to the

‘genera’. From these facts, it is difficult to enumerate the structure of the cell walls

as an important diagnostic feature to discriminate Meringium from the other groups

belonging to Hymenophyllum s.1.

The contents of cells are variously illustrated by van den Bosch (1861),

though they have to be observed in the living condition to evaluate the taxonomic

significance. In this study the living materials were available only for a few species.

68 Gardens’ Bulletin, Singapore — XXX (1977)

3. HAIRS ON RHIZOME AND FROND

The so-called articulated hairs are found on the upper surface of the axes in

H. levingei and in Buesia, though this kind of ‘hairs’ is observed as to be identical

with the ‘scales’ on the lower surface. The discussion on such ‘hairs’ has been given

in the section on denticulation and is not appropriate to note in the section on

hairs, |

At the margin of the accessory wing on the veins in Amphipterum, there are

the hairs identical with those found on the axes and rhizome of most species of

Meringium and Amphipterum, The general morphology of this type of hairs is

described in the foregoing pages in the section of general morphology. The basal

cell of the hairs is attached to the axes or marginal cells of the accessory wing at

the base or at the middle portion (Fig. 35) and the hairs are adpressed. Hairs

of this type are also found in the species of Hemicyatheon, Hymenophyllum s.str.,

Leptocionium, and several species of Mecodium. In these species as well as in

Meringium, the hairs are restricted to the axis and never observed at the margin

of lobes. In this respect it is rather peculiar to observe the hairs at the margin

of the accessory wing in Amphipterum. In some species of Meringium, the fronds

are nearly glabrous, or the hairs are restricted to the very young portion of fronds

and rhizome, though the structure of the hairs if any is the same in all the species

belonging to the ‘genera’ cited above.

On the veins or especially at their junction, most species of Mecodium

bear rather sparsely, smal! multicellular hairs consisting of several sub-transparent

cells with thin walls (Fig. 39). This kind of hairs is quite different from the above

described hairs found in Meringium and ‘genera’ allied to this. The stellate hairs

of Sphaerocionium (Fig. 36) were observed in detail by Morton (1947) who

applied the distribution of hairs to the subdivision of that ‘section’. The stellate

hairs are only known in Sphaerocionium, though the hairs of some species of

Microtrichomanes were referred to the former hairs (Iwatsuki, 1975).

Marginal setae of H. johorense, H. reductum, and H. armstrongii

The morphology of H. johorense was well described by Holttum (1929, 1955)

and Copeland (1937), although the marginal setae were illustrated only by the

line drawing. The marginal setae are in two to six cells, with oblique septa giving

hooked appearance of setae, dark brown, polished, with thick walls and pointed

apex (Figs. 37-38). The setae are similar to those of Trichomanes digitatum group

in appearance except for the multicellular construction in contrast with the

unicellular setae of the latter. Similar setae are also found in Didymoglossum which

belongs to Trichomanes s.1,

The fronds of H. johorense are small, simple to 5-6 lobed, branched nearly

dichotomously, comparable to Trichomanes digitatum in this frond form, though

this is also known in various dwarfed species as noted in the first part of this

series. The margin of lobes is entire in H. johorense, not specialized except for

bearing setae, the cell walls are thick and coarsely pitted, and the receptacles are

extruded. We have no species of Meringium comparable to this species except for

H., reductum and the systematic position of them is unknown at present. Copeland

considered H. johorense as a member of Microtrichomanes but transferred H.

reductum to Meringium without any reasonable interpretation.

H. armstrongii is a small fern in New Zealand, having the fronds simple to

four-lobed, arranged subdichotomously or flabellately. Morton treated this species

as a second member of subgen. Craspedophyllum, and his treatise seems to be

probable considering only thin cell walls and deeply cleft bivalvate involucre,

Meringium and allies 69

though the presence of marginal setae makes it doubtful to this conclusion. At

margin of the ultimate segments of H. armstrongii are the setae of the same type

as those of H. johorense. The marginal cells of the lobes are polished dark-brown,

similar to the cell walls of the setae. In Craspedophyllum, however, the cells of a

marginal row of the lobes are specialized without any marginal setae. The marginal

cells of H. armstrongii are not specialized in structure, differing only in the dark-

brown coloration. Anyway, it will be advisable at present to exclude H. johorense,

H. reductum, and H. armstrongii from Meringium on the basis not only of the

marginal setae but also of various other features.

4. Sorus

The position of sorus is paratactic in all the species in question. The involucre

is obconic in the lower half with bivalvate upper portion. In this diagnosis the

involucre is similar to that of Crepidomanes especially when pl. 16 of Copeland

(1937) is compared with pl. 27 of Copeland (1933) as an example. The involucre

of Crepidomanes is usually longer as a whole with longer tube and entire lips

usually having the pseudoveins. In many cases the lips of involucre are denticulate

at margin in Meringium, but the denticulation at lips of involucre is not in

accordance with the denticulation at margin of segments and wings. On the surface

of involucre, usually on the lower obconic part, there are the accessory projections

in some species, especially in H. denticulatum, H. acanthoides and others. The

depth of cleft is variable to some extent, and the obconic portion is either narrowly

winged or not.

I have made a preliminary observation on the development of sorus in H.

polyanthos and found that at first a small cup-shaped involucre is developed

which becomes deeply cleft at maturity. Further observation is necessary to

elucidate the character phylogeny of sorus.

The receptacles are clavate to cylindrical, growing indefinitely, and extruded

from the lips of involucre. The form of receptacles is variable among the species

of Mecodium, including capitate and clavate receptacles and longer involucre, but

never extruded. The extrusion of the receptacles is various according to the species,

although it will be notable that the exserted receptacles are found only for the

sorus with obconic base. In Mecodium and Hymenophyllum s.str., the involucre is

cleft nearly to the base forming no obconic basal portion, and the receptacles are

never extruded even in the case when they are clavate in structure.

Hymenophyllum s.str. is discriminated from Meringium by the structure of

sori, though the distinction is in some cases obscure. I have examined only a small

number of specimens of American species, although I can point out that several

species which are referred to Meringium have deeply cleft bivalvate involucre

which forms a tube only at the basal portion. New Caledonian H. dimidiatum was

included in Meringium by Copeland, and this is followed by Morton, as an isolated

species, even with the description ‘receptacle, so far as seen, included’. Copeland

noted that ‘sori .... cleft about half-way down’, though in fact the sori are deeply

cleft nearly to the base and form obconic base when the sori are rather deeply

placed at apex of the ultimate segments. From these facts, H. dimidiatum seems

to be better placed in Hymenophyllum s.str. Contrary to that case, Australian H.

cupressiforme is said to belong to Hymenophyllum s.str., by both of the above

authors, though the receptacles are clavate and are larger, extruding in some cases

from the lips of involucre. The small extrusion seems to depend upon the larger

size of clavate receptacles, different from the long extruded receptacles typical

for Meringium.

70 Gardens’ Bulletin, Singapore — XXX (1977)

Classification of Meringium and ‘genera’ allied to this

It is recommended that all the species ever described be examined before

proposing a system in the categories lower than family. In this study, however, the

observation was made only for some representative species, pending the detailed

comparison for every species concerned. The discussion on the system is, therefore,

possible only in a limited sense, and a few comments will be made on the ever

proposed systems.

Before the discussion on the relationship among the species of Meringium

group, the systems proposed by Copeland and Morton will be summarized in a

table:

Copeland (1947)

I. Gen. 1 Mecodium, 2 Craspedophyllum, 3 Hemicyatheon

II. Gen. 4 Sphaerocionium, 5 Apteropteris, 6 Microtrichomanes

UI. Gen. 7 Hymenophyllum

IV. Gen. 8 Meringium, 9 Amphipterum, 10 Myriodon, 11 Buesia, 12 Leptocionium,

13 Rosenstockia.

Morton (1968)

Gen. HI. Rosenstockia

Gen. IV. Hymenophyllum

Subgen. | Hymenophyllum: Sect. 1 Hymenophyllum, 2 Buesia, 3 Ptychophyllum,

4 Eupectinum, 5 Myriodon

Subgen. 2 Sphaerocionium: Sect. 6 Sphaerocionium (Subsects. Ciliata, Plumosa,

Hirsuta, and Leptocionium), 7 A pteropteris

Subgen. 3 Craspedophyllum: Sect. 8 Craspedophyllum

Subgen. 4 Hemicyatheon: Sect. 9 Hemicyatheon

Subgen. 5 Mecodium: Sect. 10 Mecodium (Subsects. Mecodium, Amphipterum, and

Diplophyllum).

1. MERINGIUM AND HYMENOPHYLLUM S.STR.

In definition Hymenophyllum s.str. is distinguished from Meringium ‘by the

more deeply cleft involucre, the shorter receptacle, the absence of peculiarly

(pitted) thickened cell wall, and usually smaller size’ (Copeland, 1838: p. 38).

There are definitions by various authors. but the above seems to be a representative

figuration. Morton cites the direction of sori for Hymenophyllum s.str., but this is

true for only a few number of species.

Copeland (1938) enumerated 14 species for Hymenophyllum s.str. by his

definition. One of them was transferred to sect. Ptychophyllum by Morton (1968)

who added 11 species to sect. Hymenophyllum mostly following the suggestion

given in the Index of Copeland (1938). I can not say exactly at present how many

species belong to Hymenophyllum s.str., which may be classified into two: the

group of H. peltatum represented by entire involucral lips include H. antarcticum,

H. perfissum, H. subdimidiatum, and H. wilsonii; the other is the group of H.

thunbridgense with denticulate involucral lips, including H. barbatum, H. dimi-

diatum, H. revolutum, and H. simonsianum. H. cupressiforme seems to belong to -

the former, though the lips are slightly dentate with shortly extruded receptacles,

deeply cleft involucre, and thin but pitted internal cell walls. I have no sufficient

knowledge on the southern species, although Copeland pointed out that the

dwarfed species in the Far South, e.g. H. minimum, H. pumilo, and H. pumilum,

were the representatives which were difficult to be distinguished between Meringium

and Hymenophyllum s.str., the former two being actually included in sect.

Ptychophyllum by Morton.

Meringium and allies 71

H. levingei has by far no direct relationship to H. thunbridgense. In having

the ‘trichomanes’ on the axes of fronds, this resembles Buesia, though this particular

feature may have been evolved independently in H. levingei and in Buesia. H.

levingei is similar to Buesia in this feature but has entire margin of ultimate

segments and deeply cleft involucre with clavate but not extruded receptacles. This

species should be segregated from Hymenophyllum s.str., probably at the same level

as Buesia from Meringium.

The cell walls are not distinctly different between Meringium and Hymeno-

phyllum s.str., and the pale brown multicellular hairs are found in both of them.

The sole feature to discriminate these two is the morphology of sorus, especially

in the obconic base of involucre and extruded receptacles in the former, From

these facts, Hymenophyllum s.str. and Meringium are considered to be close to each

other accepting the system of Morton (1968) at least concerning this part.

2. MERINGIUM AND MECODIUM

Morton included in sect. Ptychophyllum several species which were included

in Mecodium by Copeland, e.g. H. reinwardtii, H. samoense, H. taiwanense, and

H. thuidium. The interpretation in the structure of leaf margin was different between

the above two authors. As noted in the previous pages in the section of denticula-

tion, I prefer to follow Copeland keeping the above four species in Mecodium.

Among the Asiatic species of Mecodium, those of the group of H. javanicum

is represented by waved or crisped margin of the ultimate segments and wings,

thus representing the appearance similar to Meringium. Some of the species

belonging to this group, e.g. H. fimbriatum, H. javanicum, and H, riukiuense, have

denticulate lips of involucre and pitted internal cell walls. In these features they

share the same characteristics with Meringium, though they appear to be identical

as the result of the evolution along different courses.

H. macroglossum and H. pachydermicum were placed by Morton in sect.

Sphaerocionium subsect. Ciliata ‘without undue strain’, contrary to the treatise of

Copeland who included them in Meringium. As the hairs are quite different between

Sphaerocionium and the other ‘genera’ belonging to Hymenophyllum s.1., we

cannot accept the transferance by Morton as appropriate. We would refer to the

fact that the above two species are close to some species with dense hairs belonging

to Mecodium, e.g. H. exsertum, H. gardneri, and H. oligosorum, in hairiness,

structure of internal cell walls, entire margin of segments, texture, coloration, and

pinnation with broader wings giving the general appearance of shallow incision,

but different only in the structure of sorus. The difference in soral construction is

considered in this paper to record exactly the phylogeny among the species of the

filmy ferns, though the pale brown multicellular hairs are common in Meringium

and Hymenophyllum s.str. but not found in Mecodium except for the above species.

H, barbatum, a representative of Hymenophyllum s.str.. is also similar in appea-

rance to the above mentioned species except for the denticulation at margin of

segments and not so thick cell walls.

Morton noted that only four species of Meringium in the sense of Copeland

had segments with entire margin but in fact the latter author included four more

species without denticulate segment in the latter, and Morton enumerated these

species in sect. Ptychophyllum. As noted in the section of denticulation, the entire

margin is rather peculiar among the species of Meringium, but H. penangianum

seems to be close to H. holochilum except for the entire margin of segments. H.

pachydermicum and H. macroglossum are different in various features from H.

meyenianum, but still belong to Meringium in the broader sense.

2 Gardens’ Bulletin, Singapore — XXX (1977)

Morton distinguished Ptychophyllum and Mecodium in the level of subgenus,

based chiefly on the structure of the margin of segment. In addition to the

difference in denticulation, we can list up the difference in the structure of sorus

and hairs between Meringium and Mecodium, admitting that the difference between

them is comparatively smaller than the one between these two and Sphaerocionium.

3. THE ‘GENERA’ ALLIED TO MERINGIUM

Copeland gave a diagram of affinity of genera and showed that the close

allies to Meringium were Amphipterum, Buesia, Leptocionium, and Myriodon,

adding to them Hemicyatheon and Rosenstockia of doubtful relationship. In the

opinion of Morton, on the contrary, the closest allies to Meringium are Buesia,

Hymenophyllum s.str., Myriodon and a small group in South America named

Eupectinum. He lowered the status of Leptocionium and Amphipterum to sub-

sections under the subgenera Sphaerocionium and Mecodium respectively, that of

Hemicyatheon to a distinct subgenus but retained Rosenstockia as a separate

genus. Thus in his classification of the Hymenophyllaceae he admitted six genera.

Amphipterum. Morton considered that, only two species among four

enumerated by Copeland, belonged to this taxon separating the other two on the

basis of having the segmental margin denticulate. The systematic evaluation on

denticulation has repeatedly been discussed in this paper which concludes that we

can not distinguish any taxa of the filmy ferns in question based solely on this

feature. I found no sound reason in the treatise of Morton and consider four, or

three, species treated by Copeland are close to each other.

Thus confined, Amphipterum includes the species having the leaf margin

denticulate or entire, although the accessory wing on veins not in the plane of the

lamina is common to the species in question. The structure of sorus, hairs, internal

cell walls, and the general appearance are not different between Meringium and

Amphipterum, and it will be accepted here to keep Amphipterum as a distinct

taxon very close to Meringium.

Buesia, Five species were referred to this by Morton (1968) who noted that

he was uncertain that it even needed to be distinguished as a section. As was

suggested by him, Buesia should better be compared with the species having the

accessory wing, according to the speculation that the ‘scales’ may be the broken

accessory wing.

Myriodon. The multi-directed projections of Myriodon were inferred as having

been resulted from the occurrence of laminae in more than two planes and the

splitting of laminar surface or reduction of lamina, remaining only the teeth.

According to the above interpretation, that feature is particular, but Myriodon is

quite similar to Meringium except for the above mentioned feature. Myriodon is

accepted here as a monotypic taxon very close to Meringium, the conclusion

being identical with that given by Copeland and by Morton.

Leptocionium. Indicated by the morphology of setae, this is considered as a

relative of Sphaerocionium but not so close to Meringium. In this I prefer to follow

not Copeland but Morton.

Hemicyatheon. This is recognized by Copeland on the basis of two species

which were considered by him closely related to each other but had fallen into

Meringium and Mecodium by definition, respectively. Morton maintained Hemi-

cyatheon as a subgenus based on the type species, referring another species to

subgen. Hymenophyllum.

H. deplanchei is not very close to H. meyenianum, though there is no problem

to include the former in Meringium, especially by such features as denticulate

margin of ultimate segments, thick and pitted internal cell walls, obconic involucre

Meringium and allies 73

with denticulate lips cleft to the half-way, and exserted receptacles. Copeland’s

negotiation to place H. deplanchei in Meringium was based on his speculation that

this species was close to H. bailayanum which was different from Meringium in

entire margin of ultimate segments, although we refuse to separate any species from

Meringium based solely on this feature. Judging from the soral structure and

morphology of cell walls, H. bailayanum seems to belong to Meringium in broader

sense, probably next to H. deplanchei, with the result again that the entire segment

is not so particular to discriminate the taxa in a rank higher than species. Both

Copeland and Morton consider that H. bailayanum is close to Mecodium and

distinct from Meringium based on the entire margin of segments, although it is

similar to Meringium in soral structure and cell walls which actually indicate

the relationship among the species of the filmy ferns in question. Thus the two

species included in Hemicyatheon by Copeland may better be transferred to

Meringium.

Rosenstockia. I have little information to add to this monotypic ‘genus’,

pending the further study when fresh materials are available.

4. SUBDIVISION OF MERINGIUM

There are several species groups separable from H. meyenianum and its close

allies, although it is rather difficult to diagnose the groups within a few words.

Moreover, there are several species which are included in Meringium by Copeland

but are to be separated from this.

H. johorense, H. reductum, and H. armstrongii are distinct from Meringium

in having dark brown marginal setae. As noted in the section of hairs, these three

are distinct from the other members of Meringium. H. johorense and H. reductum

form a small taxon not so far from Microtrichomanes. I have little information

to add at present to H. armstrongii pending further investigation. It is doubtful

that the latter is referable to Craspedophyllum as done by Morton.

Both H. lobbii and H. blandum are dwarfed species and apparently similar to

each other, although it is difficult to know whether their resemblance is the result

of parallel evolution or of the phylogenetic similarity. It is possible to recognize

the similarity between these dwarf forms and the larger species without any actual

evidence. [ am sure that these species are not close to H. johorense and H. reductum

irrespective of their apparent similarity.

H. pachydermicum and H. macroglossum form a group distinct from the

others in having dense hairs on axes beneath, broader wings of axes and short

entire segments which are broader towards base, giving an appearance of fronds

less dissected, and brownish in dried condition. I have not seen actually the

materials of H. pulchrum which seems to belong here by description and figure

given by Copeland.

Many of the Southeast Asian species referred to Meringium are close to H.

meyenianum having the characteristics described for typical Meringium. I am not

sure at present whether the group of H. holochilum is separable from H.

meyenianum group or not. H. penangianum and H. edentulum with entire or

subentire margin of ultimate segments belong to the group of H. meyenianum in

very strict sense.

I know little at present about the American species, and the southern species

are still not observed in the living condition. H. deplanchei and H. bailayanum

form a separate group even when they are included in Meringium, and no close

allies can be represented here.

74 Gardens’ Bulletin, Singapore — XXX (1977)

5. CONCLUSIVE REMARK

It is necessary to make a detailed revision of the species concerned before

having a systematic conclusion of Hymenophyllum s.1., although it will be still useful

to give here a system rather diagrammatically summarizing the above discussion for

the basis of further investigation. This is a tentative scheme and is a modification

of the systems given by Copeland and by Morton.

Hymenophyllum s.1.

Hymenophyllum

Hymenophyllum s.str. (Hymenophyllum s.str. with two subgroups, H. levingei,

and Eupectinum)

Meringium (Meringium s.str. with several subgroups, Amphipterum, Buesia, and

Myriodon)

Mecodium

Sphaerocionium

References

van den Bosch, R. 1861. Hymenophyllaceae javanicae. Verh. kon Acad. Wetensch.

Amsterdam [X-—6: 1-67.

Clarke, C. B. 1880. A review of the ferns of Northern India. Tr. Linn, Soc. II.

Bot, b. 425-61.

Copeland, E. B. 1933. Trichomanes. Phil. J. Sci. 51: 119-280.

1937. Hymenophyllum, Phil. J. Sci. 64: 1-188.

1938. Genera Hymenophyllacearum. Phil. J. Sci. 1-110.

———— 1947. Genera Filicum. Waltham, Mass. 247 pp.

Holttum, R. E. 1929, New species of ferns from the Malay Peninsula. Gard. Bull.

S.S. 4: 408-410.

1955. A revised flora of Malaya II. Ferns of Malaya. Singapore. 643 pp.

[watsuki, K. 1968. Contributions to the classification of the filmy ferns (2). Acta

Phytotax. Geobot, 23: 117-125, in Japanese.

——-— 1975. Studies in the systematics of filmy ferns I. A note on the identity

of Microtrichomanes. Fern Gaz. 11 (2 & 3): 115-124.

Morton, C. V. 1947. The American species of Hymenophyllum section Sphaero-

cionium, Contr. U.S. Nat. Herb. 29: 139-201.

1968. The genera, subgenera, and sections of the Hymenophyllaceae.

Contr. U.S. Nat. Herb. 38: 153-214.

Tagawa, M. 1940. Studies on Formosan ferns 2, Acta Phytotax. Geobot. 9: 139-148.

Plate I. Denticulation. Figs. 19. Margin of ultimate segment, x 100: Fig. 1, H. poly-

anthos (Iwatsuki et al. M 13034); Fig. 2, H. taiwanense (type); Fig. 3, H. fimbriatum (Tagawa

3347); Fig. 4, H. barbatum (Tagawa et al. T 1502); Fig. 5, H. cupressiforme (Muir 765):

Fig. 6, H. penangianum (Mizutani 252245); Fig. 7, H. meyenianum (Iwatsuki et al. M 13549);

Fig. 8, H. denticulatum (Iwatsuki et al. T 14579 bis); Fig. 9, H. acanthoides (Iwatsuki et al.

M 13235). Figs. 10-11. Accessory wing of H. fuscum (Iwatsuki et al. S 1063): Fig. 10,

ultimate segment with accessory wing, x 100; Fig. 11, Accessory wing, x 400, compare

with II-23. Figs. 12-15. ‘Scales’ and ‘hairs’ of H. levingei (Kanai et al. 8249), x 100: Fig. 12,

‘scales’ on lower surface; Fig. 13, ‘hairs’ on upper surface; Figs. 14 & 15, comparison of

‘scales’ and segment, with focus on ‘scales’ in 14 and on segment in 15.

Plate II. Internal cell walls, all x 400: Fig. 16, H. meyenianum (Iwatsuki et al.

M 13549); Fig. 17, H. denticulatum (Iwatsuki et al. T 14579 bis); Fig. 18, H. blandum

(Iwatsuki ei al. M 13515); Fig. 19, H. multifidum (Pichi Sermolli 6257); 20, A. bontocense

(Tagawa et al. T 4815); Fig. 21, H. johorense (Mizutani 2724 a); Fig. 22, H. deplanchei

(Balansa 1641); Fig. 23, H. fuscum (Iwatsuki et al. S 1063); Fig. 24, H. barbatum (Tagawa

et al. T 1502); Fig. 25, H. simonsianum (Kanai et al. 725175); Fig. 26, H. wilsonii (Jermy

et al. 8473); Fig. 27, H. cupressiforme (Muir 765); Fig. 28, H. levingei (Kanai et al. 8249);

Fig. 29, H. polyanthos (Iwatsuki et al. M 13034); Fig. 30, H. fimbriatum (Tagawa 3347);

Fig. 31, H. oligosorum (Iwatsuki 6657); Fig. 32, H. exsertum (Kanai et al. 725170); Fig. 33,

H. hirsutum (Standley 21479).

Plate III. Hairs and denticulation. Figs. 34-39. Hairs: Fig. 34, multicellular hair of

H. exsertum (Kanai et al. 725170, x 100; Fig. 35, multicellular hair at margin of accessory

wing of H. fuscum (Iwatsuki et al. S 1063), x 100; Fig. 36, stellate hairs of H. hirsutum

Standley 21479), x 100; Fig. 37, marginal setae of H. johorense (Mizutani 2724 a), x 100;

Fig. 38, oblique septa of Fig. 37, x 400; Fig. 39, multicellular hair of H. polyanthos

(Iwatsuki et al. M 13034), x 400. Figs. 40-41. H. acanthoides (Iwatsuki et al. M 13235):

Fig. 40, a plant, x 5; Fig. 41, segment with crisped denticulation, x 100.

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Calamus caesius and Calamus trachycoleus Compared

J. DRANSFIELD

Royal Botanic Gardens, Kew

Summary

Two closely related rattan palms, Calamus caesius and C. trachycoleus are shown to have

remarkably different habits; the silvicultural significance of the difference in habit is discussed.

Throughout the Lower Barito area of South Kalimantan, Indonesian Borneo,

a rattan species receives intensive cultivation. This species, Calamus trachycoleus,

is the only species cultivated on a large scale (i.e. thousands of hectares). It is

closely related to Calamus caesius which also receives some degree of cultivation

in the area and elsewhere. Though so similar to Calamus caesius, C. trachycoleus

differs in one feature of its growth form; this difference has a major effect on the

habit of the rattan and is one of the most important features of the species which

has led to its ultimate supremacy as a plantation rattan. Holttum (1955) in his

classic paper on growth forms of monocotyledons has indicated the importance

of an understanding of the basic growth forms of monocotyledons; here is an

example from the rattans where a difference in the degree of growth of one feature

of a basically similar growth pattern differentiates between two species and

accounts for the economic success of one over the other in cultivation.

DESCRIPTION OF THE GROWTH FORMS OF CALAMUS CAESIUS AND

C. TRACHYCOLEUS

CALAMUS CAESIUS BLUME

This rattan is very widespread being found in Sumatra, the Malay Peninsula

including Southern Thailand, and Borneo, at altitudes ranging from near sea-level

in coastal peat swamp forest to 1000 m or more in the hills; the wide range may

in part be due to the fact that it receives a rudimentary cultivation and is some-

times planted in primary forest near villages (Dransfield pers. obs. and hearsay

from villagers in Borneo). It also seems to tolerate a great range of soil conditions

from seasonally flooded alluvial clay soils, peat-swamp soils to well-drained steep

soils on varying substrata in Hill Dipterocarp Forest. It is however commonest

as a lowland plant on alluvial flats beside rivers. Throughout Borneo, the indigenous

people often plant a few clumps of this and occasionally other species at the edge

of their villages or near their longhouses. Calamus caesius without doubt produces

the best quality small diameter class (7-15 mm) cane of all rattan species entering

the rattan trade.

Calamus caesius produces rather dense clumps of many aerial stems, often

more than 15 in number radiating from a condensed system of short underground

rhizomes. Initial growth of the seedling produces an orthotropic stem about 1 cm

in diameter including the leafsheaths. This seedling stem branches from the basal

nodes to produce one to three sucker shoots which are also orthotropic; subsequent

suckering from these sucker shoots produces short subterranean rhizomes c 3 cm

in diameter and up to 10 cm long which bear pale brown scale leaves and short

condensed internodes. Such rhizomes eventually (growth rates of the rhizomes

are not known) metamorphose into rapidly growing orthotropic shoots of diameter

1.5 — 2.5 cm including the leaf-sheaths with lower internodes 50-100 cm or more

76 Gardens’ Bulletin, Singapore — X XX (1977)

in length bearing at the base leaves with highly reduced laminae often consisting

of rachis alone and long-sheathing bases. Subsequent leaves develop more and

more leaflets until the adult leaf shape is attained with the rachis bearing 10 or

more grouped leaflets on each side, the rachis tip terminating in a cirrus. As

the rhizome metamorphoses into an orthotropic stem, the two nodes at the area

of metamorphosis each develop a branch with the potential of continued rhizoma-

tous growth though the potential may not always be realised. Some of the

branches remain dormant as bulb-like shoots. As each rhizome grows into an

aerial stem, there is the potential for replacement by two new rhizomes and hence

exponential increase in the number of aerial stems in the clump. However, because

of the shortness of the rhizomes, many of the potential new rhizomes become

juxtaposed and squashed and in this condition remain as bulb-like dormant shoots

as described above. Further development of these dormant shoots depends on the

opening up of the clump either effected by death of orthotropic shoots or by

clearance by man of debris during cultivation of this species (pers. comm. from

villagers in Borneo).

CALAMUS TRACHYCOLEUS BECC.

This rattan is only known from South Borneo within the watersheds of the

rivers flowing into the Java Sea — i.e. the Barito, Kapuas (Kalimantan Tengah),

Kahayan, Mendawai, Sampit and Seruyan Rivers. In this area it is found growing

on seasonally-flooded riverbanks on alluvial clays and the margins of swamp forest.

It receives intensive cultivation in the Barito Selatan area upriver from Kuala

—

fais Wy

SP, o? ya

4 me My if

fha> WZ

Ay itr fos

Y, EH No aa

lt Ail,

1. The base of a clump of Calamus caesius with most roots removed; note the short

rhizomes, bulb-like dormant shoots, and the smaller diameter of the aerial shoots com-

pared with that of the rhizomes, (Dransfield 3933, Sungei Jaya, Kalimantan Tengah,

4.2.1974)

Calamus

2. A piece of a stolon (“selantar’) of

Calamus trachycoleus at the point of

metamorphosis into an orthotropic; stem

sheaths have been removed to show the

production of two branches at the point

of metamorphosis. Dransfield 3929,

Sungei Jaya, Kalimantan Tengah

4.2.1974).

Kapuas on the Barito River, being grown

in plantations on riverside seasonally-

flooded alluvia] soils. Though eminently

successful as a plantation crop and in the

wild, apparently, as a riverbank colonizer,

it is unknown elsewhere in Borneo.

Calamus trachycoleus produces a small

diameter class cane (7-15 mm) with

internodes generally shorter than those of

Calamus caesius and of not quite such

good quality; yet it accounts for about 80

per cent of the rattan trade on the Barito

River.

Calamus trachycoleus is immediately

distinguishable from C, caesius in produc-

ing diffuse open colonies rather than dense

clumps; rather than having a condensed

system of short underground rhizomes, it

spreads by means of lax above-ground

stolons. Initial growth of the seedling is

similar to that of C. caesius, the first stem

and one or two suckers being orthotropic.

Subsequent branching results in robust

stolons up to 4 cm in diameter which grow

along the soil surface or slightly raised

above ground level on short adventitious

roots. The stolons (known locally as

“selantar’) bear sheathing pale brown

scale leaves and short internodes 4-7 cm

long and like the rhizome of C. caesius

metamorphose into orthotropic stems. The

length of the stolons from point of origin

to point of metamorphosis may be 3 m

or more. As in Calamus caesius, two

branches are produced at the area of

metamorphosis, one each from adjacent

nodes, but unlike C. caesius, both branches

grow out to produce new stolons unless

damaged; no dormant branches have been

observed and this is regarded as being

correlated with the open growth of the

clump and apparent lack of competition

between the branches.

Each branch in Calamus trachycoleus

is apparently adnate to the internodes

of the proceeding leaf and is hence carri-

ed out of the scaleleaf axil; however their

position suggests an axillary origin rather

than an internodal or other anomalous

origin. This feature of Calamus trachyco-

leus deserves anatomical investigation. Of

the two branches produced at each meta-

morphosis, the proximal tends to develop

before the distal and may be already 30 cm

in length before the distal emerges

78 Gardens’ Bulletin, Singapore — X XX (1977)

from the scale leaves, Because of distortion the proximal branch of the stolon

appears to continue the growth of the subtending stolon and at first sight the

orthotropic stem appears as the branch rather than the main axis. If young material

is examined at a stage when a stolon tip begins to grow upwards, the two new

stolons are seen to be branches rather than one of them being a continuation of the

stolon.

Unlike Calamus caesius, C. trachycoleus is a rapidly invasive species and the

potential of exponential increase in number of aerial stems is usually realised, Once

a plantation of C. trachycoleus has been established very little cultivation is neces-

sary and harvests can be made after an initial 7-10 year period at 2 yearly intervals.

In C. caesius on the other hand, clumps require clearance of debris to encourage

development of new shoots and in the Barito Selatan area of South Borneo at any

rate, only two main harvests are obtained, an initial one after 7-10 years followed

by a second after a further four years; after the second harvest the clumps are

supposedly exhausted.

DISCUSSION

In his original description of Calamus trachycoleus Beccari (1913) comments

that the rattan is allied to C. caesius but most closely related to C. pogonacanthus.

Recent fieldwork in Borneo has allowed the present author to make several

collections of C. pogonacanthus which is now seen to belong to a distinct group of

species in Calamus all with the peculiar feature of bearing both a cirrus and a

flagellum, Cirrus (barbed whip at the end of the leaf representing an extension

of the leaf rachis) and flagellum (barbed whip borne on the leaf sheath and

representing a sterile inflorescence) are the two major climbing organs found in

Malesian rattans and are usually mutually exclusive. The presence of both in

C. pogonacanthus and a few other species is hence noteworthy. C. trachycoleus

bears a cirrus only and hence is here regarded as being more closely related to

C. caesius than to C. pogonacanthus. Furthermore, in the field, without reference

to the remarkable difference in habit, C. caesius and C. trachycoleus are distinguished

with some difficulty — both species are about the same size, have very similar

leaflet arrangement, and have white indumentum on the lower leaflet surfaces,

though in C. trachycoleus it is sparse and usually only present on young leaves.

The leaf sheath with its abundant minute thorns forming a scabridity between the

sparse large triangular thorns in C. trachycoleus and lacking such scabridity in

C. caesius is the only reliable character for distinguishing the two species in the

herbarium; yet in the field the two are seen to be remarkably distinct because of

their differing habits.

The striking differences in development of the suckering habit of these two

rattans illustrates how important such differences may be to the silviculturalist;

further studies of rattan suckering habits may produce other examples of closely

related species differing in habit and are seen to be of great importance in a

consideration of the ecology of rattans.

ACKNOWLEDGEMENTS

I should like to thank Sdr. Damhuri of the Herbarium Borgoriense for

preparing the figures.

REFERENCES

BECCARI, O. (1913): Asiatic Palms — Lepidocaryeae, Part 1 The species of

Calamus. Appendix. Ann. R. bot. Gard. Calcutta 11.

HOLTTUM, R. E. (1955): Growth habit of monotyledons — variations on a

theme. Phytomorphology 5 (4), 399-413.

Synaptospory: a hypothesis

A possible function of spore sculpture in pteridophytes

K. U. KRAMER

Institute for Systematic Botany

University of Ziirich

Switzerland

Summary

Part of the spore output of some ferns is shed in groups of spores, sometimes still

contained within the sporangium. It is theorized that spore sculpture, particularly a distinct,

strongly sculptured perispore, plays a part in keeping, or bringing, spores together. This is

important for increasing the chance for intergametophytic fertilization. The want of a

pronounced spore sculpture in most epiphytic ferns and the strongly sculptured spores of

heterosporous pteridophytes are brought in connexion with the phenomenon, tentatively

called synaptospory.

Until recently, pteridologists used to distinguish between fern taxa (genera

or groups of higher rank) with a perispore and taxa without one, and much

taxonomic value was attributed to this character (e.g... Kramer in Lanjouw &

collab., 1968; Wagner, 1974). Through Lugardon’s fundamental work (e.g., 1971,

1972a, 1974) it has now become known that the perispore is apparently of universal

occurrence in ferns, and it has also been found in so-called fern allies (e.g., Lugardon

1969, 1972b, 1973). It is therefore no longer correct to speak of ferns with and

ferns without a perispore. Instead, one should distinguish between spores having a

perispore that is firmly appressed to the exospore, and others where the perispore

forms a distinct, usually strongly sculptured layer. To avoid this awkwardness in

expression I shall continue to speak below of spores with and spores without a

perispore, being, of course, fully aware of the incorrectness of these terms.

In most cases a strongly developed surface sculpture of a spore was attributed

to the perispore although in some cases the exospore was held responsible for the

surface features. It now seems as if in all these cases a perispore is present, but

where it seemed to be absent it firmly adheres to the exospore and cannot readily

be distinguished except in cross-sections examined with the transmission electron

microscope. ;

Whatever the morphological interpretation, the function of this surface sculp-

ture has remained a matter of conjecture, not to say an enigma, as it remains to this

day, to a certain extent and largely as to details, in the pollen grains of seed plants.

In the following I want to put forward a hypothesis that tries to account for the

presence of spore sculpture in pteridophytes. I wish to stress that it may only be

one of several possible functions, and that the attempt at an explanation is no

more than a hypothesis, as for the moment it rests on too few observational data. I

hope, though, that it may lead to further experimental work and will either be

confirmed or, if disproved, will show to have been a stimulating error.

When we review the “‘presence”’ or ‘‘absence” of a perispore (inverted commas

because of the restrictions stated above). or, more generally, of a pronounced

external sculpture in the spores of modern ferns, it becomes immediately apparent

that large groups are without such sculpture and other, equally large and as a rule

80 Gardens’ Bulletin, Singapore — XXX (1977)

very naturally defined ones, possess it. Wagner (l.c.) regarded the groups with a

pronounced perispore as derived ones, and the perispore in general as a specializa-

tion, whatever its function. It seems to me that there is another striking correlation,

namely between the “‘presence” or “‘absence” of a perispore and the habitat of

the fern, that has, to my knowledge, not been reported before.

If we regard the spore wall of taxa of higher order among the modern ferns,

it springs to the eye that epiphytic genera, genus groups, etc., in most cases have

more or less smooth, ‘‘perisporeless’”” spores: Vittarioids, Davallioids, Grammi-

tidoids, Polypodioids, Hymenophyllaceae. Minor exceptions can be found with ease,

e.g. in the Polypodioids in Drymoglossum (see Wagner, l.c., fig. 2), in the

Davallioids in Gymnogrammitis (see Ching, 1966, and Sen & Sen, 1971), etc., and

there is one important major exception: the genus Elaphoglossum, This, though

containing some terrestrial species, as do indeed most epiphytic groups, consists

almost entirely of epiphytes; but its alliance is with a group of terrestrial or

scandent genera known as the Lomariopsidoid ferns or Lomariopsidaceae*).

‘*Perispore-less”’ spores occur, of course, in a considerable number of essentially or

entirely terrestrial fern groups. Examples are Osmundaceae, Plagiogyriaceae and

Dennstaedtioids.

A second observation, of high relevancy in this context, was recently published

by Schneller (1975). It has nearly always been assumed, explicitly or tacitly, that

fern spores are scattered invidually as far as possible. Schneller showed that in

Dryopteris spp. a not inconsiderable part of the spores is shed in groups of two,

three, four, or more.

According to Gastony’s findings (1974), in certain tree-ferns not the spore but

the sporangium, with its complete spore-content, is the unit of dispersal. The spores

germinate whilst held together by the sporangium. I have the strong impression

that at least partial retention of spores in the sporangium. with subsequent shedding

of the entire structure, is not a rare phenomenon in ferns.

The pioneer work by Naf (1961) and Voeller (1971) has shown that the

development of gametophytes and their sex organs is a process subject to physiologi-

cal interaction between closely associated prothalli. These interactions greatly tend

to favour intergametophytic fertilization and hence gene exchange. Various authors

have reported on the much greater viability of fern sporophytes produced by

intergametophytic as opposed to intragametophytic fertilization (e.g., Lloyd, 1974).

Terrestrial ferns by and large may be supposed to grow in an essentially stable

environment (there are, of course, exceptions like pioneers on bare soil, lava flows,

etc.) where the rate of mortality of individual plants is relatively low. Once they

are mature, individual plants often grow slowly and are supposed to be potentially

very long-lived, e.g., in Osmunda regalis. Except when new habitats are colonized,

sexually produced new generations will follow each other at rather long time

intervals, and it must be of great importance for the plants to maintain their genetic

variability, and to avoid the influence of recessive deleterious genes, by increasing

the chance for cross-fertilization when new plants are sexually produced from

prothalli. The coherence of some spores, whether through the action of the spore

wall or by retention in the sporangium, must greatly enhance this chance, and

external spore sculpture may be one mechanism by which the former is achieved.

Spores from one sporangium will not be all genetically identical, unless the mother

plant is one hundred percent homozygous. Schneller (1.c.) found many groups of

more than four spores being dispersed as a unit, so the tetrad is certainly not the

* On the evidence available at present I find myself unable to agree with Tutin (1964) and

Pichi Sermolli (1968) who distinguish a family Elaphoglossaceae comprising only Elapho-

glossum and satellite genera.

Synaptos pory 81

limit beyond which the spores do not cohere. And there seems no reason to suppose

why spores produced by different plants from one colony, and even from different

ones, could not meet after having been dispersed by air currents, runoff water, etc.,

for instance, in a surface-film of rainwater, cohere, and germinate together, with

subsequent cross-fertilization.

In contrast, most epiphytes inhabit a very unstable milieu, The branches and

twigs they colonize have but a limited life-span, and they are thus bound to colonize

new places continuously in order to survive as a species, Many do this by producing

long-creeping, branching rhizomes, or sometimes runners, and such ferns very often

inhabit the smaller branches and twigs of trees and shrubs, as I observed myself

in some parts of tropical America, and as can be deduced from de la Sota’s diagram

(1971, fig. 13) illustrating Costa Rican fern epiphytes; see also Johansson (1974).

The epiphytes with short rhizomes, often more massive, or even forming so-called

nests, favour the more stable habitats of the main trunks and crotches of larger

branches.

For colonizing new stations, these, as it were, pioneer epiphytes thus depend

to a great extent on spore dispersal, which must be sufficient and efficient in order

to ensure maintenance of a sufficiently large population, in spite of considerable

turnover. This must largely be achieved by dispersing individual spores, and self-

fertilization in lone prothalli originating from singly spread spores should be much

more common in these plants. Indeed, coherence of spores in groups would impede

dispersal. Wagner (l.c.) suggested that the lack of a perispore in Davallioid ferns

should be regarded as a result of secondary loss of this organ, and if that is true,

it might be regarded as an adaptation to epiphytism. With the comparatively rapid

succession of sexually produced generations, occasional cross-fertilization would

suffice for maintaining genetic variability.

In this connexion it is, I think, significant that so very few natural hybrids of

epiphytic fern species have been reported, in contrast to the many terrestrial hybrids

known. This may be due in part to the physical inability of the researcher to

sample populations of epiphytes in the same way as of terrestrial or epilithic

species, but it can hardly account for the fact as a whole. The regular development

of single, self-fertilized prothalli may well be the reason for a greatly reduced

chance for interspecific cross-fertilization.

The phenomenon of coherence of spores by means of cohering or interlocking

surface structures may be called “‘synaptospory’’, analogous to what has been called

“synaptospermy”’ in seed plants, after Murbeck (see, for instance, Zohary, 1962,

p. 180, and van der Pijl, 1969, p. 77). Further research will have to show whether it

exists as a widespread phenomenon.

One general feature observed in terrestrial heterosporous lycopodiophytes

seems to point in the same direction. The great majority of Selaginella species, and

many species of the partly terrestrial genus Jsoetes, have strongly sculptured spores.

For illustrations, see, for instance, Knox (1950) and Hellwig (1969). For sexual

reproduction these plants are, of course, entirely dependent on germination of

spores producing gametes of opposite sex in very close proximity. The surface

sculpture may here, too, serve to enhance the chance for this to occur.

Equisetum is probably another case in point. The perispore here dissolves into

hygroscopically moving bands, somewhat unfortunately termed elaters, which loosen

the spore mass but also become entangled, leading to distribution of spores in groups.

The prothalli are at least very often functionally unisexual (Sporne, 1952, p. 112;

Duckett & Duckett, 1974).

I am indebted to Mr. E. Hennipman, Leiden, for stimulating discussions.

82 Gardens’ Bulletin, Singapore — XXX (1977)

References

Ching, R. C. 1966. Gymnogrammitidaceae Ching, a new fern family. Acta Phytotax.

Sin. 11: 11-16.

Duckett, J. G., & Duckett, A. R. 1974. The ecology of Equisetum gametophytes.

Am. J. Bot. 61: 36 [brief report).

Gastony, G. J. 1974. Spore morphology in the Cyatheaceae, I. The perine and

sporangial capacity: general considerations. Am. J. Bot, 61: 672-680.

Hellwig, R. L. 1969. Spores of the heterophyllous Selaginellae of Mexico and

Central America. Ann. Missouri Bot. G. 56: 444-464.

Johansson, D. 1974. Ecology of vascular epiphytes in West African rain forest.

Acta Phytogeogr. Suec. 59: 1-129.

Knox, E, M. 1950. The spores of Lycopodium, Phylloglossum, Selaginella and

Isoetes, and their value in the study of microfossils of Paleozoic age. Trans.

Bot. Soc. Edinb. 35: 211-357; 12 plates.

Kramer, K. U. 1968. Pteridophyta, in: J. Lanjouw & collab. Compendium van de

Pteridophyta en Spermatophyta. Oosthoek, Utrecht. 342 pp.

Lloyd, R. M. 1974, Genetic and mating system studies in ferns: systematic and

evolutionary implications, Internat. Organiz. Plant Biosyst. Newsletter 9: 2-14.

Lugardon, B. 1969. Sur la structure fine des parois sporales d’Equisetum maximum

Lamk. Pollen et Spores 11: 449-474.

1971. Contribution a la connaissance de la morphogénése et de la

structure des parois sporales chez les Filicinées isosporées. Thése, Université

de Toulouse. 257 pp., 51 plates.

1972a. La structure fine de l’exospore et de la périspore des

Filicinées isosporées, I — Généralités. Eusporangiées et Osmundales. Pollen

et Spores 14: 227-261.

1972b. Sur la structure fine et la nomenclature des parois micro-

sporales chez Selaginella denticulata (L.) Link et S. selaginoides (L.) Link.

C. R. Acad. Sci. Paris Sér. D 274: 1656-1659.

1973, Nomenclature et structure fine des parois acétorésistantes des

microspores d’Isoetes. C. R. Acad. Sci. Paris Sér. D 276: 3017-3020.

1974, La structure fine de l’exospore et de la périspore des Filicinées

isosporées. II. Filicales. Commentaires. Pollen et Spores 16: 161-226.

Naf, U. 1961. Mode of action of an antheridium-inducing substance in ferns. Nature

189: 900-903.

Pichi Sermolli, R. E. G. 1968. Adumbratio Florae Aethiopicae 15. Elaphoglossaceae.

Webbia 23: 209-246.

van der Pijl, L. 1969. Principles of dispersal in higher plants. Springer-Verlag,

Berlin. 154 pp.

Schneller, J. 1975. Untersuchungen an einheimischen Farnen, insbesondere der

Dryopteris filix-mas-Gruppe. 3. Teil. Oekologische Untersuchungen. Ber.

Schweiz. Bot. Ges. 85: 110-159.

Sen, T., & Sen, U. 1971. Morphology and anatomy of the fern genus Gymnogram-

mitis. Ann. Bot. 35: 229-235.

Synaptospory 83

de la Sota, E. R. 1971. El epifitismo y las pteriddfitas en Costa Rica (América

Central). Nova Hedwigia 21: 401-465.

Sporne, K. R. 1962. The morphology of pteridophytes. Hutchinson, London. 192 pp.

Tutin, T. G. 1964. Elaphoglossaceae, in: T. G. Tutin, V. H. Heywood & collab.,

Flora Europaea Vol, 1. Cambridge. xxxii + 464 pp.

Voeller, B. 1971. Developmental physiology of fern gametophytes: relevance for

biology. BioScience 21: 266-270.

Wagner, W. H. Jr. 1974. Structure of spores in relation to fern phylogeny. Ann.

Missouri Bot. G. 61: 332-353.

Zohary, M. 1962. Plant life of Palestine. Ronald Press, New York. 262 pp.

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Spore Morphology of Malayan Dennstaedtiaceae sensu Holttum

E. SoEPADMO & E. E. KHoo

Department of Botany

University of Malaya

Kuala Lumpur

Abstract

Spore morphology of 28 species belonging to 28 genera included by Holttum (1954) in

the Dennstaedtiaceae are described. Amongst the Malaysian taxa there are at least seven

distinct spore-types. The disposition of the Malaysian genera in these spore-types does not

conform with any of the existing systems of classification. Only spores of those genera

assigned by Holttum in the subfamilies Davallioideae, Dryopteridoideae and Tectarioideae

show some degree of uniformity.

Introduction

The Dennstaedtiaceae as originally defined by Holttum (1947, 1949, 1954)

is a very large family of modern ferns, containing very diverse groups of genera

and species. In the Peninsular Malaysia alone the family comprises not less than

48 genera and 203 species out of the total 124 genera and 389 species of its fern-

flora. As has been shown in the excellent review by Pichi Sermolli (1973) and

Mickel (1973), there seems to be a great controversy over the delimitation and

classification of the taxa involved. Many authors, such as Christensen (1938),

Ching (1940), Copeland (1947), Reimers (1954), Pichi Sermolli (1958), Nayar

(1970) and others placed the 48 Malaysian genera involved in 5 or 10 separate

families, whereas Holttum (1954) included these genera in 11 subfamilies of the

Dennstaedtiaceae, though later (1973) he more or less agrees with Pichi Sermolli’s

system of classification. Furthermore, it is also evident that there is a great

difference of opinion with regard to the disposition and affinity of the genera

within each family or subfamily recognised by the above authors.

In an attempt to introduce a more natural system of classification of these

taxa, many pteridologists (e.g. Atkinson, 1973; Lovis, 1973; Manton, 1961; Mickel,

1973; Sen & Sen, 1973; Sledge, 1973; Swain & Cooper-Driver, 1973; Van Cotthem,

1973; Walker, 1973, and others) have used modern and advanced techniques in

their studies, previously unexploited by those who are responsible in drawing up

the existing schemes of classification. Amongst these modern techniques are:

anatomical, cytological, phytochemical, biochemical, developmental and palyno-

logical.

Of the palynological aspects, Erdtman & Sorsa (1971) have admirably

summarised our present knowledge on the spore morphology of those genera

included in the Dennstaedtiaceae by Holttum. This work shows that of the total

3052 species so far recognised in the 48 genera occurring in Malaysia (Willis,

1973), spore descriptions of only 449 (= 14.7%) have been published by Harris

(1955), Erdtman (1957), Nayar & Devi (1963, 1964a, 1964b, 1966, 1967,

1968a, 1968b), Nayar & Kaur (1963), Tardieu-Blot (1963a & b, 1965), and

several others. Of these descriptions only a few were based on Malaysian specimens.

86 Gardens’ Bulletin, Singapore — XXX (1977)

The present paper describes the spores of 28 species belonging to 28 genera,

and is intended as a starting point for more intensive works to be done in the

future. |

Species Investigated

1. DENNSTAEDTIOIDEAE 6. ASPLENIOIDEAE

Hypolepis bivalvis v.A.v.R. Asplenium nidus L. var. musifolia

Microlepia speluncae (L.). Moore (Sm.) Bedd.

Orthiopteris kingii (Bedd.) Holtt. gv igi comes

2. LINDSAEOIDEAE Blechnum orientale L.

Isoloma divergens (Hook. f. & Grev.) Brainea insignis (Hook. f.) Sm.

Sm.

8. LOMARIOPSIDOIDEAE

Bolbitis heteroclita (Pr.) Ching

Egenolfia appendiculata (Willd.) Sm.

Elaphoglossum callifolium (Bl.) Moore

Lindsaea scandens Hook. f.

Sphenomeris chusana (L.) Copel.

Tapeinidium pinnatum (Cav.) Christ.

3. DAVALLIOIDEAE Lomagramma sumatrana v.A.V.R.

Davallia divaricata Bl. Teratophyllum aculeatum (Bl.) Mett.

Humata heterophylla (Sm.) Desv. var. montanum Holtt.

4. OLEANDROIDEAE 9. DRYOPTERIDOIDEAE

Nephrolepis biserrata (Sw.) Schott. Dryopteris sparsa (Don) O. Ktze.

Oleandra pistillaris (Sw.) Christ. 10. TECTARIOIDEAE

5. PTERIDIOIDEAE Arcypteris irregularis (Pr.) Holtt.

Acrostichum aureum L. Heterogonium pinnatum (Copel.)

Histiopteris stipulacea (Hook. f.) Holtt.

Copel. 11. ATHYRIOIDEAE

Pieris vittata L.

Athyrium pinnatum (Blanco) Copel.,

Stenochlaena palustris (Burm.) Bedd.

Cystopteris tenuisecta (Bl.) Mett.

Materials and Methods

In this study both fresh and herbarium materials were used. The fresh

specimens were collected from various localities in Selangor and Pahang, and were

identified with the help of Holttum’s keys (1954). Voucher specimens are now

deposited in the Herbarium, Department of Botany. University of Malaya, Kuala

Lumpur (KLU). Dried and preserved materials were obtained from herbarium

specimens available in the above Herbarium.

Spores containing materials were acetolysed, stained lightly with safranin, and

mounted in glycerin jelly. The slides were then sealed permanently with paraffin

wax. In some species, during acetolysis the perine was easily dissolved and the

spores become shrivelled. In such cases, a lesser amount of concentrated sulphuric

acid was added in the acetolysing mixture and heating was skipped. Although by

this treatment, the spores did not become transparent, the perine remained intact

and could be studied. Observation, measurement and photomicrography was

carried out with a Leitz Dialux Microscope fitted with occular micrometer,

Combiphot Automatic Camera System, achromatic condensor with N.A, 0.90,

phaco achromatic objectives 40x /0.65 and 100X /1.30, and halogen lamp-housing

model 100 Z.

For each species, measurement is based on 50 readings. This measurement is

expressed in the text as length of polar axis (P) x longitudinal equatorial diameter

(El) X transverse equatorial diameter (E2) in monolete-bilateral spores and as

P x E1l in trilete-tetrahédral spores. All measurements are exclusive of perine and

any other processes. :

Malayan Dennstaedtiaceae | 87

Description of Spores

DENNSTAEDTIOIDEAE

1. Hypolepis bivalvis (Gunong Brinchang, Perak; Poore KLU 468; Plate 8,

Figs. 4-7).

Spores monolete-bilateral, 30-46 x 48-61 x 33-46 ». Amb oblong; proximal

face flat, distal face hemispherical. Laesura 36-40 yp, margins thickened into

protruding lips about 4 to 7 » broad on either side of the laesura. Exine

2 p» thick, sexine thicker than nexine, with fine spinulose pattern. Perine thin and

provided with minute spine-like processes.

2. Orthiopteris kingii (Cameron Highlands, Pahang; Turnau KLU 2936; Plate 2,

Fig 8).

Spores trilete-tetrahedral, 29-32 x 31-39 ». Amb triangular in outline, with

convex sides and broadly rounded corners. Proximal face conical, distal face

convex to hemispherical. Laesura 15 yw, margins thickened into lips about

3 » broad and tapering towards the tips. Exine 2 » thick, sexine nearly as thick

as nexine, smooth. Perine absent.

3. Méicrolepia speluncae (Templer Park, Selangor; Abd. Samat b, Abdullah KLU

2686; Plate 2, Figs. 6-7).

Spores trilete-tetrahedral, 25-33 x 30-33 yw. Amb triangular with strongly

concave sides and broadly rounded corners, Proximal face slightly convex to

flat; distal face hemispherical. Laesura 13 y», margin thickened into lips of

2 » broad and tapering towards the tips. Exine 2 » thick, sexine as thick as nexine,

densely granulose. Perine absent.

LINDSAEOIDEAE

4. Lindsaea scandens (Gunong Ulu Kali, Pahang; Abd. Samat b. Abdullah KLU

1846; Plate 2, Figs. 2-3).

Spores trilete-tetrahedral, 22-25 x 24-27 uu. Amb triangular, with slightly

convex to straight sides. Proximal face conical with angular pole; distal face

convex. Laesura 13 », margins thickened into lips of 1 yw broad. Exine

1 uw thick, not clearly subdivided into sexine and nexine, densely granulose. Perine

absent,

5. Sphenomeris chusana (Fraser’s Hills, Pahang; Turnau KLU 2701; Plate 9,

Fig. 3).

Spores monolete-bilateral, 43-63 x 65-76 x 40-51 u. Amb oblong. Proximal

face flat; distal face hemispherical. Laesura 51 u, margins thickened into protruding

lips of 8 » broad. Exine 2 » thick, sexine much thicker than nexine, smooth.

Perine smooth and thin, partially peeling off on acetolysis.

6. Isoloma divergens (Genting Simpah, Selangor; Mohd. Kassim b. Rajab KLU

139; Plate 2, Figs. 4-5).

Spores trilete-tetrahedral, 19-22 x 20-25 uw. Amb triangular with straight to

slightly convex sides and broadly rounded corners. Proximal face conical with

angular pole; distal face convex. Laesura 12 », margins thickened into lips of

1 » broad. Exine 1.5 » thick, sexine much thicker than nexine, smooth. Perine-like

layer present, smooth, very closely adherent to the exine, and disintegrates very

easily on acetolysis.

7. Tapeinidium pinnatum (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12307;

Plate 9, Fig. 4).

88 Gardens’ Bulletin, Singapore — XXX (1977)

Spores monolete-bilateral, 26-32 x 28-46 x 27-32 ». Amb. oblong. Proximal

face flat to slightly convex; distal face hemispherical. Laesura 32 y», margins

thickened into protruding lips of 6 » broad. Exine 1.5 » thick, sexine thicker

than nexine, smooth. Perine thin with fine reticulate pattern, disintegrates easily

on acetolysis.

DAVALLIOIDEAE

8. Davallia divaricata (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12309; Plate

6, Figs. 1-3).

Spores monolete-bilateral, 27-35 x 36-50 x 24-34 ». Amb oblong. Proximal

face concave; distal face convex. Laesura 28 », margins slightly thickened into lips

of 2 » broad. Exine 2.5 » thick, sexine thicker than nexine, verrucose to rounded

tuberculose; verrucae or tubercles crowded, 5-7 p» in diameter and 2 uw» high.

Perine absent.

9. Humata heterophylla (Lombong, Johore; Abd. Samat b. Abdullah KLU 1849;

Plate 6, Figs. 4-8).

Spores monolete-bilateral, 22-36 <x 38-43 x 28-32 ». Amb oblong. Proximal

face flat; distal face hemispherical. Laesura 18 », margins slightly thickened into

lips of 1 » broad. Exine 3 » thick, sexine thicker than nexine, verrucose-tuber-

culose; tubercles crowded, round, 2 » high, 5-7 » broad and 10-16 wp long.

Perine absent.

OLEANDROIDEAE

10. Oleandra pistillaris (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12310;

Plate 4, Figs. 5-6).

Spores monolete-bilateral, 19-26 x 27-32 x 17-23 ». Amb oblong. Proximal

face flat; distal face hemispherical. Laesura 17 », margins slightly thickened into

lips of 1 » broad. Exine 4-5 yp» thick, sexine thicker than nexine, smooth. Perine

loosely wrinkled, anastomosing to form a reticulate patterns; folds about 4 » high.

spinulose; spinules minutes, less than 1 p» high, with pointed tips. Perine disinte-

grates very easily on acetolysis.

11. Nephrolepis biserrata (Damansara Road, Kuala Lumpur, Selangor; Khoo Eng

Ee KLU 12305; Plate 7, Figs. 3-4).

Spores monolete-bilateral, 25-30 x 33-40 x 23-29 ». Amb oblong. Proximal

face flat to slightly concave; distal face convex. Laesura 20 u, margins thickened

into lips of 1 » broad, tapering towards the tips. Exine 2 » thick, sexine thicker

than nexine, verrucose to tuberculose; verrucae 1 p» high and 46 y» broad.

Perine thin, more or less verrucose and easily disintegrates on acetolysis.

PTERIDIOIDEAE

12. Histiopteris stipulacea (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12306;

Plate 7, Figs. 1-2).

Spores monolete-bilateral, 38-48 x 59-65 x 43-46 ». Amb oblong. Proximal

face slightly concave; distal face convex. Laesura 40 p, margins thickened slightly

into lips of 6-10 » broad and tapering towards the tips. Exine 5 p» thick, sexine

thicker than nexine, densely verrucose-tuberculose; verrucae or tubercles granulose,

5 p» high, 6 » broad, and 8-22 » long. Perine absent.

13. Pteris vittata (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12306; Plate 1,

Figs. 1-3).

Spores trilete-tetrahedral, 48-53 x 61-65 yw. Amb triangular with nearly

straight sides and rounded corners. Proximal face flat to slightly convex; distal

Malayan Dennstaedtiaceae 89

face hemispherical. Laesura 27 », margins thickened into lips of 3 « broad and

tapering towards the tips. Exine 3 « thick, sexine much thicker than nexine, densely

rugulose; rugulae up to 6 u tall, sometimes coalescing to form a coarse reticulum.

On the proximal face there are 3 ridges parallel and close to the laesura arms.

Equator collar present; this is a collar-like ridge which protrudes as a flange

around the spores, separating the proximal and the distal parts of the spores, up

to 7 » high, often interrupted. Perine absent.

14. Acrostichum aureum (Lukut, Negeri Sembilan; Turnau KLU 2925: Plate 2,

Fig. 1).

Spores trilete-tetrahedral, 42-49 x 50-60 «. Amb rounded triangular with

slightly convex sides and broadly rounded corners. Proximal face conical with

straight sides and angular pole; distal face hemispherical. Laesura 21 «, margins

thickened into lips of 1 « broad and tapering towards the tips. Exine 2 « thick,

sexine thicker than nexine, granulose. Perine not seen.

15. Stenochlaena palustris (Damansara Road, Kuala Lumpur, Selangor; Khoo

Eng Ee KLU 12311; Plate 7, Figs. 5-6).

Spores monolete-bilateral, 30-37 x 39-53 xX 30-36 x. Amb oblong. Proximal

face flat to slightly concave; distal face hemispherical. Laesura 29 xu, margins

thickened into lips of 1 « broad. Exine 3 uw thick, sexine thicker than nexine,

provided with conical tubercles of 2 » tall with rounded tips and base diameter

of 2 uw. Perine absent.

ASPLENIOIDEAE

16. Asplenium nidus var. musifolia (Damansara Road, Kuala Lumpur; Khoo Eng

Ee KLU 12304; Plate 3, Figs. 1-2).

Spores monolete-bilateral, 27-36 x 30-SO X 27-32 u. Amb oblong. Proximal

face flat to slightly concave; distal face hemispherica]. Laesura 20 xu, margins

thickened slightly into lips of 3 u broad. Exine 1 w thick, sexine as thick as

nexine, smooth. Perine finely granulose, wrinkled into discontinous, closely sinuous

folds protruding up to 3 » from the exine surface, with smooth crests.

BLECHNOIDEAE

17. Blechnum orientale (Ulu Gombak Road, Selangor; Khoo Eng Ee KLU

12308; Plate 3, Figs. 3-4).

Spores monolete-bilateral, 34-39 x 44-51 X 3646 u. Amb oblong. Proximal

face flat; distal face hemispherical. Laesura 29 », margins thickened into lips

3 » broad and tapering towards the tips. Exine 1 « thick, sexine as thick as

nexine, smooth. Perine sparsely granulose, loosely folded into sparse, thin indistinct

folds protruding up to 3 u« from the exine surface, with smooth crests: disintegrates

easily on acetolysis.

18. Brainea insignis (Tanjong Selantai, Johore; Abd. Samat b. Abdullah KLU

1869; Plate 10, Figs. 4-6).

Spores monolete-bilateral, 31-42 x 35-SO X 29 43 x. Amb oblong to sub-

spherical. Proximal face slightly convex; distal face hemispherical. Laesura 27 x,

margins thickened into lips of 1 « thick and tapering towards the tips. Exine

2 m thick, sexine as thick as nexine, smooth. Perine densely granulose, closely

adherent to the exine nearly without folds or only with one or two indistinct short

folds which protrude up to 2 « from the exine surface; disintegrates easily on

acetolysis.

90 Gardens’ Bulletin, Singapore — XXX (1977)

LOMARIOPSIDOIDEAE

19. Egenolfia appendiculata (Kuah, Langkawi, Kedah; Abd. Samat b. Abdullah

KLU 2316; Plate 10, Fig. 3).

Spores monolete-bilateral, 32-36 x 31-39 x 26-37 ». Amb subspherical to

circular in outline. Proximal face flat to slightly convex; distal face hemispherical.

Laesura 19 yp, slightly thickened into lips of 1 » broad; in some spores forked in

one end. Exine 1 » thick, sexine as thick as nexine, smooth. Perine with finely

reticulate patterns, loose, lightly folded into very many elongated sharp ridges most

of which radiate from the centre to the periphery of the spores and protruding up

to 7 » from the exine surface; disintegrates on acetolysis.

20. Bolbitis heteroclita (Jengka Forest Reserve, Pahang; Turnau KLU 2894;

Plate 10, Figs. 1-2).

Spores monolete-bilateral, 31-37 x 35-42 x 35-38 yw. Amb oblong to

subspherical. Proximal face flat to convex: distal face hemispherical. Laesura 21 uy,

margins slightly thickened into smooth lips of 1 » broad. Exine 1 yw» thick, sexine

as thick as nexine, smooth. Perine sparcely covered with minute spinules, highly

folded into sinuous and elongated ridges radiating from the centre to the periphery

of the spores; folds protrude up to 12 u from the exine surface, with undulating

crests; disintegrates on acetolysis.

21. Teratophyllum aculeatum var. montanum (Tringkap, Pahang Abd. Samat b.

Abdullah KLU 2288; Plate 5, Figs. 1-3).

Spores monolete-bilateral, 47-60 Xx 66-71 uw (polar view not obtained).

Proximal face convex to hemispherical; distal face hemispherical. Laesura 27-30 up.

Exine very thin, less than 1 » thick, not subdivided into distinct sexine and nexine.

Perine densely crowded with minute spinules less than 1 » high, very sparsely

folded into faint folds which protrude up to 19 » from the exine surface; with

smooth crests; cracked up on acetolysis,

22. Lomagramma sumatrana (22nd mile Bentong Road, Selangor; Turnau KLU

2914; Plate 9, Figs. 1-2).

Spores monolete-bilateral, 31-43 x 50-58 x 35-40 ». Amb oblong. Proximal

face flat to slightly concave; distal face hemispherical. Laesura 29 ,, margins

slightly thickened into lips of 1 » broad. Exine 2 » thick, sexine slightly thinner

than nexine, finely granulose to spinulose. Perine absent.

23. Elaphoglossum callifolium (Tanah Rata, Pahang; Mahmud b. Sider KLU

4612; Plate 4, Figs. 1-2).

Spores monolete-bilateral, 18-30 x 23-40 x 18-22 ». Amb oblong. Proximal

face flat to slightly concave; distal face hemispherical. Laesura 18 yw. Exine

1 » thick, sexine and nexine are not clearly differentiated, smooth. Perine minutely

spinulose, sparsely folded into thin, sharp sinuous ridges protruding up to 7 » from

the exine surface; disintegrates on acetolysis.

DRYOPTERIDOIDEAE

24. Dryopteris sparsa (Maxwell Hills, Perak; Evans KLU 2493; Plate 3,

Figs. 5-6). .

Spores monolete-bilateral, 22-27 x 38-46 x 28-31 ». Amb oblong. Proximal

face flat to slightly concave; distal face convex. Laesura 20 », margins thickened

very slightly into lips of 1 » broad. Exine 1 ,» thick, sexine as thick as nexine,

smooth. Perine smooth, fairly closely adherent to the exine, folded into lobate

crowded ridges protruding up to 11 » from the exine surface; with many undulating

crests; disintegrates on acetolysis.

Malayan Dennstaedtiaceae 9]

TECTARIOIDEAE

25. Arcypteris irregularis (Ulu Gombak Road, Selangor; Wee & Lim KLU 1975;

Plate 5, Figs. 4-6).

Spores monolete-bilateral, 20-29 x 32-39 x 23-26 ». Amb oblong. Proximal

face flat; distal face hemispherical. Laesura 20 », margins faintly thickened into

lips of 1 » broad. Exine 1.5 » thick, sexine as thick as nexine, smooth. Perine with

very minute spinules, folded into a few elongated sharp, thin ridges radiating from

the centre to the periphery of the spores; folds protrude up to 10 » from the exine

surface; disintegrates easily on acetolysis,

26. Heterogonium pinnatum (Templer Park, Selangor; Abd. Samat b. Abdullah

KLU 2636; Plate 4, Figs. 3-4).

Spores monolete-bilateral, 283-40 x 40-50 x 33-36 u. Amb oblong. Proximal

face flat to slightly concave; distal face hemispherical. Laesura 27 uw, margins

faintly thickened into lips of 1 » broad. Exine 2 wu thick, sexine as thick as

nexine, smooth, breaking up on acetolysis. Perine minutely spinulose (spines less

than 1 » tall), highly folded up into elongated, closely sinuous ridges protruding

up to 6 » from the exine surface; disintegrates on acetolysis.

ATHYRIOIDEAE

27. Athyrium pinnatum (Genting Simpah, Selangor; Turnau KLU 2785; Plate

9, Figs. 5-6).

Spores monolete-bilateral, 20-27 x 33-44 x 24-25 uw. Amb oblong. Proximal

face flat to convex; distal face hemispherical. Laesura 23 yw, margins faintly

thickened into lips of 1 » broad. Exine 2 yw thick, sexine as thick as nexine,

smooth. Perine faintly granulose, closely adherent to the exine, with a few folds;

folds appear as irregular ridges protruding up to 2 u» from the exine surface;

disintegrates on acetolysis.

28. Cystopteris tenuisecta (Cameron Highlands, Pahang: Poore KLU 256; Plate

8, Figs 1-3).

Spores monolete-bilateral, 30-45 x 45-60 x 31-45 u. Amb oblong. Proximal

face flat to slightly concave; distal face hemispherical. Laesura 30 yu, margins

slightly thickened into lips of 1 « broad. Exine 2 ,» thick, sexine thicker than

nexine, densely finely spinulose. Perine folded; folds anastomose to form dense

reticulum, provided with numerous rod-like protuberances about 3-4 .» high; tips

of rods peltately lobed.

Discussion

Spore morphology of 28 Malaysian species included by Holttum (1954) in

the Dennstaedtiaceae shows a great diversity of form, size and structure. Based

on whether the spores are monolete-bilateral or trilete-tetrahedral, with or without

perine, and perine if present smooth or variously patterned, etc., there seem to be

at least seven distinct types of spores. These spore-types are:

Type-1: Spore trilete-tetrahedral, provided with well-developed equatorial collar

(Pteris vittata).

Type-2: Spore trilete-tetrahedral, without equatorial collar, without or occasionally

with a thin spinulose perine (Acrostichum aureum, Isoloma divergens,

Lindsaea scandens, Microlepia speluncae, and Orthiopteris kingii).

Type-3: Spore monolete-bilateral, provided with folded or reticulate perine, plano-

convex or concave-convex in equatorial view (Asplinium nidus var.

musifolia, Blechnum orientale, Dryopteris sparsa, Elaphoglossum calli-

folium, Heterogonium pinnatum, Oleandra pistillaris, Arcypteris irregu-

laris, and Teratophyllum aculeatum).

92 Gardens’ Bulletin, Singapore — XXX (1977)

Type-4: Spore monolete-bilateral, without or occasionally with very thin perine,

exine provided with low, flattish, roundish or conical protuberances

(Davallia divaricata, Histiopteris stipulacea, Humata heterophylla,

Nephrolepis biserrata, and Stenochlaena palustris).

Type-5: Spore monolete-bilateral, plano-convex, perine provided with spine-like

or rod-like blunt processes (Cystopteris tenuisecta, Hypolepis bivalvis).

Type-6: Spore monolete-bilateral, concave-convex or plano-convex, with or with-

out perine; perine if present thin, hardly folded and easily dissolved on

acetolysis (Athyrium pinnatum, Lomagramma sumatrana, Sphenomeris

chusana, and Tapeinidium pinnatum).

Type-7: Spore monolete-bilateral, biconvex or sometimes plano-convex, provided

with folded, reticulate or finely patterned perine (Bolbitis heteroclita,

Brainea insignis, and Egenolfia appendiculata).

In the Dennstaedtioideae, Holttum included Dennstaedtia, Hypolepis, Micro-

lepia and. Orthiopteris. Hypolepis with a monolete-bilateral spore of type-5 seems

to be out of place in this subfamily, since the other three genera have trilete-

tetrahedral spores of type-2. Furthermore, the perine in Hypolepis is provided with

spine-like or rod-like minute processes, while the perine of Dennstaedtia and

Orthiopteris, if present, is not so patterned, and in Microlepia the spores are with-

out perine. Erdtman & Sorsa (1971) included these four genera in the family

Pteridaceae in which 52 out of 65 genera have exclusively trilete-tetrahedral

spores, 7 possess mainly trilete but occasionally also monolete spores, and 6

genera with exclusively monolete-bilateral spores. Apart from Hypolepis, the other

five genera which have exclusively monolete-bilateral spores are:— Humblotiella,

Tapeinidium, Sphenomeris, Paesia, and Lonchitis. Perine of these genera, if present,

is very thin, hardly folded and is easily detached from the exine on acetolysis,

unlike that of Hypolepis. In Holttum’s classification, Tapeinidium and Sphenomeris,

on the other hand, were grouped together with Lindsaea, Schizoloma and Isoloma

in the Lindsaeoideae. Spore morphology does not support this grouping, since

both Isoloma and Lindsaea have a trilete-tetrahedral spore of type-2, whereas

Sphenomeris and Tapeinidium have a monolete-bilateral spore of type-6.

Of the Davallioideae, the four genera (i.e. Araiostegia, Davallia, Humata,

and Leucostegia) assigned by Holttum (1954) in this subfamily have a more or

less similar spore-type, namely monolete-bilateral, without perine, and the exine

is provided with low flattish verrucae or tubercles (spore type-4). For these

features, the spores of Nephrolepis, Histiopteris and Stenochlaena agree very well.

Holttum (1954), on the other hand, placed Nephrolepis together with Oleandra in

the Oleandroideae. Spores of Oleandra are, however, different from those of

Nephrolepis by being provided with folded perine, the detailed structure of which

matches very well with that of Elaphoglossum, Arcypteris, and Teratophyllum (spore

type-3). Histiopteris and Stenochlaena together with Acrostichum, Pteridium and

Pteris were included by Holttum (l.c.) in the Pteridioideae. Acrostichum and

Pteridium possess a trilete-tetrahedral spores of type-2, whereas Pteris has spores

of type-1.

The genus Asplenium (included by Holttum in the Asplenioideae) has spores

which are very much similar to those of Blechnum and Heterogonium (spore

type-3). However, Blechnum was assigned by Holttum in the Blechnoideae together

with Brainea and Woodwardia. While spores of Woodwardia match very well with

those of Blechnum, that of Brainea being monolete-bilateral, biconvex and provided

with folded, granulose-reticulose perine, agree well with the spores of Bolbitis and

Egenolfia (spore type-7). The last two mentioned genera were placed together

with Elaphoglossum, Lomagramma, Lomariopsis and Teratophyllum in the

Lomariopsidoideae by Holttum. While spores of Elaphoglossum and Teratophyllum

Malayan Dennstaedtiaceae 93

are very much similar to that of Oleandra and Tectaria (spore type-3), those of

Lomagramma, being without perine, agree well with the spores of Athyrium,

Sphenomeris and Tapeinidium (spore type-6). According to Erdtman & Sorsa

(1971), some species of Lomariopsis possess perinous spores similar to

those described as type-3, while others have a spore of type-6.

In the Dryopteridoideae, Holttum assigned Acrophorus, Diacalpe, Didymo-

chlaena, Dryopteris, Polystichum and Polystichopsis. These genera have more or

less uniform spores which match very well with those of Asplenium, Blechnum

and Heterogonium (Spore type-3). Similarly, the spores of ten genera included

by Holttum in the Tectarioideae (Arcypteris, Ctenitis, Cyclopeltis, Heterogonium,

Hypodematium, Lastreopsis, Pleocnemia, Pteridrys, Quercifilix and Tectaria)

agree well with each other and more or less similar to those of spore type-3.

Of the Athyroideae, Holttum place two Malaysian genera, namely Athyrium

and Cystopteris. As has been indicated above, the spores of Athyrium belong to

spore type-3, whereas those of Cystopteris to spore type-5.

The above discussion seems to indicate that apart from the Davallioideae,

Dryopteridoideae and Tectarioideae, the other subfamilies of the Dennstaedtiaceae

as defined by Holttum, show very diverse types of spores and that spore morpho-

logy does not lend any support to the disposition of various genera either in the

different subfamilies as recognised by Holttum (1954) or in the various families as

subscribed by Copeland (1947), Reimers (1954), Pichi Sermolli (1958) and

others.

However, this does not imply that the classification of the genera included by

Holttum in the Dennstaedtiaceae should match well with the spore types, but

rather that it demonstrates that more palynological studies, especially on those of

the tropical taxa should be done, so that morphological characters of the spores

could be considered along with other attributes in constructing a more natural

system of classification.

Finally, it should be emphasised here that, since the number of species of

which the spores have been studied in detail, is very small, and that, it is also well

known that even within a single genus the spore morphology may vary conside-

rably, the above findings should be considered with great caution.

Acknowledgement

The authors are very grateful to Mrs. Babe Foo, Messr. Mahmud b. Sider

and Michael Arokian for their technical help. We are also very much obliged to

Miss Amy Leong for typing the manuscript, and to Dr. Liew Fah Seong for going

through the manuscript and for his constructive criticisms.

References

Atkinson, L. R., 1973. The gametophyte and family relationships. In A. C. Jermy

et al (Ed.) — The phylogeny and classification of the ferns, 73-90.

Ching, R. C, 1940. On natural classification of the family Polypodiaceae.

Sunyatsenia 5, 201-268.

ee C., 1938. Filicinae. In F. Verdoorn (Ed.) — Manual of Pteridology.

22-550.

copeeesls E. B., 1947. Genera Filicum. Chronica Botanica. Waltham Mass..

Erdtman, G., 1957. Pollen and Spore Morphology/Plant Taxonomy — Ul. —

Gymnospermae, Pteridophyta, Bryophyta.

94 Gardens’ Bulletin, Singapore — XXX (1977)

—____—_— & P. Sorsa, 1971. Pollen and Spore Morphology/Plant Taxonomy

— IV. — Pteridophyta.

Harris, W. F., 1955. A manual of the spores of New Zealand Pteridophyta. N.Z.

Dept. Sci. Industr. Res. Bull, 116, 186 pp. Willington.

Holttum, R. E. 1947. A revised classification of Leptosporangiate ferns. J. Linn.

Soc. (Bot.) Lond. 52, 123-158.

1949. The classification of ferns. Biol, Rey, 24, 267-295.

1954. A Revised Flora of Malaya — II. Ferns of Malaya, 299-574.

Govt. Printing Office Singapore.

1973. Introduction to Pteridophyta. In J. C. Willis (Revised by

H. K. Airy Shaw) Dict. FI. Pl. & Ferns. 8th. edition. Cambridge University

Press.

Lovis, J. D., 1973. A biosystematic approach to phylogenetic problems and its

application to the Aspleniaceae. In A. C. Jermy et al (Ed.) — The phylogeny

and classification of the ferns, 211-227.

Manton, [., 1961. Evolution in the Pteridophyta. In P. J. Wanstall (Ed.) — A

Darwin Centenary Bot. Soc. Brit. Isle Conf. Rep. 6, 105-119

Mickel, J. T., 1973. The classification and phylogenetic position of the Dennstaed-

tiaceae. In A. C. Jermy et al (Ed.) — The phylogeny and classification of

the ferns, 135-144.

Nayar, B. K., 1970. A phylogenetic classification of the homosporous ferns. Taxon

19, 229-236.

Nayar, B. K. & S. Devi, 1963. Spore morphology of some Japanese Aspidiaceae.

Pollen Spores 5, 354-372.

1964 a. Spore morphology of Indian ferns-I. Aspidiaceae. Grana

Palynol. 5, 80-120.

1964 b. Spore morphology of Indian ferns-II Aspleniaceae &

Blechnaceae. Grana Palynol. 5, 222-242.

1966. Spore morphology of the Pteridaceae-I. The Pteridoid ferns

Grana Palynol. 6, 476-497.

1967. Spore morphology of the Pteridaceae — II. The Gymno-

grammoid ferns. Grana Palynol. 7, 568-600.

1968a. Spore morphology of the Pteridaceae. III. The Dicksonioid,

Dennstaedtioid and Lindsayoid ferns. Grana Palynol. 8, 185-203.

1968b. Spore morphology of the Pteridaceae. IV. Taxonomic and

phyletic considerations. Grana Palynol. 8, 517-535.

Nayar B. K. & S, Kaur, 1963. Spore morphology of some Indian members of the

Lomariopsidaceae. Pollen Spores 5, 87-94.

Pichi Sermolli, R. E. G., 1958. The higher taxa of the Pteridophyta and their

classification. Uppsala Univ. Arsskrift., 6, 70-90.

1973. Historical review of the higher classification of the Filicopsida.

In A, C. Jermy et al (Ed.) — The phylogeny and classification of the ferns,

11-40

Reimers, H., 1954. Pteridophytg. Farnpflanzen. In Engler’s Syllabus der Pflanzen-

familien, 12 Aufl. I, 269-311. Berlin.

Malayan Dennstaedtiaceae 95

Sen, U. & T. Sen, 1973. Anatomical relationships between the Oleandra and

Nephrolepis groups. In A. C. Jermy et al (Ed.) — The phylogeny and classifi-

cation of the ferns, 155-172.

Sledge, W. A., 1973. Generic and family boundaries in the Aspidiaceae and

Athyriaceae. In A. C. Jermy et al (Ed.) — The phylogeny and classification

of the ferns, 203-210.

Swain, T. & G. Cooper-Driver, 1973. Biochemical systematics in the Filicopsida.

In A. C. Jermy et al (Ed.) — The phylogeny and classification of the ferns,

111-134.

Tardieu-Blot, M. L., 1963a. Sur les spores de Lindsaeaceae et de Dennstaedtiaceae

de Madagascar et des Mascareignes, étude de palynologie appliquée a la

systematique. Pollen Spores 5, 69-86.

1963b. Sur les spores de Pterideae Malgaches. Pollen Spores §,

337-353.

1965. Sur les spores d’Adiantaceae, Aspleniaceae, Thelypteridaceae

et Athyriaceae de Madagascar. Pollen Spores 7, 319-338.

Van Cotthem, W. R. J., 1973. Stomatal types and systematics. In A. C. Jermy

et al (Ed.) — The phylogeny and classification of the ferns, 59-71.

Walker, T. G., 1973. Evidence from cytology in the classification of ferns. In A. C.

Jermy et al (Ed.) — The phylogeny and classification of the ferns, 91-110.

Willis, J. C., 1973. Dict. Fl. Pl. & Ferns. (Revised by H. K. Airy Shaw). 8th ed.

Cambridge Univ. Press.

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ne . .

aii OF perine tay

obl.e.v.; 3 — detailed structure of distal | :

(abbrev. in legends: pr.p.v. = proximal polar view, obl.e.v.

view, d = distal)

Plate 2. Fig. | — Acrostichum aureum, pr.p.v.; figs. 2 & 3 — Lindsaea scandens, pr.p.v. &

e.v. respectively; figs.-4 & 5: Isoloma divergens, 4 — pr.p.v., 5 — optical section of

4: fis. 6 & 7: Microlepia speluncae, 6 — pr.p.v., 7 — optical section of 6; fig,.8:-=

Orthiopteris kingii, optical section of pr.p.v.

2 — pr.p.v.;

Plate 3. Figs. 1 & 2: Asplenium nidus var. musifolia, | — e.v.,

Blechnum orientale, 3 — obl.pr.p.v., 4 — e.v. with broken perine;

Dryopteris sparsa, 5 — obl.pr.p.v., 6 — e.v.

Plate 4. Figs. 1-2: Elaphoglossum callifolium, 1 — e.v., 2 — d.p.v.; figs. 3 & 4: Heterogonium

pinnatum, 3 — left, e.v., right, d.p.v., 4 — pr.p.v.; figs. 5 & 6: Oleandra pistillaris,

d.p.v., 5 — focus on perine, 6 -— optical section.

Plate 5. Figs. 1-3: Teratophyllum aculeatum var. montanum, 1 — d.p.v. showing detailed

structure of perine, 2 — pr.p.v., 3 — obl.e.v.; figs. 4-6: Arcypteris irregularis,

4 — obl.pr.p.v., 5 — pr.p.v. in optical section, 6 — d.p.v.

Plate 6. Figs. 1-3: Davallia divaricata, 1 — pr.p.v. in optical section, 2 — detailed structure

of exine in pr.p.v., 3 — e.v. in optical section; figs. 4-8: Humata heterophylla,

4 — obl.e.v., 5 — pr.p.v., 6 — d.p.v., 7 — e.v., 8 — e.v. in optical section.

Plate 7. Figs. | & 2: Histiopteris stipulacea, | — e.v., 2 — obl.pr.p.v.: figs. 3 & 4 Nephrolepis

biserrata, 3 — pr.p.v., 4 — e.v.; 5 & 6: Stenochlaena palustris, 5 — pr-.p.v. in optical

section, 6 — e.v. in optical section.

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Plate 9. Figs. 1-2: Lomagramma sumatrana, 1 — e.v., 2 — obl.pr.p.v.; fig. 3 — Sphenomeris

chusana, obl.e.v.; fig. 4 — Tapeinidium pinnatum, obl.e.v.; figs. 5 & 6: Athyrium

pinnatum, 5 — obl.pr.p.v., 6 — e.v.

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An Enumeration of the Malesian Species of Aralia L.

W.R. PHILIPSON

Department of Botany

University of Canterbury, New Zealand

The species of Aralia in South-east Asia and the adjacent islands have been

subject to considerable misunderstanding. The present account takes a broad view

of specific limits, following along lines originally proposed by van Steenis (Bull.

Bot. Gard. Buitenz. ser. 3, 17 (1948) 391).

Aralia

Linn. Sp. Pl. (1753) 273. — Acanthophora Merr. Philip. J. Sc. 13 (1918) 316;

Steen. Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 390.

Sparingly branched shrubs or small trees, or climbing, rarely (extra Malesia)

herbaceous, glabrous or hairy, often prickly. Leaves pinnate to tri-pinnate, usually

with leaflets at the insertion of the lateral pinnae; leaflets serrate; petiole with a

sheathing base. Inflorescence a terminal panicle; flowers sessile or pedicellate, with

an articulation below the flower; calyx with 5—6 teeth; petals 5-6, imbricate; ovary

2-6 celled; styles 2-6 free or shortly connate below. Fruit a fleshy drupe; pyrenes

cartilaginous compressed; endosperm uniform.

Distr. More than 30 species in North America and Asia. In Malesia 6 species,

extending from Sumatra and the Malay Peninsula through Borneo and Celebes to

the Philippines and West New Guinea.

Eco. Usually on scrubby hillsides and in second growth, often in ravines or

near streams, or in thickets near or above the limit of tree-growth. Occurring at

low altitudes (100 m) but usually in the montane zone, up to 3000 m.

Notes. Hui-Lin Li in Sargentia 2 (1942) 101, treats some species that extend

into Malesia. Merrill considered that the climbing habit and recurved spines of

Acanthophora justified its separation as a distinct genus, but more recent authors

have not agreed.

KEY TO THE SPECIES

1. a. Flowers sessile (capitate) or very shortly pedicellate .....................:0.00 2

i rowers peaicellate iCumbellate) 45.00. Lh ocean ee eeees 3

2. a. Flowers sessile, underside of leaf + densely tomentose, hairs of the

branches and inflorescence + appressed and felted, bracts around the

PREM I TEN PAT oe. soaks. Tadeo eee ceca eens 1. A. dasyphylla

b. Flowers short pedicellate, underside of leaf sparsely tomentose, hairs of the

branches and inflorescence + patent, bracts around the capitula less densely

I Re ed i ov cciaa « siildnweu ag gh~s omansed dp adaeee 2. A. javanica

3. a. Climbing or scrambling liane, spines curved .................. 3. A. scandens

b. Erect shrubs or small trees, spines straight ................cccscseseceeeeeeeeeeees 4

98 Gardens’ Bulletin, Singapore — XXX (1977)

4. a. Leaflets glaucous beneath, margins with few crenations, fruit small (+

3 MM TOME) .....00.00s00svaveenenpussvaes sys peeudbnaie sen 4. A. bipinnata

b. Leaflets green (or with fawn pubescence) beneath, margins serrate, fruit

rather larger (4-6 mm long):).,6..5000)s.80100. nema ey 5

5. a. Young parts and under-surface of leaves glabrous (but with small spines)

stebeccesbocsnsdtwbanna sec see ekg hey te MehtOA te onkhs the niuics + aitinnntna an 5. A. ferox

b. Young parts and under-surface of leaves pubescent ............ 6. A. montana

1. Aralia dasyphylla Mig. Fl. Ind, Bat. 1, 1 (1856) 751. — A. urticifolia BI.

ex Mig. Ann, Mus. Bot, Lugd. -Bat. 1 (1863). — A. dasyphylla var. strigosa Miq.

Ann. Mus. Bot. -Lugd. — Bat. 1 (1863) 9. — A. dasyphylla var, latifolia Miq.

Ann, Mus. Bot. Lugd. -Bat. 1 (1863) 9. — A. beccarii Ridl. J. Mal. Br. R. As.

Soc. 87 (1923) 64. — A. dasyphylla var. urticifolia (Bl. ex Miq.) Bakh. in Blumea

6 (1947) 367. — A. dasyphylla var. typica Bakh, -Oostr, in Back. Bekn, Fl. Java,

(em. ed.) 7 (1948) fam. 159, 18. — A. dasyphylla var. dasyphylla Back. & Bakh.

Fl. Java, 2 (1965) 710.

Prickly shrub or small tree, often unbranched, to about 5 m high, young parts

densely brown pubescent. Leaves forming large rosettes at the summit of the stems,

about 1 m long (or more), bi- or tripinnate, the petiole, rhachis and lateral

rhachides prickly or unarmed, densely pubescent; leaflets subsessile or petiolule

c. 5 mm long (or longer), usually densely pubescent on the lower surface, less

dense above, ovate to oblong-ovate, c. 5-14 (18) x 3-5 (10) cm, base rounded to

sub-cordate, apex accuminate, margin finely or sometimes coarsely serrulate.

Inflorescence a large terminal panicle, densely brown pubescent, ultimate branches

arranged racemosely, ending in heads of several sessile flowers.

Distr. Malay Peninsula, Sumatra, West and Central Java. Also northwards to

southern China.

Eco. Primary forest and second growth in deep ravines or open hillsides,

occurring from low altitudes (+ 100 m) to 2,500 m.

Notes. The capitulate flowers are characteristic (see also under A. javanica).

The presence of this species in the Malay Peninsula has often been overlooked,

though it extends into southern China. Variability is discussed by van Steenis

(Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 391).

2. Aralia javanica Miq. Pl. Jungh. 3 (1855) 420.

A shrub or small tree, often unbranched, young parts covered with brown

pubescence which persists on the stems and rhachides of the inflorescence as +

patent hairs. Leaves tufted at the ends of the branches, bipinnate; leaflets variable

in size, subsessile or the petiolules up to c. 2 cm long, blade ovate to elliptic, up

to 18 X 8 cm (usually smaller), both surfaces with sparse short appressed bristly

hairs, base cuneate to truncate, apex accuminate, margin finely and unevenly

serrulate. Inflorescence a large terminal panicle; the ultimate branches bearing

heads (or sub-umbellules) of c. 10 flowers, surrounded by an involucre of small

linear bracts.

Distr. Central and West Java (Dieng, Surakarta, G. Papandayan, Mt Malabar,

G. Prahu). |

Eco.. Mountain forests, 2000-3000 m.

Notes. This imperfectly known species may prove to be a form of the wide-

spread A. dasyphylla, from which it appears to differ in the shortly pedicellate

flowers, the sparser leaf-tomentum, the more patent hairs on the inflorescence

branches and the bracts of the umbellules less thickly enveloped in hairs.

Malesian Aralia 99

3. Aralia scandens (Merr.) Ha in Nov. Sist. Vyssh. Rast. 11 (1974) 229. —

Acanthophora scandens Merr, in Philip. J. Sc. 13 (1918) 316. — ‘‘Aralia ferox

Mig.” King in J. As. Soc. Beng 67 (1898) 45.

Prickly scandent shrub, glabrous, reaching a height of 10 m or more, stems

c. 2.5 cm thick. Leaves dispersed (+ 30 cm apart), up to 1.5 m long, tri- or

quadri-pinnate, prickly on the petiole, rhachides, and sometimes on the leaf veins;

leaflets ovate to elliptic-ovate or ovate-lanceolate, base rounded or sub-cordate,

apex acuminate, margins finely spinulose-denticulate. Inflorescence a large terminal

spiny panicle, the main rhachis to c. 60 cm long, bearing secondary branches singly

or in whorls, up to 50 cm long; the ultimate branches 1-4 cm long, subtended by

lanceolate bracts, racemosely arranged, ending in umbellules; umbellules 10-20

flowered, pedicels slender 10-12 mm long.

Distr. Malay Peninsula: Perak, Selangor, Pahang. Sabah: Mt. Kinabalu.

Celebes: Menado, South Celebes. Philippines: Luzon, Panay, Catanduanes,

Mindanao.

Ecou, Thickets on mountainsides, often near streams, or among second growth.

Between 180-1550 m altitude.

Notes. The only species with the habit of a liane, with spaced leaves, and

recurved spines. The flowers are whitish or yellowish, and slightly fragrant. Visited

by numerous small bees. Fruit fleshy, purple. Scrapings of the bark are applied to

wounds and a decoction of the boiled bark is drunk to releave internal pain

(Mindanao).

4. Aralia bipinnata Blanco, FI. Filip. (1837) 222. — A. hypoleuca Presl. Epim.

(1851) 250. — “A. javanica Miq” F. Vill. Nov. App. (1880) 101. — A. glauca

Merr. in Philip. J. Sc. 2 (1907) 291. A. apoensis Elmer, Leafl. Philip. Bot. 7 (1914)

2325. A. bipinnata f. inermis Steen, in Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 392.

A shrub or small sparsely branched tree to 7 m high, with prickly stems, Leaf

to 1.5 m or more long, forming large crowns at the ends of the branches, bipinnate,

with some prickles, especially on the petiole or unarmed; leaflets sessile or with a

short petiolule, ovate to lanceolate, apex acute or acuminate, base rounded to

cordate, margin conspicuously crenate, upper surface green, glabrous, lower surface

glaucous, pubescent along the veins and sometimes sparingly on the mesophyll, or

almost glabrous throughout. Inflorescence a large terminal panicle, the peduncle

and also usually the main branches prickly, the whole either almost glabrous or

pubescent; tertiary branches usually 5-10 cm long, ending in umbellules, and

bearing a small number of lateral umbellules, or branches; umbellules with many

radiating pedicels (about 20-30); pedicels 5-10 mm long.

Distr. Philippines: Luzon, Leyte, Negros, Mindoro, Mindanao. West New

Guinea: Vogelkop. Also in Taiwan and Okinawa.

Eoo.. In rather open forests, ravines, and in thickets and second-growth;

occurring at about 700 m, but chiefly between 1000-2450 m.

Notes. van Steenis discusses the variability in pubescence and the development

of spines (Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 392.

5. Aralia ferox Miq. FI. Ind. Bat. 1, 1 (1856) 750. — A. filicifolia Ridl. J. Fed.

Mal. St. Mus. 8 (1917) 42.

Spiny shrub or small tree, usually unbranched, to about 10 m high. Leaves

forming a large rosette at the summit of the stem, up to about 1 m long, bi- or

tri-pinnate, prickly on the petiole, main rhachis, and often on the lateral rhachides;

leaflets sessile or petiolules to c. 5 mm, ovate or ovate-oblong, variable in size,

base truncate to rounded or cuneate, apex acute acuminate, margin sharply serrate,

100 Gardens’ Bulletin, Singapore — XXX (1977)

both surfaces with small bristle-like spines, especially on the veins, Inflorescence a

large terminal panicle, 25-50 cm long, glabrous, tertiary branches disposed singly

or in sub-verticils, ending in umbellules; flowers about 10-12 per umbellule.

Distr. Sumatra: West Coast, Mt Kerintje. Java: Gedeh, Patuha and Tang-

kuban Prahu.

Ecot. In montane scrub and among scattered trees; occurring between

1900-2900 m.

forma nana Steen. in Bull, Bot, Gard. Btzg. ser 3, 17 (1948) 394, fig. 1.

Smaller, probably 0.5-1.0 m tall, leaves tri-pinnate, 30 cm long, spiny all over,

leaflets 4-13 X 2-7 mm, rhachides of the ultimate pinnae winged.

Distr. Sumatra: known only from Mt Talang.

EcoL. Growing about 2500 m altitude.

Notes. van Steenis considers this interesting dwarf form to represent the

extreme of a series in size variability.

6. Aralia montana Bl. Bijdr. (1826) 870. — Panax armatus Wall. Cat (1832),

4933 nomen nudum; G, Don Gen. Syst. 3 (1834) 386. — Aralia montana var.

acutata Miq. Ann. Mus. Bot. Lugd. -Bat., 1 (1863) 9. — A. armata (Wall.) Seem.

Journ. Bot. 6 (1868) 134. — A. thomsonii Seem. Journ. Bot. 6 (1868) 134. — A.

montana var. crassifolia Bakh. — Oostr. in Back. Fl. Java (em. ed.) 7 (1948) fam.

159, 18.

A shrub or small tree, frequently unbranched, with prickly stems. Leaves to

1 m or more long, forming large crowns at the ends of the branches, bipinnate,

usually with some prickles, especially on the petiole, or unarmed. Leaflets sessile

or with a short petiolule, ovate, up to 14 X 7 cm, apex acute to acuminate, base

truncate or rounded, oblique in lateral leaflets, margin sharply serrate, upper surface

with the remains of a strigose tomentum, often + rugose, lower surface often

with a + velvety tomentum, or with more harsh hairs + confined to the veins.

Inflorescence a large terminal panicle, the peduncle and branches tomentose,

prickles, if any, confined to the peduncle and main rhachis; secondary branches at

intervals along the main rhachis, c. 35 cm long, bearing numerous tertiary branches

along their length; tertiary branches usually c. 6 cm long, ending in umbellules,

and often bearing a number of lateral umbellules; umbellules with many radiating

pedicels; pedicels usually 12-15 mm long, occasionally shorter, pubescent. Lobes 5,

5-celled, glabrous.

Distr. Malay Peninsula, Sumatra, Java, Sarawak, Sabah, Celebes.

EcoL. Primary and secondary forest, bamboo forest and low-lying moist

ground, from near sea level to 2600 m.

Notes. This species is considered to include all west Malesian examples with

pedicellate flowers and pubescent leaves, This broad concept is contrary to former

treatments which have recognized several species (A. thomsonii, A. armata), The

alliance with A. chinense Linn, is also very close. The application of names to this

and other Javanese species has been very confused. The position is ably discussed

by Koorders and Valeton Bijdr. 7 (1900) 52 et seq. and by van Steenis Bull. Bot.

Gard. Btzg Ser 3, 17 (1948) 391. In most specimens the lower leaf surface and the

pedicels are densely tomentose, but examples occur in which the leaf is only

sparsely hairy and the pedicels may be glabrous. A. armata appears to be within

the range of viaration of the complex as also does A. decaisneana Hance of China.

The most aberrant specimens are those with glabrous umbellules which are mostly

from Sumatra but also from Java: they may indicate that A. foliolosa Seem. should

also be included in this complex.

Scanning Electron Microscopical Studies on the

Spores of Pteridophytes

XI. The Family Oleandraceae (Oleandra, Nephrolepis and Arthropteris)

LIEW FAH SEONG

Department of Botany

University of Malaya

Abstract

The monolete bilateral spores of three genera of ferns belonging to the Oleandraceae

(Oleandra, Nephrolepis and Arthropteris) were examined by scanning electron microscope.

It is found that the spores of Oleandra are characterized by a large folded perine, and densely

echinate exine bearing pointed spines or blunt supporting rods. Arthropteris possesses spores

with more or less similar perine characteristics, and considering its morphological similarities

with those of Oleandra, this genus may well be included in the family Oleandraceae. Spores

of Nephrolepis, on the other hand, being characterized by insulate or verrucose perines,

thus resemble more closely those of the Davalliaceae, the closest relative of Nephrolepis.

However, many of its morphological and anatomical characteristics are so distinctive and

unique that it clearly constitutes a natural group by itself, and should, as proposed by Ponce

de Leon (1953), be treated as the only member of an independent family, the Nephrolepidaceae.

Introduction

The taxonomic and phylogenetic position of the fern genera Oleandra, Nephro-

lepis, Arthropteris and Psammiosorus have long been debated and remain contro-

versial. Oleandra Cav. is a genus of the tropics and has about 40 species. It differs

from other fern genera in many features, and for that reason its taxonomic position

is not fully established. Attributing great importance to the articulation of the stipe

to the stem J. Smith (1866) first segregated Oleandra from the Aspidiaceae and

considered it the sole genus of the new tribe of Oleandreae. After the detailed

anatomical study of the Japanese species O. wallichii by Ogura (1938), Oleandra

was considered as an isolated genus, and treated as the only genus in the family

Oleandraceae (Ching, 1940). Later, many authors accepted it as a monogeneric

family (Dickason, 1946; Ponce de Leon, 1953; Pichi-Sermolli, 1965). Others

however, have either treated it as “a genus incertae sedis’ (Bower, 1928), or placed

it among many other groups of ferns. Of the latter Diels (1889) arranged it in the

independent monotypic Tribus Oleandreae of Polypodiaceae, and Christensen

(1938) put it in the monotypic subfamily Oleandroideae of the same family. Still

other pteridologists included it in the Dennstaedtiaceae (Holttum, 1947), in the

Davalliaceae (Copeland, 1947; Alston, 1956; Tardiue-Blot, 1958), or in the sub-

family Oleandroideae of Davalliaceae (Tindale, 1961; Crabbe, Jermy & Mickel,

1975).

The genus Nephrolepis Schott has about 30 species distributed in the

tropics, Japan and New Zealand. Pres] (1836) included Nephrolepis in the section

Nephrodiariae of his tribe Aspidiaeae. Most workers placed it either in the family

Davalliaceae (Ching, 1940; Dickason, 1946; Copeland, 1947), or included it in a

separate group placed near the Davalliaceae (Nayar, 1970; Holttum, 1971, 1973).

However, Ponce de Leon (1953) established the family Nephrolepidaceae for this

group of ferns. Nephrolepis is usually classified with Oleandra because “they are

101

102 Gardens’ Bulletin, Singapore — XXX (1977)

probably more nearly related together than either is to any other genus” (Holttum,

1954).

Arthropteris J. Smith is a small genus of about 20 species and distributed in

the Old World tropics and southwards to Australia and New Zealand. ““The species

of Arthropteris are in utter confusion” (Copeland, 1958). The affinity of Arthrop-

teris is not clear either (Holttum, 1966). It has certain characteristics which are

similar in some extent, to the genera Oleandra, Nephrolepis, Davallodes, Elaphog-

lossum, Thelypteris, Teratophyllum, Ctenitis and Tectaria, It is usually placed

near Nephrolepis, but may not be its nearest relative (Holttum, 1966).

The monotypic genus Psammiosorus C. Chr. of Madagascar is peculiar and

shows features in common with Arthropteris (Copeland, 1947) and the Davallioid

ferns (Christensen, 1932; Holttum, 1966). It is always grouped with the Davallioids

or Oleandroids (Pichi-Sermolli, 1965), but its actual taxonomic position and

affinities are not clear.

These four genera have in the past and recently been variously grouped

(Holttum, 1966, 1971; Nayar, 1970; Sen & Sen, 1973; Crabbe, Jermy & Mickel,

1975): in one family, the’ Oleandraceae; in two separate families, the

Oleandraceae (Oleandra) and Davalliaceae (Nephrolepis, Arthropteris and

Psammiosorus) or in more than two families (Oleandraceae, Nephrolepidaceae

and Davalliaceae). In view of the controversy regarding the taxonomic problems

and phylogenetic relationships of these genera, it was thought that a detailed study

of them is a project worthwhile to undertake.

Spores of ferns are becoming increasingly important in fern taxonomy (Nayar,

1964; Wood, 1973; Moe 1974; Liew, 1976c). Because they offer both reliable and

stable morphological characters in (i) distinguishing species in some genera, (ii)

in differentiating genera, subgenera and characterizing families (Brown, 1960; Liew,

1976b) and (iii) in the tracing of possible phylogenetic relationships and trends of

evolution between taxa (Brown, 1960; Sorsa, 1964; Nayar & Dewi, 1968; Wagner,

1974) the present author decided to investigate this character and use it as a means

to tackle the problems raised above. Scanning electron microscopical observation

is preferred and undertaken because it offers the combined superiority of greater

depth of field, high resolution, and visually presents three dimensional topographical

information which reveals subtle and spectacular structures of spore surfaces

previously unobtainable with the optical microscope.

Materials and Methods

Spores of the three genera of ferns belonging to the Oleandraceae (Oleandra,

Nephrolepis and Arthropteris) were taken either from fresh plant material collected

in the fields or from herbarium sheets deposited in many institutions (NY, SING,

TAI, KLU and HK; see Table I for details). Fresh fronds were carefully dusted

for their spores and spores from herbarium specimens were obtained with the use

of clean tooth picks.

Before making scanning electron microscopical observation spores of each

species are dispersed in a drop or two of Hoyer’s solution (Anderson, 1954) and

checked preliminary for their proper identity under an ordinary light microscope.

Details of the method of preparation and observation on scanning electron micro-

scope were as described before (Liew, 1976a). Many models of scanning scopes

were employed in the present investigation, e.g., JOEL JSM 15, JSM U3, Cambridge

S4-10, and Hitachi MSM 4. Black and white photomicrographs were recorded on

ordinary 120 negative or on Polaroid films with (105 P/N, 55 P/N) or without

(42, 107) negatives. Sizes of spores were measured either in polar x equatorial axes

(P x E) or in polar x longest equatorial x shortest equatorial axes (P x El x E2).

Values given are averages of five to ten readings.

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104 Gardens’ Bulletin, Singapore — XXX (1977)

Results

The sizes and general characteristics of the spores of all the species of these

Oleandroid genera studied in the present investigation are given in Table I.

The insulate or verrucose spores of Nephrolepis clearly form a natural group

by themselves (Plates I and IL). These ellipsoid bilateral spores are monolete and

mostly concavo-convex in lateral view. A few of them are plano-convex. The

average size of the spore measures about 17-29 x 23-38 x 14-29 yw. Under the

SEM, a broken outer sporoderm will reveal the inner exine layer of the spore,

which is psilate, slightly scabrate or undulating (Plate I, fig. 8). The laesura is

slightly raised, thin and with a short ridge in some species (e.g., in N. pectinata and

N. davallioides) or thick and forming long and rough ridges in others (e.g. in

N. acutifolia and N. biserrata). The sculptine or sporoderm has on its surface

elements of various sizes, from small grains or humps to large tubercules, or with

irregular excrescences or raised ridges of various sizes.

The ellipsoid spores of Oleandra are totally different from Nephrolepis (Plates

III and IV). They are monolete bilateral spores, and are plano-convex to slightly

biconvex in lateral view. The amb is oblong in shape, All the spores have a thick,

prominent, loose and wrinkled perine which anastomoses to form regular or

irregular polygons or lophate sculptures. Depending upon the species concerned,

the perine may have spinulose, blunt or dentate excrescences. The size of the spore

is 25-42 x 30-55 x 26-32 yw. Scanning electron micrographs of broken sporoderms

reveal that the outer layers are built on stilts, and are supported by numerous

rods connected to the layer(s) down below. These micrographs also reveal the

smoothness of the inner exine in many cases (Plate IV, fig. 40). Spores of Oleandra

may differ in sculptural details, even within the same species, but they nevertheless

form a group with a unique and distinctive general pattern.

Spores of Arthropteris have a thick perine of various sculpturing ornamenta-

tions (Plates V and VI). Foldings may be high or low in profile, and are covered

with minute projections or interlacing networks. Some Arthropteris spores (e.g.

A. articulata) closely resemble those of Oleandra species. The outer perine of most

of the spores of Arthropteris sloughs off easily, leaving behind the naked exine with

smooth surface and laesura. The size of the spores range 28-56 x 40-65 x 25-30 un.

It is ellipsoid and sometimes roundish in polar view, and plano-convex to concavo-

convex in lateral view.

Discussion

I. Nephrolepis

From our studies on eleven taxa of Nephrolepis gathered from various parts of

the world it is demonstrated that the bilateral spores of Nephrolepis species are

rather uniform and form a clear-cut group by themselves. They have thin perine -

and verrucose sculptine with elevations of various sizes, from slightly raised ridges —

to conspicuous tubercules.

In the literature, spores of many species of Nephrolepis have been studied.

Holttum (1954) mentioned that spores of Nephrolepis have slightly and irregularly

roughened surfaces, and are without distinctive markings. Hannig (1911) pointed

out that “‘a perispore is unlikely to occur” in Nephrolepis (N. exaltata). This is

contradicted here as shown in Plate I, fig. 8, which clearly shows the presence of an

outer layer. Among the many authors who have examined the spores of Nephrolepis

exaltata (L.) Schott was Selling (1946) who investigated Hawaiian species and

described it as having very irregular warts and short, irregular ridges, with a size

of 24-28 x 41-50 yu. Earlier workers have similarly investigated the spores of this

species but did not mentioned the sculpture patterns. Marquesan (southeastern

Spores of Oleandraceae 105

Polynesia) spores of this taxon are + 34 w in length, and with the surface

conspicuously tuberculate (Brown & Brown, 1931). Erdtman and Sorsa (1971)

described the spores from British West Indies as convex-plane, 22 x 37 x 23 u

(perine not included) with the laesura about 21 «, and the perine similar to that

in N. biserrata. The exine stratification is obscure. Spores from North America

(Florida) are subpsilate to tuberose, ellipsoid, monolete, and measure 38-55 x

21-31 » (Kremp & Kawasaki, 1972).

Harris (1955) described the spores of the New Zealand Nephrolepis cordifolia

as having verrucate surfaces with elongated projections and of 18-40 xu. Spores of

the same species in India (Darjeeling) are reported to be bilateral and possess a

very thin perine forming small humps (Sen & Sen, 1973).

Some other species of Nephrolepis have also been described by Erdtman and

Sorsa (1971). For example, the spores from Java of N. davallioides (labelled as

N. acuminata) are provided with a more or less verrucose, dark brownish, cf.

perinous layer ca 1.0 » broad. The spores are reported to be concavo-convex and

of 17 x 34 x 23 » in size (perine not included). The Ceylonese species of N.

biserrata has convex-concave spores and measured 23 x 39 x 26 » (perine included).

It has a perine ca 1.5 » thick, is verrucose; warts are dark brownish ca 1.3 x

(2.0-6.0) » high. The Japanese species of N. falcata has a more or less perinous

thickening which is columnar, wart-like and about (3.0-4.0) x (4.0-7.0) yu. The

spores are 30 x 45 x 35 uw (perine not included). The Ceylonese species of N.

hirsutula has convex-plane or slightly biconvex spores, 21 x 35 x 25 » thick and

finely verrucose.

The bilateral and more or less verrucate sculptine patterns of the spores is

similar, if not identical, in all species of Nephrolepis so far examined, and thus

form a rather homogeneous group by themselves. Though a few of the Nephrolepis

species are remarkably variable, poorly defined or with bizarre features, the

uniformity and uniqueness in their vegetative characters, however, has clearly

demonstrated the naturalness of the whole group (Pichi-Sermolli, 1965). “It

therefore constitutes a clearly natural genus’ (Copeland, 1958).

Traditionally Nephrolepis has been associated with the Davallioids, Dick-

sonioids or the Aspidioid ferns. Copeland (1947) commented that this genus has

many morphological and anatomical characters in common with the Davalliaceae

and affinity seems certain, but many are not close. In chromosome number

(n= 41), scales (peltate) and indusium (reniform), Nephrolepis resembles

Oleandra; but in many other characters, such as habit, anatomy of the stem and

stipe, articulation of the stipe, the feature of the rhizome, venation, position of the

sori and structure of the spores, they differ considerably (Pichi-Sermolli, 1965).

However, as a group, this genus has spores similar to its closest relative of

Davalliaceae ferns, Davallia, Humata, Araiostegia, Leucosteiga and Davallodes,

all of which possess spores with markings of varying degree. Undoubtedly, the

phylogenetic position of Nephrolepis is thus in the Davalliales.

Since features of some Nephrolepis species are similar to some of the fern

genera and families, especially members of the Davalliaceae, and have in the mean-

time shown to be distinctive in other important characters, it might be better to

group all the species of Nephrolepis in a separate and distinctive group, the family

Nephrolepidaceae, as was suggested by Ponce de Leon (1935). The phylogenetic

affinity of this family is with the family Davalliaceae.

Il. Oleandra

The bilateral spores of the genus Oleandra are also homogenous in having a

distinctive, thick, smooth or wrinkled perine with a few or many protruding folds.

The folds may form an irregular network having large meshes (lophate). Spinulose

excrescences distributed on the surface may be few or numerous. Usually these

106 Gardens’ Bulletin, Singapore — XXX (1977)

are broader and thicken at the base tapering to sharp apices. Some of these

structures have blunt rods.

In the literature, all workers who have done palynological work on Oleandra

have reported similar findings. For example, Erdtman and Sorsa (1971) described

the convex-plane spores of O. musifolia of Ceylon as being 28 x 40 x 30 p

(perine not included), the perine with folds of ca 5.0-10.0 » high, and anastomosing

to form irregular polygons with broadly conical and blunt crests. The perine is

sparsely spinulose or dentate. It is provided with densely spaced, intra-perinous

rods about (2.0-5.0) x (0.5-1.0) » and spaced ca 1.0-2.0 p» apart. As described

by Kremp and Kawasaki (1971) the ellipsoid spores of the Javanese Oleandra

neriiformis measure 43-63 x 36-49 », and are monolete and perisporate or psilate

(in this case, presumably the perine has sloughed off!). Erdtman (1957) also

illustrated the spore of this species, Copeland (1947) mentioned that the spores

are bilateral or angular by shrinkage of the epispore and are apparently smooth.

Sen and Sen (1973) illustrated and described the spores of Indian species of

O. wallichii as monolete, bilateral plano-convex in lateral view, and oblong in polar

view. The perine is said to be folded and spiny.

The habit of growth of Oleandra is almost unique and has been described as

the only shrubby fern, although all species included in it are not necessary shrubby

(Nayar, Bajpai & Chandra, 1968). The genus also shows homogeneity in many

features, such as spore sculpture, etc. It differs from all other ferns in the anatomy

of the stem, the position of the sori, the presence of peculiar aerial roots, and also

in the structure of the spores (Pichi-Sermolli, 1965; Braggio, 1966). There are

thus enough justification to treat Oleandra as an isolated genus in the independent

family Oleandraceae Ching ex Pic. Ser., as agreed by many pteridologists (J. Smith,

1866; Diels, 1889; Christensen, 1938; Ching, 1940).

The ancestry of Oleandra is unknown (Copeland, 1947). Its remarkable

resemblance in many features with some species of Davallia and Humata suggests

clearly its close affinity with the Davallioid ferns. Nowadays, most workers place

it under the Davalliales.

In ferns, bilateral perinate spores are characteristic of the Aspidioids,

Asplenioids, Lomariopsidoids and the Thelypterioid ferns, and the presence of

spinulose excrescences over the perine is found especially common among the

Aspidioid genera (Nayar & Devi, 1964; Nayar & Kaur, 1965). When the characters

of the spores are taken into consideration with anatomical, morphological and

cytological features, it is suggested that this unique group of ferns has a closer

phylogenetic relationship with the Aspidioid ferns, as has been suggested by many

investigators such as Presl (1836), Dickason (1946), Nayar, Bajpal and Chandra

(1968), and Pichi-Sermolli (1965). The last author has also emphasized that its

affinity in the second place is with the Polypodiales, rather than with the

Dicksoniales to which affinity was alluded by most authors in the past.

Ill. Arthropteris

Copeland (1947) referred to the spores of Arthropteris as oblong and with

epispore. In his study on the genus of Arthropteris in Malesia, Holttum (1966)

mentioned that the spores of this genus have folded perispores.

However, there are some differences of opinion concerning the sculptine

structures even within the same species. For example, Harris (1955) reported the

New Zealand species of A. tenella as having bilateral, winged spores with cristate

perispore separated from the spore wall and both of these structures differ in

contour. The surface is striate-scabrate and with a few crests. The crests, when

seen in oblique or transverse view, form irregular, almost spinulose projections,

9-14 » in height. The contour or outline is deeply resected. However, spores of

Spores of Oleandraceae 107

the same species from the same country were reported by Sen and Sen (1973) to

be monolete-bilateral and possessing a thin and granulose (in the description) or

verrucose (in the diagram) perine. Furthermore, Kremp and Kawasaki (1972)

reported this same species to have monolete ellipsoid spores of 34-62 x 27-43 xu,

and having a thin perine with psilate surface. Our SEM investigation on 4.

tenella, however, confirmed the correct observation and description of Harris

(1955). The verrucose spores of Sen and Sen (1973) were most probably due to

contamination or observation on abnormal spores. Those observations by Kremp

and Kawasaki (1972) could have been either too immature or mature spores with

ablated perine, a phenomenon frequent among many Arthropteris species.

Spores of several other species of Arthropteris have been studied e.g. A.

altescandens of Juan Fernandez was reported to have spores with short elliptic to

subcircular amb and as having a very thin (ca 0.3 «) and folded perine (folds up

to 10 » high). Those of A. obliterata (labelled as Nephrolepis ramosa Moore) of

Ceylon was said to have no perine, were biconvex and 39 x 55 x 40 «. The psilate

spores of A. monocarpa of Africa and Madagascar and the spores with a large

and folded perine and smooth exine of the African (Congo) A. orientalis have

also been studied by Braggio (1966).

The affinity of Arthropteris is not clear. Copeland (1947) placed it tentatively

in the Oleandra-Davallia series. Collective similarities in aspects of frond structure,

articulation of pinnae, terminal sori on veins, are evidence of affinity with the

Nephrolepis. Holttum (1966) does not think that Nephrolepis is its nearest relative

despite the placement near Arthropteris. He, however, agreed with J. Smith in

accepting Arthropteris as a natural genus, and believed that its peculiar combina-

tion of characters may give significant hints as to interrelationships among various

groups of ferns. But, he is not yet clear as to how such a relationship may best be

formally expressed in a scheme of classification. From studies on the spores of two

Arthropteris species of Africa and Madagascar, Braggio (1966) emphasized that

Arthropteris differs from both Oleandraceae and Davalliaceae, and that it ought

to be referred to the order of Aspidiales because of its spore characteristics (with

psilate exine and a wide and thin perine, with or without ornamention).

From our observation on spore structure Arthropteris species is closer to the

Oleandroids rather than to the Nephrolepis and Davalliaceae. This is especially

evident from the spores of A. articulata which are almost indistinguishable from

those of some Oleandra species (Plate V, figs. 46-49). Spores of other species of

Arthropteris may not be more distant as they possess a smooth exine and a thick

and variously folded perine with different sculptured patterns (Plate VI). However,

they still display numerous structural features in common with the Oleandra

spores. Pichi-Sermolli (1965) also pointed out the similarities of both genera, such

as the articulation of the stipe, the shape of the sorus and indusium, the presence

of a wide perine, and the chromosome number. When everything is taken into

consideration, the disposition of Arthropteris in the Davalliales is undoubtedly

natural.

The proper systematic position of the peculiar monotypic genus Psamyniosorus

(endemic in Madagascar) is not fully established (Christensen, 1932; Copeland,

1947; Tardieu-Blot, 1964; Pichi-Sermolli, 1965; Braggio, 1966; Holttum 1966).

In conclusion, the family Davalliaceae sensu Copeland (1947) can now be

envisaged as the order Davalliales which comprises three natural groups, the family

Davalliaceae s.s, containing more strictly Davallioid genera, the family Oleandra-

ceae, with the genera Oleandra and Arthropteris, and the family Nephrolepidaceae,

with the monotypic genus Nephrolepis. The proper phylogenetic position of

Psammiosorus cannot be proposed until further study. At the level of the Order,

the affinity of the Davalliales is close to the Aspidiales.

108 Gardens’ Bulletin, Singapore — XXX (1977)

Acknowledgements

The author gratefully acknowledges his indebtedness to all directors and

curators in charge of institutions for making herbarium materials available for the

present investigation (NY, SING, TAI, KLU & HK). He also thanks Mr. and Mrs.

C. S. Wang and C. F. Lo; Miss S. C. Wang, Mr. Y. C. Huang and Mr. K. W. Ho

for their cooperation and technical assistance in the use of different models of

SEMs at the various institutions and universities.

Literature Cited

Alston, A. H. G. 1956. The subdivision of the Polypodiaceae. Taxon, 5: 23-25

Anderson, L. E. 1954, Hoyer’s solution as a permanent mounting medium.

Bryologist, 57: 242-244.

Bower, F. O. 1928. The Ferns (Filicales). Vol. If1. The Leptosporangiate Ferns.

Cambridge Univ. Press, Cambridge.

Braggio, G. 1966. Morfoligia delle spore sistematica delle Davalliales. Webbia,

21: 725-767.

Brown, C. A. 1960. What is the role of spores in fern taxonomy? Amer. Fern. J.,

50: 6-14.

Brown, E. D. W. & F. B. H. Brown. 1931. Flora of Southeastern Polynesia.

II. Pteridophytes. Bishop Mus. Bull, 89.

Case, I. M. 1932. The development of the sorus in some species of Nephrolepis

with some observations on points of anatomical interest. Trans, Roy. Soc.

Edinb., 57: 258-276.

Ching, R. C. 1940. On the natural classification of the family Polypodiaceae.

Sunyatsenia, 5: 201-268.

Christensen, C. 1932. The Pteridophyta of Madagascar. p. 70-74. Hayerup’s

Boghandel, Kobenhavn.

1938. Filicinae, In F. Verdoorn (ed.), Manual of Pteridology. p.

522-550. Nijhoff, the. Hague.

Copeland, E. B. 1947. Genera Filicum. Chronical Botanica, Waltham, Mass.

1958. Fern Flora of the Philippines. Vol. I. Nat. Inst. Sci. Tech.,

Manila.

Crabbe, J. A., A. C. Jermy & J. T. Mickel. 1975. A new generic sequence for the

pteridophyte herbarium. Fern Gaz., 11: 141-162.

Dickason, F. G. 1946, A phylogenetic study of the ferns of Burma. Ohio J. Sci.,

46: 73-108.

Diels, L. 1899. Polypodiaceae. In A. Engler & K. Prantl, Die Naturlichen Pflanzen-

familien, I (4): 139-339. Engelmann, Leipzig.

Erdtman, G. 1957. Pollen and Spore Morphology/Plant Taxonomy. (Corrected

reprint 1972) Hafner Pub. Co., New York.

Erdtman, G. & P. Sorsa. 1971. Pollen and Spore Morphology/Plant. Taxonomy.

IV. Pteridophyta. Almqvist & Wiksell, Stockholm.

Hannig, E. 1911. Uber das Vorkommen von Perisporien bei den Filicinen nebst

ge gi uber die systematische Bedeutung derselben. Flora, 103:

321-346.

ae eS

Spores of Oleandraceae 109

Harris, W. F. 1955. A manual of the spores of New Zealand Pteridophyta. New

Zealand Dept. Sci. Ind. Res. Bull. 116: 1-186.

Holttum, R. E. 1947. A revised classification of leptosporangiate ferns. Bot. J.

Linn. Soc., 53: 123-158.

1949. Classification of ferns. Biol. Rev., 24 267-296.

1954. A Revised Flora of Malaya. Vol. Il. Ferns. (2nd ed., 1968)

Government Printing Office, Singapore.

—_—_———— 1966. The genus Arthropteris J. Sm. in Malesia. Blumea,

14: 225-229.

1971. The family names of ferns. Taxon, 20: 527-531.

1973. Introduction to Pteridophyta (and) Family names of

Pteridophyta (+ all pteridophyte entries). In J. C. Willis, A Dictionary of

the Flowering Plants and Ferns, 8th ed. Revised by H. K. Airy Shaw.

Cambridge Univ. Press, Cambridge.

Kremp, G. O. W. & T. Kawasaki. 1972. The Spores of the Pteridophytes. Hirokawa

Book Co., Tokyo.

Liew, F. S. 1976a. Scanning electron microscopical studies on the spores of

Pteridophytes. I. The genus Pyrrosia Mirbel (Polypodiaceae) found in Taiwan.

Quart. J. Taiwan Mus., 24: 191-216.

1976b. A scanning electron microscopical study on the spores of

Lycopodium found in Taiwan. Sci. Agr., 24: 162-172.

—_—_ 1976c. Scanning electron microscopical studies on the spores of

pteridophytes. VII. The family Plagiogyriaceae, Taiwania, 21: 37-49.

Moe, D. 1974. Identification key for trilete microspores of Fennoscandian

Pteridophyta. Grana Palyn., 14: 132-142.

Nayar, B. K. 1964. Palynology of modern Pteridophyta. In B. K. Nair (ed.),

Advances in Palynology, p. 100-141. United Block Printers, Lucknow.

1970. Classification of homosporous ferns. Taxon, 19: 229-236.

Nayar, B. K. & S. Devi. 1964. Spore morphology of Indian ferns. I. Aspidiaceae.

Grana Palyn., 5: 80-120.

1968. Spore morphology of the Pteridaceae. IV. Taxonomic and

phyletic considerations. Grana Palyn., 8: 517-535.

Nayar, B. K. & S. Kaur, 1965. Spores morphology of the Lomariopsidaceae.

J. Palyn., 1: 10-26.

Nayar, B. K., N. Bajpai & S. Chandra. 1968. Contributions to the morphology of

the fern genus Oleandra. Bot. J. Linn. Soc., 60: 265-282.

Ogura, Y. 1938. Anatomy and morphology of Oleandra wallichii (Hk.) Presl.

Jap. J. Bot., 9: 193-211.

Perez Arbelaez, E. 1928. Die naturliche Gruppe der Davalliaceen (Sm.) Kaulf. Bot.

Abhandl. K. Geobel, 14:1-96.

Pichi-Sermolli, R. E. G. 1959, Pteridophyta. In W. B. Turril (ed.), Vistas in

Botany, p. 421-493. Pergamon Press, London.

—————. 1965. Adumbratio Florae Aethiopicae. II. Oleandraceae. Webbia,

20: 745-769.

110 Gardens’ Bulletin, Singapore — XXX (1977)

1970. A provisional catalogue of the family names of living

pteridophytes. Webbia, 25: 219-297.

Pokrowskaila, I. M. 1958. Analyse Pollingue. Ann. Serv. Inform. Geol. B.R.G.G.M.,

24: 1-435.

Ponce de Leon, A. 1953. Nociones de botanica Sistematica. VIII. Division

Pteridophyta. Revta Soc. Cuban Bot., 10: 37-45.

Presl, C. B. 1836. Tentamen Pteridographiae. Haase, Prague.

Selling, O. H. 1946. Studies in Hawaiian pollen statistics, Part I. The spores of the

Hawaiian pteridophytes. Bishop Mus. Spec. Pub, 37: 1-87.

Sen, T., U. Sen & R. E. Holttum. 1972. Morphology and anatomy of the genus

Davallia, Araiostegia and Davallodes, with a discussion on their affinities.

Kew Bull., 27: 217-243.

Sen. U. & T. Sen, 1973. Anatomical relationships between the Oleandra and

Nephrolepis groups. Bot. J. Linn. Soc. 67, Suppl. 1: 155-172.

Smith, J. 1866. Ferns, British and Foreign. p. 73. London.

Sorsa, P. 1964. Studies of the spore morphology of Fennoscandian fern species.

Ann. Bot. Fenn., 1: 179-201.

Tardieu-Blot, M. L. 1958. Polypodiacees (sensu lato). Dennstaedtiacees. Aspidia-

cees. In H. Humbert (ed), Flore de Madagascar et des Comores (Plantes

Vasculaires). Fam. 5. 1: 1-391. Typographie Firmin-Didot, Paris.

1964. Sur les spores de Davalliaceae et Vittariaceae Malgaches.

Pollen Spores, 6: 537-544.

Tindale, M. D. 1961. Pteridophyta of South Eastern Australia. Davalliaceae.

Contr. N.S. Wales Nat. Herb. Fl. Ser. 208-211: 1-4.

Wagner, W. H., Jr. 1974. Structure of spores in relation to fern phylogeny.

Ann. Missouri Bot. Gard., 61: 332-353.

Walker, J. W. & J. A. Doyle. 1975. The bases of Angiosperm phylogeny: Palynology.

Ann. Missouri Bot. Gard., 62: 664-723.

Wood, C. C. 1973. Spore variation in the Thelypteridaceae. Bot. J. Linn. Soc. 67.

Suppl. 1: 191-202.

Scale for figures of Plates I to VI

For values of magnifications in the legend. / cm on the micrographs approximates to:—

67u (x 150) IS5u (x 750) 6.7u (x 1500)

40u (x 250) 10x (x 1000) 4u (x 2500)

20u (x 500) 8u (x 1250) 2.5u (x 4000)

WG QE ‘

: Sk TAM

. WS \ x S

S WG NS S y . \ XC

SSS

NSS

SAY

Plate I. Spore characteristics of Nephrolepis.

Figs. 7: Group of spores shown at different views: 2, 3, 4: typical bilateral spores of

Nephrolepis, with surfaces more or less smooth (2) and rough (3, 4); 5, 6: spores in polar

view, showing laesurae thin (5) and thick (6); 7-/0: close-up views of the spore surfaces,

showing more or less smooth (7), moderately roughened (8, 9) and rough (/0) surfaces. Note

that 8 is of a broken spore of N. exaltata, showing the outer rough surface (perine) and the

inner smooth surface (exine).

N. pectinata: figs. J] (x 750), 2 (x 1000), 7 (x 2500); N. cordifolia: 3 (x 1500), 5 (x 1500);

N. hirsutula: 4 (x 1250), 9 (x 2500), 7/0 (x 5000); N. biserrata: 6 (x 1500); N. exaltata: 8

(x 2500).

Plate II. Some representative spores of Nephrolepis species.

N. acutifolia: figs. 1] (x 1250), 12 (x 500); N. biserrata: 13 (x 1500); N. cordifolia: 14

(x 1500), 75 (x 1250); N. davallioides: 16 (x 1250); N. dicksonioides: 17 (x 1250); N. exaltata:

18 (x 1000); N. falcata: 19 (x 1250); N. hirsutula: 20 (x 1250), 21 (x 1000); N. lauterbachii:

22 (x 1250); N. pectinata: 23 (x 1000); N. radicans: 24 (x 1250).

WSS

SSS

Plate III. Spore characteristics of Oleandra.

Figs. 25: Group of spores within a broken sporangium: 26: typical structure of a spore

with broken perine; 27: a partly spinulose and partly ridged spore: 28: a wholly spinulose

spore; 29: another type of spinulose spore with thick ridges; 30, 3/, 32: close-up views of

some broken perines, showing erect blunt rods and details of surrounding structures: 33, 34

close-up views of some unbroken perines. showing holes and short spines; 35: structur

the short spines, etc. on the perine.

O. undulata: 25 (x 150), 26 (x 1000), 29 (x 500), 37 (x 1250), 32 (x 4000), 35

O. wallichii: 27 (x 1000), 28 (x 750); O. madagascarica: 33 (x 500); O. pistillaris: 30 (

34 (x 500).

re oO

43 44 45

Plate IV. Some representative spores of Oleandra species.

O. Madagascarica: figs. 36 (x 500), 37 (x 1250); O. neriiformis: 38 (x 1000), 39 (x 1500),

40 (x 1000); O. pistillaris: 41 (x 500); O. undulata: 42 (x 1000); O. wallichii: 43 (x 1000),

44 (x 1000), 45 (x 750).

Plate V. Spore characteristics of Arthropteris.

Figs. 46: Group of spores seen under low magnification; 47, 48: normal spores and

part of an ablated spore; 49, 50, 5/, 52: close-up views of the spores, showing detailed

structures of various kinds of outgrowths on the perines; 53, 54: broken spores, showing

rough perines and smooth exines.

A. articulata: figs. 46 (x 250), 47 (x 750), 48 (x 750), 49 (x 2500); A. pallisotit: 50 (x 2500),

53 (x 750); A. tenella: 51 (x 2500); A. beckleri: 52 (x 1250); A. dolichopoda: 54 (x 1250)

Plate VI. Some representative spores of Arthropteris species.

A. beckleri: figs. 55 (x 500), 56 (x 500); A. dolichopoda: 57 (x 500). 58 (x 500); A.

monocarpa: 59 (x 250); A. obliterata: 60 (x 750); A. pallisotii: 61 (x 750), 62 (x 500); A.

repens: 63 (x 500); A. tenella: 64 (x 1000); A. wollastonii: 65 (x 500), 66 (x 500).

SAURAUIAE GERONTOGEAE

I. Notes on Malayan species

R. D. HOOGLAND

Australian National University

Canberra

Abstract

Saurauia tristyla has been erroneously recorded from Malaya; the correct placement of

these records is discussed. §. roxburghii Wall. and S. napaulensis DC. are new records for

the area. S. pentapetala (Jack) Hoogl. is proposed as a new combination, with basionym

Ternstroemia pentapetala Jack. Three new species are described: S. fragrans Hoogl., S.

mahmudii Hoogl., and S. malayana Hoogl. S. cauliflora var. calycina King is placed in

synonymy under S. leprosa Korth., and S. grandis Ridl. in synonymy under S. vulcani Korth.

The altitudinal distribution of the 10 species found in Malaya is discussed.

Introduction

Under the general title SAURAUIAE GERONTOGEAE the author hopes

to present a number of papers on the Old World species of Saurauia. The present

must be seen only as one which is preliminary to a fully documented account of

the Malayan species to be presented in the near future. Other accounts currently

being prepared cover the species from continental South-East Asia, Java, and the

Lesser Sunda Islands. Considering the high degree of local endemism in the genus,

the preparation of separate treatments for the major islands or island groups of

Malesia offers the opportunity to gradually compound our knowledge of the genus

in this region.

The author is grateful to the authorities of the following herbaria for their

ala in allowing the study of their collections during visits or by way of

oans:

Botanical Section, Department of Agriculture, Bangkok (BK).

Forest Herbarium, Royal Forest Department, Bangkok (BKF).

British Museum (Natural History), London (BM).

Herbarium Bogoriense, Bogor (BO).

peor ecamem Herbarium, Botanical Survey of India, Sibpore, Howrah

Herbarium Australiense, C.S.I.R.O., Canberra (CANB).

Royal Botanic Garden, Edinburgh (E).

Conservatoire et Jardin botaniques, Genéve (G).

Royal Botanic Gardens, Kew (K).

Forest Research Institute, Kepong (KEP).

Department of Botany, University of Malaya, Kuala Lumpur (KLU).

Rijksherbarium, Leiden (L).

Laboratoire de Phanérogamie, Muséum National d’Histoire Naturelle.

Paris (P).

Botanic Gardens, Singapore (SING).

De ore we Universiti Kebangsaan Malaysia, Kuala Lumpur

111

112 Gardens’ Bulletin, Singapore — XXX (1977)

Fieldwork in Malaya was made possible with generous assistance from the

University of Malaya, Kuala Lumpur; the Forest Research Institute, Kepong; the

Universiti Kebangsaan Malaysia, Kuala Lumpur; and the Botanic Gardens, Penang.

I am indebted to Mrs. Cathy Porter for the illustrations of Saurauia mahmudii

and §. malayana, and to Miss Jill Ruse for those of S. fragrans.

I. Saurauia tristyla auct. non DC. in Malaya

The type of Saurauia tristyla DC, [in Mem. Soc. Phys, Hist. Nat. Genéve 1

(1822) 423, t. vii] originated from the Moluccas and represents a species which

has subsequently been collected repeatedly there and in Celebes (Sulawesi). It has

also been recorded from a wide geographical area including West New Guinea,

Sumatra, Borneo, Malaya, Burma, Thailand, Indo-China, and Southern China

(including Taiwan). Most, if not all, of these further records must be discarded

as being referable to other species. The present note concerns itself only with the

records and collections from Malaya and Peninsular Thailand. The main literature,

under this name and for this area, is contained in King [in J. Asiat. Soc. Beng.

59 II (1890) 197], Ridley [Fl..Mal. Pen. 1 (1922) 207], Craib [Fl]. Siam. Enum.

1 (1925) 129], and Keng [in Fl. Thail. 2 (1972) 110].

A study of the material in CAL, seen and annotated by King, revealed that

he included specimens of three species for which I accept the names Saurauia

fragrans Hoogl., S. pentapetala (Jack) Hoogl., and S. roxburghii Wall. In addition,

two specimens which I refer to §. pentapetala were annotated S. cauliflora DC.

(Scortechini 99) and S. pendula Bl. (Scortechini 331) whereas one collection

(Scortechini s.n.: CAL 49323/4) of S. roxburghii was annotated “quite different

from S. tristyla DC., G. King’. None of these appear to have been considered

further in King’s account of the genus in the Materials for a Flora of the Malayan

Peninsula. The same three species were available to Ridley in the herbaria of

Singapore and Kew and included by him in S. tristyla, but as S. fragrans has not

been found in Thailand the accounts of S. tristyla by Craib and Keng include only

the other two species.

1. Saurauia pentapetala (Jack) Hoogl., comb. nov.

Basionym: Ternstroemia pentapetala Jack in Mal. Misc. 1 (5) (1821) 40; cf.

Merr. in J. Arn, Arb. 33 (1952) 247. = Cleyera pentapetala (Jack) Spreng., Linn.

Syst. Veg. (ed. 16) 2 (1825) 596. = Scapha pinangiana Choisy in Mém. Soc. Phys.

Hist. Nat. Genéve 14 (1855) 119, nom. illeg. — TYPE: ‘“‘Native of Pulo Penang”’,

lost; NEOTYPE: Hoogland 12601, SE. slope of Penang Hill, Penang, 8 April

1975, on forest edge at ca 420 m altitude, in CANB, duplicates being distributed

to K, KEP, L, SING, US.

Synonym: Saurauia nudiflora var. angustifolia Craib, Fl. Siam. Enum, 1 (1925)

128. — TYPE: ‘“PATTANI. Bachaw, c. 600 m., by stream in evergreen forest,

Kerr, 7196”; HOLOTYPE in ABD, n.v., ISOTYPES in BK, BM, E, K.

Misapplied name: Saurauia tristyla auct. non DC.: Wall., Numer. List (1829)

no. 1466 et Pl. Asiat. Rar. 2 (1831) 40; Dyer in Fl. Br. Ind. 1 (1874) 287, p.p.;

King in J. Asiat. Soc. Beng. 59 II (1890) 197, p.p.; Ridl., Fl. Mal. Pen. 1 (1922)

207, p.p.; Craib, Fl. Siam. Enum. 1 (1925) 129, p.p.; Burk. & Henders., Gard.

Bull. Straits Settl. 5 (1925) 349, p.p.; Henders., ibid. 4 (1928) 226, p.p.; Burk.,

Dict. Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966) 2002, p.p.; Keng in FI.

Thail. 2 (1972) 110, p.p.

The two earliest names, validly published for Old World species of Saurauia,

are those given by W. Jack under the generic name Ternstroemia to one Sumatran

and one Malayan species.’ The Sumatran species was correctly renamed Saurauia

rubiginosa (Jack) Merr., but the required new combination has never been proposed

Malayan Saurauiae 113

for the Malayan species. The first author to have placed it in Saurauia appears to

have been Dyer (1874) who lists it in synonymy under S. tristyla DC. This species

had already been recorded from Penang, the type area for Jack’s name, by Wallich

(1829, 1831) on tke basis of a specimen collected by Porter. All subsequent

authors followed this placement of the name. Merrill, in his account of William

Jack’s genera and species of Malayan plants (1952), did not specifically place it

beyond saying that it “belongs clearly in the group with S. tristyla DC.”

Saurauia pentupetala is probably the most common and most widely distri-

buted species in Malaya, having been found in most of the mainland states and

on Penang and Singapore islands. In Peninsular Thailand it is known from Ranong

and Surat Thani districts southward. The species is most commonly found in the

lowlands, up to ca 500 m altitude, with an occasional occurrence at higher altitudes

(up to 1100 m).

2. Saurauia roxburghii Wall., Numer. List. (1829) no. 1467, nom. nud., et PI.

Asiat. Rar. 2 (1831) 40. — TYPE from India.

Misapplied name: Saurauia tristyla auct. non. DC.: King in J. Asiat. Soc. Beng.

59 II (1890), 197, p.p.; Ridl. in Trans. Linn, Soc. London II Bot. 3 (1893) 277,

et Fl. Mal. Pen. 1 (1922) 207, p.p.; Craib, Fl. Siam. Enum. 1 (1925) 129, p.p.;

Burk. & Henders., Gard. Bull. Straits Settl. 3 (1925) 349, p.p.; Henders., ibid. 4

(1928) 226, p.p.; Burk., Dict. Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966)

2002, p.p.; Keng in Fl. Thail. 2 (1972) 110, p.p.

Saurauia roxburghii is a widespread species of continental South-East Asia,

extending from eastern Nepal through southern China to the Ryukyu Islands,

through Burma, Thailand, and Indo-China, and is now for the first time recorded

for Malaya. Here it is less widespread than S. pentapetala, being absent from

Malacca, Johore, Singapore, and Penang, and, though it has been found at quite

low altitudes, tends to be more frequent between 500 and 1000 m where it may be

common in regrowih vegetation; the maximum recorded altitude is 1250 m.

3. Saurauia fragrans Hoogl., sp. nov. (fig. 2)

Arbor parva usque 12 m alta, 20 cm diametro. Folia oblonga vel obovato-

oblonga, plerumque 18-30 cm longa, 6-11 cm lata, 14-17-nervata, apice acuta +

acuminata, basi acuta vel obtusa, petiolo 24 cm longo. Flores solitarii vel usque

20 fasciculati, interdum inflorescentia cymosa paucifiora interspersa, in axillis

foliorum vel cicatricium foliorum. Pedunculus usque 5 mm _ longus, pedicellus

§-22 mm longus. Sepala suborbicularia, 3-5 mm longa et lata. Corolla late

campanulata (fere plana), 2 mm.alta, 13 mm diametro; tubus 14-2 mm longus;

lobi 51-84 mm longi, 44-74 mm lati, apice bilobulati. Stamina plerumque 20-23,

filamentis subaequalibus 23-34 mm longis, antheris versatilibus dorsifixis 1}-2 mm

longis. Ovarium subglobosum, 1.8-2.3 mm altum, 1.8-2.7 mm diametro, dense

sericeo-hirsutum, triloculare; styli 3-44 mm longi, basi } mm connatl.

Squamae triangulares usque 0.7 mm longae, 0.3 mm latae, apice acuminatae,

superficie dense trichomatosae, in ramellis, pendunculis, pedicellis, et foliis,

praesertim facie inferiori; inter squamas pili crassiusculi usque 0.3 mm_ longi,

frequenter stellatim collocati.

TYPE: Hoogland 12571, along Kuala Lumpur—Raub road between m.s. 53 and

54 (ie. near The Gap), Selangor, 27 March 1975, in fairly tall secondary forest

on roadside at ca 720 m altitude. HOLOTYPE in CANB, ISOTYPES being

distributed to A, BO, E, G, K, KEP, L, MEL, SING, US.

Further collections examined: Whitmore FRI 2622 (L. SING), Kasim &

Rahim K 750 (UKM), Kunstler 10638 (CAL, K), Teo & Pachiapa 340 = KL

2904 (K, L, SING), Lye L 68/71 (SING), Poore 856 (KLU), Burkill SF 3184

(SING), Stone 6894 (KLU).

114 Gardens’ Bulletin, Singapore — XXX (1977)

Misapplied name: Saurauia tristyla auct.

non DC.: King in J. Asiat Soc. Beng. 59

II (1890) 197, p.p.: Ridl., Fl. Mal. Pen. 1

(1922) 207, p.p.; Burk., Dict. Econ. Prod.

Mal. Pen. (1935) 1967 et ed. 2 (1966)

2002, p.p.

Saurauia fragrans is known only from

a few localities in Selangor: The Gap

(straddling the Selangor—Pahang bound-

ary), Kepong, Ulu Bubong, and Ulu

Gombak, and one in Negri Sembilan:

Gunong Tampir, occurring at elevations

from 75 to 750 m. Outside Malaya it has

also been found in Northern Sumatra.

The specific epithet has been chosen

as, in contrast with other . species of

Saurauia encountered by me in India,

Malaya, Java, Flores and New Guinea,

the flowers of the type specimen were

heavily sweet-scented, a feature made

even more noticeable because of the very

large number of flowers present on the

tree. The fragrance of the flowers was

also noted in Lye L 68/71.

The differences between Saurauia

roxburghii, S. pentapetala, and S. fragans

are set out in the accompanying table.

S. fragrans

1 (-3)-flowered, with at most short peduncle;

at least those below the leaves in cluster of up

to 20.

trichomes on base of and between scales.

S. pentapetala

INFLORESCENCE

1-3 (-5)-flowered, in most plants predominantly

PEDUNCLE AND PEDICELS

1-flowered, with at most short peduncle; at least

those below the leaves in cluster of up to 15.

incorporated in base of scales and sparse or

mostly absent between scales.

S. roxburghii

With tomentum of very short villose hairs and | With broad-based scales, with villose tomentum | With long fairly broad based scales and short

peduncle; at least those below the leaves in

(1-) 3-7-flowered, often open with relatively long

cluster of up to 4.

occasionally a few small elongated scales.

=—_—

115

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116 Gardens’ Bulletin, Singapore — XXX (1977)

II. Saurauia in the Cameron Highlands

During a short visit to the Cameron Highlands in May 1973 I was able to

collect a number of specimens of Saurauia. On closer examination it became clear

that amongst these collections two undescribed species were represented. When IL

tevisited the area in March-April 1975, I took the opportunity of collecting further

specimens of these two species and making notes of their altitudinal occurrence.

Similarly, the altitudinal zonation of the other species in the area and along the access

road from Tapah to Tanah Rata was noted and in part documented by the collect-

ing of herbarium specimens. It appeared that out of ten species now known to

occur in Malaya, eight are found in the area covered. Only S. rubens Ridl. and

S. fragrans Hoogl., both of limited known occurrence, are absent.

altitude in metres

o0osl

°o

009

fragrans

leprosa

mahmudii

Fig. 1. Known alti-

malayana tudinal range of

Saurauia species in

the Cameron High-

napaulensis lands (solid block)

and elsewhere in Ma-

laya (open block).

nudiflora

pentapetala

roxburghii

rubens

vulcani

In fig. 1 the observed altitudinal range in the Cameron Highlands and along

the road from Tapah is presented along the altitudinal range outside the Cameron

Highlands of all species known from Malaya as ascertained from data available of

herbarium specimens. The’ species are arranged in alphabetical order; apart from

those already treated the following are involved:

ie Ah oo -

_ -

117

Malayan Saurauiae

; b. and c. open flower, x 2;

d. flower bud just before opening, x 3; e. longitudinal section of flower, x 2; f. stamens,

Saurauia fragrans Hoogl. a. flowering branch, x

Fig. 2.

x 4, All after Hoogland 12571.

118 Gardens’ Bulletin, Singapore — XXX (1977)

4. Saurauia leprosa Korth., Verh. Nat. Gesch. Kruidk. (1842) 131. — TYPE from

Java.

Synonym: Saurauia cauliflora var. calycina King in J, Asiat. Soc. Beng. 59 II

(1890) 199; Ridl., Fl. Mal. Pen. 1 (1922) 208; Craib, Fl. Siam. Enum, 1 (1925)

127; Henders. in J. Mal. Br. Roy. Asiat. Soc. 17 (1939) 25, 37; Keng in Fl, Thail.

2 (1972) 110. — TYPE: ‘‘Perak: Batu Kurau. Scortechini, No. 1614.”; HOLO-

TYPE in CAL, ISOTYPE in SING.

Misapplied name: Saurauia cauliflora auct. non DC.: Burk. & Henders, in Gard.

Bull. Straits Settl. 3 (1925) 336, 349; Henders. ibid. 4 (1928) 226; Burk., Dict.

Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966) 2002.

This is in Malaya a rather rare mainly lowland species found only in Perak,

Pahang, and Selangor, and occurs outside Malaya in Peninsular Thailand, Sumatra,

and Java. Throughout its range some individuals are almost exclusively cauliflorous,

others are exclusively ramiflorous or carry the inflorescences in the leaf axils, while

few individuals show the whole range of inflorescence positions.

5. Saurauia mahmudii Hoogl., sp. nov. (fig. 3)

Arbor parva usque 6 m alta. Folia obovata vel obovato-oblonga, plerumque

20-30 cm longa, 10-15 cm lata, 20-28-nervata, apice obtusa late acuminata, basi

acuta subdecurrentia, petiolo 24-44 cm longo. Inflorescentiae solitariae vel 2-4

fasciculatae in axillis foliorum vel cicatricium foliorum, pro parte majori dichasia

triflora, nonullae biflorae vel uniflorae, usque 7 cm longae. Pedicellus 2-5 cm

longus. Sepala 9-12 mm longa, 8-10 mm lata, basi 3 mm crassa. Corolla late

urceolata, 13-15 mm longa, 15 mm diametro; tubus 7-9 mm longus, 12 mm

diametro; lobi 8-10 mm longi, 74-11 mm lati, recurvati, apice bilobulati. Stamina

plerumque 25-32, filamentis subaequalibus 5-53 mm longis, antheris versatilibus

dorsifixis 24-3 mm longis. Ovarium semi-globosum 3-34 mm altum, 5 mm diametro,

glabrum, triloculare; styli 7 mm longi, basi 14-2 mm connati.

Squamae anguste triangulares, 0.5-1.0 mm longae, 0.2-0.3 mm latae, acutae,

margine praesertim) inferiori piloso. In ramis, foliis, pedunculis, pedicellis, bracteis,

et bracteolis sparsae; in sepalis absunt, interdum uno vel duabus basi exceptis.

TYPE: Hoogland 12579, along road to summit of Gunong Brinchang near m:s.

46. Cameron Highlands, Pahang, 1 April 1975, in gully on steep slope at ca 1770m

altitude. HOLOTYPE in CANB, ISOTYPES being distributed to BRI, K, KLU,

L, SING, US.

Further collections examined: Whitmore FRI 15449 (KEP), Hoogland 12326

(CANB, K, KEP, L), 12327 (CANB, SING), /2328 (CANB, SING). 12580

(CANB. L), 12581 (A, BO, CANB, KEP), Kasim & Zainudin K 988 (UKM),

Stone 5595 (KLU); all from the Gunong Brinchang area.

The species closely resembles §. nudiflora from which it is readily distinguished

by the glabrous ovary as against densely sericeous. In S. nudiflora nearly all flowers

are solitary though a few 2- or 3-flowered inflorescences occur in most plants; in

S. mahmudii the inflorescences are mostly 3-flowered with fewer 2- or 1-flowered

ones interspersed. Individuals with predominantly 3-locular ovaries are found in

S. nudiflora, though only one such collection is known to me (Hoogland 12570) in

which the flowers are smaller, with a smaller than usual number of stamens for this -

species. In all other collections examined the ovary is predominantly 5-celled, with

occasional flowers with 4- or 6-celled ovary. Var. tetragyna Boerl., originating

in Java, is reported to have a 4-celled ovary. In all collections of S. mahmudii the

ovary is 3-celled (one 2-celled ovary was found in 130 flowers analysed).

I have dedicated this species to Mahmud bin Sider, of the University of

Malaya, who assisted me in collecting in the Cameron Highlands in 1973 and 1975.

Malayan Saurauiae 119

a f VE, a

Camp ?y

iy: se Oa ‘

=4 AY, Dy we tae c =

DAG 4

i¢

Fig. 3. Saurauia mahmudii Hoogl. a. flowering branch, x 4; 5. open flower, x 2;

os Fs eerseeg section of flower, x 2; d. stamens, x 2; e. fruit, x 2. All after Hoogland

120 Gardens’ Bulletin, Singapore — XXX (1977)

6. Saurauia malayana Hoogl., sp. nov. (fig. 4)

Frutex vel arbor parva usque 12 m alta, 20 cm diametro. Folia obovato-

oblonga, plerumque 14-32 cm longa, 6-15 cm lata, 12-20-nervata, apice obtusa

breviter acuminata, basi rotundata vel obtusa, petiolo 14-44 cm longo. Inflores-

centiae solitariae ve] 2-3 fasciculatae in axillis foliorum vel cicatricium foliorum,

dichasiales, 3—20-florae, usque ad 20 cm longae. Pedunculus 1-10 cm longus.

Pedicellus 5-40 mm longus. Sepala 2 exteriora late triangulari-ovata, 8-9 mm

longa, 9-12 mm lata; 2 interiora suborbicularia, 8-10 mm longa, 9-12 mm lata; basi

2 mm crassa. Corolla campanulata, 10-15 mm longa, 16-20 mm diametro; tubus

5-7 mm longus, 10-13 mm diametro; lobi 84-11 mm longi, 9-13 mm lati,

aliquantum reflexi, apice bilobulati. Stamina plerumque 30-40, filamentis

subaequalibus 34-54 mm longis, antheris versatilibus dorsifixis 2-3 mm _longis.

Ovarium semi-globosum 24-34 mm altum, 44-6 mm diametro, dense hirsutum,

(4-) 5 (-7)-loculare; styli 54-74 mm longi, basi 1-24 mm connati.

Squamae anguste triangulares acuminatae, superficie trichomatosae; inter

squamas pili gracilcs stellatim collocati. Squamae ramorum usque 4 mm longae,

i mm latae (plerumque minores), foliorum usque 3 mm longae, pedunculorum et

pedicellorum usque 14 mm longae, faciei exterioris sepalorum exteriorium usque

1 mm longae. Pili graciles usque 1 mm longi.

TYPE: Hoogland 12591, along road from Ringlet to Tanah Rata between m.s.

29 and 30, Cameron Highlands, Pahang, 2 April 1975, in open secondary forest in

gully on steep slope at ca 1150 m altitude. HOLOTYPE in CANB, ISOTYPES

being distributed to A, G, K, KEP, L, SING, US.

Further collections examined: Hoogland 12332 (CANB), 12335 (CANB,

KLU, L), 12576 (A, BO, CANB, E. KEP), 1/2586 (BO, CANB, E, KLU, L,

US), 12587 (CANB, L), 12588 (CANB, L), 12590 (CANB, G, K, SING), 12595

(BH, E, CANB, KLU), Turnau 789 (KLU).

Amongst its Malayan congeners this species stands out by the rather long,

open inflorescences with large flowers. Other species in the area with many-

flowered inflorescences are S. napaulensis with much smaller flowers and S. vulcani

with the flowers in a much more compact inflorescence. Both these species have

generally many more flowers to each inflorescence, and also differ substantially in

the nature of their scales,

Saurauia malayana has been found only in the Cameron Highlands and is

particularly common around the Ringlet between 1100 and 1300 m altitude,

whereas a few trees were also found near Tanah Rata at 1400 m.

7. Saurauia napaulensis DC. in Mém. Soc. Phys. Hist. Nat. Genéve 1 (1822) 421.

— TYPE from Nepal.

This is another widespread species of continental South-East Asia, extending

from north-western India (Himachal Pradesh) to southern China, through Burma,

Thailand, and Indo-China, and now, like S$. roxburghii, recorded for the first time

for Malaya. Compared to that species, it ranges further West in the foothills of the

Himalayas, but not as far East in the Chinese region. While I have no hesitation in

accepting S. roxburghii as indigenous in Malaya, there must be some reservations

about this for S. napaulensis. The species was not collected in the Cameron .

Highlands (the only area in Malaya where it has been found until 1936) but it is

now extremely common, particularly in disturbed areas between Tanah Rata and

Brinchang and on the summit of Gunong Brinchang.

The species is grown as an ornamental in the tea-growing areas of Ceylon and,

in particular, of Southern India and the possibility of its introduction into the

Cameron Highlands for the same purpose cannot be discarded. On the other hand,

Malayan Saurauiae 121

Fig. 4. Sauranuia malayana Hoogl. a. flowering branch, x 4; 6. open flower, x y

c. longitudinal section of flower, x 2; d. stamens, x 2; e. imature fruit, x 2. All after

Hoogland 12591.

122 Gardens’ Bulletin, Singapore — XXX (1977)

it may have occurred there naturally and extended its area substantially with the

opening up of the highland country, The nearest known natural occurrences

outside Malaya are in Thailand on Doi Inthanon in Chieng Mai district and Phu

Miang in Phitsanulok district, some 1500 km to the North.

8. Saurauia nudiflora DC. in Mém. Soc. Phys. Hist. Nat. Genéve 1 (1822) 422,

t. v; King in J. Asiat. Soc. Beng. 59 II (1890) 198; Ridl. in Trans. Linn. Soc.

London II Bot. 3 (1893) 277, in J. Fed. Mal. St. Mus. 4 (1909) 6, et Fl. Mal.

Pen. 1 (1922) 207; Burk. & Henders. in Gard. Bull. Straits Settl. 3 (1925) 336, 349;

Henders., ibid. 4 (1928) 226; Burk., Dict. Econ. Prod. Mal. Pen. (1935) 1967 et

ed. 2 (1966) 2002. — TYPE from Java.

Common in north-western Malaya, where it has been found in Kedah, Perak,

Pahang, and Selangor. Outside Malaya it occurs in Sumatra and Java.

9. Saurauia rubens Ridl. in J. Roy. Asiat. Soc. Straits Br. 61 (1912) 1, et FI.

Mal. Pen. 1 (1922) 208. — TYPE: “Selangor: Semangkok Pass, at the foot of

the hill leading to the Sempang mines.”; HOLOTYPE in SING (Ridley 15771,

‘““Sempang track, Ap. 1911’’), ISOTYPE in BM.

Common in the Fraser’s Hill area, also collected once in Trengganu

(Mandi Angin). Endemic to Malaya.

10. Saurauia vulcani Korth., Verh. Nat. Gesch. Kruidk. (1842) 128. — TYPE

from Sumatra.

Synonym: Saurauia grandis Ridl. in J. Fed. Mal. St. Mus. 4 (1909) 6 et Fl. Mal.

Pen. 1 (1922) 207. — TYPE: ‘“‘Tel6m River banks.”; HOLOTYPE in SING

(Ridley 13580, ‘““‘Telom, Nov. 1908”), ISOTYPES in BM, K.

Apart from its abundant occurrence in the Cameron Highlands, this species

is known from Malaya only from a recent collection from the Genting Highlands.

It spreads readily on disturbed roadsides and on landslides.

The Cyperaceae Tribe Cypereae of Ceylon

TETSuO0 KOYAMA

The New York Botanical Garden, Bronx, New York

Abstract

This taxonomic treatment, revising all the Ceylonese taxa of the Cyperaceae tribe

Cypereae, enumerates 59 species of four genera, Cyperus, Pycreus, Mariscus and Kyllinga. The

following new names are proposed: Cyperus compressus ssp. micranthus, C. diffusus ssp.

macrostachyus, C. nutans ssp. eleusinoides, Pycreus flavidus, P. pumilus ssp. membranaceus,

Mariscus cyperinus ssp. laxatus, M. pedunculatus, M,. sumatrensis, and Kyllinga odorata

ssp. cylindrica.

In the course of my preparation of a taxonomic account of the Cyperaceae for

the Flora of Ceylon, a project under the auspices of the Smithsonian Institution

(Koyama 1970), I have had the opportunity of revising important historical

collections of Ceylonese Cyperaceae that are housed in the Herbarium of the

National Botanic Gardens, Peradeniya. These old collections, of which only a part

is represented at Kew and elsewhere, have not received the attention of specialists

since Trimen published his Handbook (1900). In addition, a series of field expedi-

tions recently sent to Ceylon (Koyama 1970, 1974) resulted in an accumulation

of a substantial number of critical gatherings, among which are several cyperoid

taxa that are new to the flora of Ceylon. In the present paper 59 species of the

tribe Cypereae are recognized as valid for the flora of Ceylon and are classified

under four genera, Cyperus, Pycreus, Mariscus and Kyllinga. The treatment not

only identifies the collections mentioned above but also updates the nomenclature

with particular emphasis on the coordination to the two recent imporant works on

the Cypereae (Kiikenthal 1935-36; Kern 1974). The remarkable richness of

cyperoid taxa in Ceylon rests on the fact that besides the so-called Indo-Malesian

and the Pantropic elements, on this little island there occur Western Asiatic elements

reaching Ceylon as well as those interesting endemic ones which are confined to

the southern part of Deccan Peninsula and Ceylon. Special attention was paid to

these entities of phytogeographic interest.

I appreciate the assistance of Dr. F.. Raymond Fosberg, who has given me

this opportunity of studying Ceylonese Cyperaceae. Messrs R. Cooray, T. Herat,

N. Balakrishnan, A. H. M. Jayasuria and K. Sumithraarachchi certainly deserve

my sincere thanks for their help in the field. The financial assistance from the

Smithsonian Office of International Program is gratefully acknowledged for their

research grant in defraying my field expenses.

KEY TO GENERA

1. Rhachilla not articulated, persistent, hence glumes falling apart from rhachilla

2. Achenes triquetrous or dorsi-ventrally flattened with one side facing

PEE ae eR WE this dw cn in Res Viwddiey tn onan- ss ess. i ARNE TO fo Cyperus

2. Achenes laterally flattened with one angle facing rhachilla ......... Pycreus

1. Rhachilla articulated, hence spikelets falling in entirety

3. Achenes triquetrous with one side facing rhachilla .................. Mariscus

3. Achenes laterally flattened with one angle facing rhachilla ......... Kyllinga

123

124 Gardens’ Bulletin, Singapore — XXX (1977)

CYPERUS L.

1. Stigmas 3, or style more or less undivided (sp, no. 33)

2. Spikelets spicately disposed on conspicuously elongated rhachis, thus

forming spikes

3. Rhachilla of spikelets winged with base of glumes, which is decurrent

along the rhachilla internode

4. Spikelets many to numerous; spikes with long rhachis, cylindrical,

normally 3-7 cm long

5. Leaves with elongated blade

6. Spikelets linear, terete or subterete, 0.5-1 mm wide

7. Spikelets patent to spreading; spikes as a rule peduncl-

ed; leaves 4-15 mm wide; culm smooth on angles

Pe ERC Ta ae EE Penne VET 1 C. digitatus

7. Spikelets erect to suberect; spikes subfastigiate, sub-

sessile; leaves 20-30 mm wide; culms scabrous on

upper,angles 14:03..0-00.n2 zene 2 C. platyphyllus

6. Spikelets lanceolate to lance-oblong, ca, 1.55 mm wide,

flattenéd 27.28.2508). QIAN. AR RA 3 C. exaltatus

5. Leaves reduced to bladeless sheaths ............... 4 C. papyrus

4. Spikelets several to teens, occasionally up to 30; spikes with rather

short rhachis broadly ellipsoid or ovoid, normally 1-4 cm long

8. Plants with creeping rhizomes or stolons (except in C. bulbosus,

which bears tunicate bulb at base of culm)

9. Culms thick, normally more than 5 mm wide and 60 cm

tall, clothed at base with bladeless sheaths

10. Culms obtusely trigonous to subterete; glumes lance-

ovate, acute-tipped; bracts much shorter than corymb

11. Culms not septate ............... 5 C. corymbosus

11. Culms transversely septate ...... 6 C. articulatus

10. Culms acutely triquetrous; glumes oblong, obtuse at

apex; bracts much surpassing corymb ...................+

9. Culms slender, mostly 1-2 mm wide and less than 60 cm

tall; leaves with elongated blade

12. Culms arising from bulbose base, tufted ...............

a pod ea eins 04 Wags csborase skp su aaw) eens ann 8 C. bulbosus

12. Stolons present; culm bases forming a corm-like

enlargement |

13. Glumes 2-2.5 mm long; spikelets slightly swollen;

leaves normally as long as the culm ...............

« « na'ybamcaiene Qanicle eoniesalt ne tama 9 C. stoloniferus

13. Glumes 3-4 mm long; spikelets strongly flattened; .

leaves normally much shorter than the culm

14. Culms 10-30 cm tall; bracts as long as or

shorter than the inflorescence ...............

2D. LEE. ROG 10 C. rotundus

14. Culms 30-60 cm tall; bracts 2 to 3 times

as long as the inflorescence

Ceylon Cypereae

125

15. Spikelets linear-lanceolate, gradually

narrowed to an acute apex, 2.2-3 mm

wide; glumes ovate

16. Glumes 3-3.5 mm _ long, red-

brown, close together not ex-

posing achene between two

glumes; spikelets 2.2-2.5 mm

a 11 C. tuberosus

16. Glumes 4 mm long, yellow-

brown, slightly spaced exposing

portion of achene between two

glumes; spikelets 3-3.2 mm wide

CER er AT aes 12 C. retzii

15. Spikelets linear, rather suddenly

obtusish at apex, 1.5-2 mm _ wide

glumes oblong ......... 13 C. scariosus

8. Plants without creeping rhizomes or stolons

17. Spikelets linear to lanceolate, 15-2 mm wide; weakly

flattened with two acute edges; glumes close together,

acute at apex

18.

Glumes distinctly colored with red-brown, brown or

dark brown; plants perennial with culms more or

less enlarged at base

19. Glumes 2.2-2.5 mm long, pale-green and strong-

ly colored with dark or purple-brown ............

acetate Ae es fade desis n>» <a 14 C. sphacelatus

19. Glumes 3-3.5 mm long, straw-colored and

tinged with reddish brown or brown ............

OE ny 15 C. tenuiculmis

Glumes pale or stramineous-green, not at all brown-

colored; plants annual with fibrous roots only; culms

not thickened at base .................. 16 C. zollingeri

Spikelets filiform, subterete, less than 1 mm thick; glumes

spaced, obtuse at apex

20.

Spikelets spreading, ca. 0.75 mm wide; glumes

rounded-obtuse at apex with moderately narrow

ee eee 17 C. distans

Spikelets erect-patent, ca. 1 mm wide; glumes shallow-

ly emarginate at apex with very broad hyaline mar-

(ER Sp Os 18 C. nutans

3. Rhachilla of spikelets without conspicuous wings, base of glumes not

decurrent along the rhachilla internode

Plants perennial with stolons

22. Rhachis of spikes glabrous; glumes obtuse at white-hyaline

21.

apex

EE M5, OV cca dwsaccanatacvevans 19 C. procerus

22. Rhachis of spikes densely hispid; glumes acute at apex

without broad hyaline apex ......................45 20 C. pilosus

Plants annual, with fibrous roots only

126 Gardens’ Bulletin, Singapore — XXX (1977)

23. Spikelets 1-2.5 cm long, 2-3 mm wide; glumes 2-3.5 mm

long, acute at apex, tightly disposed

24. Glumes 3.5 mm long; achenes 1.5 mm long; inflores-

cence usually open with elongated rays ..................05

i edienetcLial cicette 21b C. compressus ssp. compressus

24. Glumes 2 mm long; achenes 1 mm long; inflorescence

congested in a head-like cluster....0. .::.:2 1:1; «ease

ey eee 21b C. compressus ssp. micranthus

23. Spikelets 0.5-1 cm long, 1-1.5 mm wide; glumes 0.75—1.25

mm long, obtuse at apex, loosely disposed ......... 22 C. iria

2. Spikelets digitately disposed or capitately congested at apices of inflores-

cence rays, thus forming heads without rhachis

25. Inflorescences open with elongated rays

26. Large-sized perennials with conspicuous rhizome; at least some

leaves and bracts more than 10 mm wide; culms 30-90 cm tall

27. Leaves with elongated blade; leafy bracts 5-10

28. Achenes with acute angles; spikelets in clusters of 2 or

3, sometimes solitary, weakly compressed

28a. Spikelets 4-6 mm long, up to 12-flowered ......

NitttUaaesieye. cade 23a C. diffusus ssp. diffusus

28a. Spikelets 10-20 mm long, up to 40-flowered ......

qui kh tains 23b C. diffusus ssp. macrostachyus.

28. Angles of achenes spongy-thickened; spikelets in

clusters of 5 to 8, strongly compressed ..................

ae alah « caeiiaclhs 5o. ait ania ie 24 CC. platystylis

27. Leaves reduced to bladeless sheaths, leafy bracts more than

1S: SRR eee 25 C. alternifolius ssp. flabelliformis

26. Small to medium-sized annuals or short-lived perennials; leaves

and bracts at most 5 mm wide; culms up to 40 cm tall

29. Umbel rays longer than or nearly as long as the culm ......

LL SLE RS ae FB ND os iacaeteseecene: a an

29. Umbel rays much shorter than the culm

30. Glumes awnless; achenes obovate; leaves flattish

31. Spikelets digitate in clusters of 3-15; glumes

acutish at apex

32. Roots brownish; basal sheaths straw-brown

or pale-brown; spikelets dark green .........

een, ee eS 27 C. pulcherrimus

32. Roots purplish; basal sheaths purple-brown;

spikelets brown or purple-brown

Ceylon Cypereae 127

33. Glumes densely imbricate and close

together so that achenes not shown

between glumes; rhizome often con-

spicuous, rather frequently elongated

op te a a. Be 28 C. haspan

33. Glumes sublaxly imbricate and slightly

spaced, thus achenes exposed between

glumes; plants tufted with fibrous roots

Cy ee ee 29 C. tenuispica

31. Spikelets many, congested in globose heads ......

eee. “a EE hs ee Ss eee 30 C. difformis

30. Glumes awned at apex; achenes oblong; leaves

canaliculate

34. Glumes with straight awn ...... 31 C. castaneus

34. Glumes with recurved awn ...... 32 C. cuspidatus

25. Inflorescences congested a single head

35. Style 3-cleft; achenes acute on angles

36. Rhizome with long creeping stolons; culms 5-20 cm tall;

hess \1—2 om wide 2.4.2.4 su e..:. 33 C. arenarius

36. Rhizome short, culms tufted, 20-60 cm tall; heads more

Watt cml acres “lS... 34 C. conglomeratus

35. Style scarsely divided at apex; achenes with spongy-thickened

angles; plants floating on water .................. 35 C. cephalotes

1. Stigmas 2

37. Tall perennial; culms ca. 100 cm tall; inflorescences umbelliform with

elongated rays up to 18 cm long ........................... 36 C. alopecuroides

1. Cyperus digitatus Roxburgh, [Hort. Beng. 81, 1814, nomen, &] Fl. Indica

ed. 1, 1: 209, 1820; C. B. Clarke, Fl. Brit. India 6: 618, 1893; Trimen & Hook.

f., Handb. Fl. Ceylon 5: 36, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 55,

1935; Kern, Fl. Males. I, 7 (3): 601, 1974.

‘Cyperus venustus R. Br.’ sensu Nees, Contrib. Bot. India 86, 1834.

‘Cyperus neesii Kunth’ sensu Thwaites, Enum. Pl. Zeyl. 344, 1868. ——- Cyperus

hookeri Bockeler, Linnaea 36: 308, 1870. ——- Cyperus digitatus Roxb. var.

hookeri (Béckir.) C. B. Clarke, Fl. Brit. India 6: 618, 1893; Trimen & Hook. f.,

Handb. Fl. Ceylon 5: 37, 1900.

Jaffna District: Ca. 5 miles W of Paranthan, south shore of Jaffna Lagoon,

T. Koyama et al. 14043 (NY, PDA, US); Paranthan, Clayton 5263 (K, PDA).

Anuradhapura District: Wilpattu National Park, East Intermediate Zone, along

128 Gardens’ Bulletin, Singapore — XXX (1977)

<< rr ti “‘_OS:;~(‘(‘i~S

maton Hey _

Fig.l. Cyperus platyphyllus Romer & Schultes. AA, habit. B, spikelet with its prophyll.

C, glume. D, dorsal view of achene. E, ventral view of achene with style. Scales = 1 mm.

Ceylon Cypereae 129

the Wildife Dept. trail, ca. 2 miles from barrier gate, T. Koyama & Jayasuria 13965

(NY, PDA, US). Kandy District: Ramboda, Thwaites C. P. 3043 ex p. (K, PDA):

Pusselawa, Thwaites C. P. 3043 ex p. (PDA). Colombo District: Ferguson s.n.

(PDA).

Distribution. Pantropic with its range extending north into southern China

and Formosa; abundant in South America and southern Asia. relatively scarce in

tropical Africa and Australia.

Plants from India and Ceylon have often been treated as var. hookeri, which

was segregated by its more copiously spiculose spikes. My observation of this

species on a world-wide basis does not support this variety.

2. Cyperus platyphyllus Romer & Schultes, Syst. Veg. 2: 876, 1817: C. B.

Clarke, Fl. Brit. India 6: 618, 1893, & Illustr. Cyper. ¢. 20, f. 3-4, 1909: Trimen &

Hook. f., Handb. Fl. Ceylon 5: 38, 1900; C. E. C. Fischer in Gamble, Fl. Madras

9: 1642, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 59, 7. 8&7, 1935. — Fig. 1.

“Cyperus elatus L.,” sensu Roxburgh, Fl. Indica ed. 1, 1: 207, 1820.

Cyperus roxburghii Nees in Wight, Contrib. Bot. India 84, 1834; Thwaites, Enum.

Pl. Zeyl. 70, 1864. Cyperus eminens Klein ex Kunth, Enum. Pl. 2: 70,

Papyrus latifolia Willdenow. Abhandl. Akad. Berlin 1812/13:

Trincomalee District: Trincomalee, Glenie in 1864 (PDA). Matara District:

Kukul Corale, Thwaites C. P. 304] ex p. (K. PDA); Matara, Trimen 23 Feb. 1881

(PDA). Amparai District: Mile 205 on Rd. A 4,ca. 3 miles N of Pottuvilp,

T. Koyama et al. 14011 (NY, PDA, US).

Distribution. Confined to southern India and Ceylon. Marshy places at low

altitudes.

This rather rare species is easily discernible by its sessile or nearly sessile

spikes forming digitate secondary corymbs, and very broad leafy bracts and leaf

blades. The distribution range of this species, covering the southern part of Indian

Peninsula and Ceylon only, shows the same pattern as those of Mariscus clarkei,

Fimbristylis tenuifolia, Carex lobulirostris and Carex leucostachya.

3. Cyperus exaltatus Retzius, Obs. Bot. 5: 11, 1789: Thwaites, Enum. Pl. Zeyl.

343, 1864: C. B. Clarke, FI. Brit. India 6: 617, 1893: Trimen & Hook. f.. Handb.

Fl. Ceylon 5: 32, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 64, f. 9 A-F,

1935; Kern Reinwardtia 2: 99, f. 2, 1952 & Fl. Males. L, 7 (3): 602, 1974.

Cyperus venustus R. Brown, Prodr. Fl. Nov. Holl. 217, 1810; Thwaites, Enum.

Pl. Zeyl. 432, 1864. Cyperus altus Nees in Wight, Contrib. Bot. India 84,

1834. Cyperus exaltatus Retz. var. amoenus C. B. Clarke, Journ. Linn. Soc.

21: 187, 1886; Trimen & Hook. f., Handb. Fl. Ceylon 5: 33, 1900.

Jaffna District: Jaffna, Silva 110 (PDA); Murukan, Silva in 1917 (PDA).

Vavunia District: Ilukkulama, ca. 6 km SW of Vavunia, 7. Koyama & Herat

13603 (NY, PDA, US). Anuradhapura District: Anuradhapura, at margin of

Nuwara Wewa Tank, T. Koyama et al. 13939 (NY, PDA, US); Anuradhapura,

Brodin C. P.3040 ex p. (PDA): Minneriya, ca. 15 miles E of Habarana, T. Koyama

et al. 13568 (NY, PDA, US). Mannar District: ca. 10 miles S E of Mannar, 7.

Koyama et al. 13922 (NY, PDA, US). Polonnaruwa District: Yoda Ela, Silva in

1905 (PDA); Mannampitiya, ca. 5 miles ESE of Polonnaruwa, 7. Koyama &

Herat 13572 (NY, PDA, US). Kandy District: Peradeniya, Trimen C.P. 3040 in

part (PDA). Badula District: ca. 8 miles E of Mahiyangana, 3.5 miles N. of Mile

53 on Mahiyangana-Padiyatalam Rd., T. Koyama et al. 13980 (NY, PDA, US).

Trincomalee District: Trincomalee, Glenie C.P. 3788 (PDA). Batticaloa District:

Batticaloa, Trimen C.P. 3040 in part (PDA). Monaragala District: E of south end

of Jayanthiwewa, T. Koyama et al. 13988 (NY, PDA, US). Ratnapura District:

130 Gardens’ Bulletin, Singapore — XXX (1977)

Fig. 2. Cyperus pangorei Rottboell. AA, habit. B, portion of spikelet showing two glumes

and their bases forming wings on the rhachilla. C, spikelet. D, prophyll. E, glume. F & G,

achenes. H & J, prophylls at the base of umbel ray, Scales = 1 mm.

Ceylon Cypereae 131

Ratnapura, Trimen C.P. 3040 in part (PDA); Ranwala, Amaratunga 670 (PDA).

Kurunegala District: Hettipola, Amaratunga 670 (PDA); Uhuniya, Amaratunga

182 (PDA). Ruhuna National Park: Andunoruwa Wewa, Cooray 16 Dec.

1969 (US).

Distribution. Pantropic, with its range extending northwards to eastern China

and Japan. Rare in South America, In Ceylon very common in wet places at low

altitudes, and often noted in rice fields.

4. Cyperus papyrus L., Sp. Pl. ed. 1, 47, 1753.

Kandy District: cultivated in the Botanic Gardens, Peradeniya, Amaratunga

197 (PDA). Native of East Tropical Africa with a subspecies in Madagascar.

Widely cultivated in Ceylon as an ornamental.

5. Cyperus corymbosus Rottboell, Descr. Icon. Rar. Nov. Pl. 42, t. 7 f. 4, 1773;

' Thwaites, Enum. Pl. Zeyl. 344, 1864; C. B. Clarke, Fl. Brit. India 6: 612, 1893;

Trimen & Hook. f., Handb. Fl. Ceylon 5: 29, 1900; C. E. C. Fischer in Gamble,

Fl. Madras 9: 1641, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 80, f. 10, 1936.

Anuradhapura District: between Ramboda and Madawachchiya, 7. Koyama

& Herat 13602 (NY, PDA, US). Matale District: between Naula and Dambulla,

vicinity of Mile 40, Jayasuria 56 (US, PAD). Kandy District: Peradeniya,

Amaratunga 199 (PDA). Kegalle District: Baddewela, T. Koyama & Samarakoon

13546 (NY, PDA, US). Badula District: 5 miles WNW of Mahiyangana, Davidse

7595 (MO, NY, PDA). Batticaloa District: Wandeloos Bay, T. Koyama & Herat

13579 (NY, PDA, US). Monaragala District: E of south end of Jayanthiwewa,

T. Koyama et al. 13987 (NY, PDA, US). Galle District: Moragoda, left bank of

Bentota Ganga, Amaratunga 2357 (PDA). Puttalam District: Irranonillu Madampe,

Amaratunga 2574 (PDA).

Distribution. Tropical West Africa, Madagascar. Nepal, India, Burma,

northern Australia, West Indies and tropical South America (Venezuela and

Brazil). Marshy places at low altitudes.

In Ceylon this species called Gal-éhi is occasionally more or less cultivated,

because its straight culms are preferred by natives as material for making a kind

of rough mats. The culms of Scirpus grossus L.f. are also used for the same purpose

but is not cultivated.

6. Cyperus articulatus L., Sp. Pl. ed. 1, 44, 1753; Thwaites, Enum. Pl. Zeyl. 343,

1864; C. B. Clarke, Fl. Brit. India 6: 611, 1893; Trimen & Hook. f., Handb. FI.

Ceylon 5: 29, 1900; C. E. C. Fischer in Gamble, Fl. Madras 9: 1641; Kiikenthal,

Pflanzenr. 4 (20), 101 Heft: 77, 1936.

Polonnaruwa District: Kentalai, Trimen in 1885 (PDA). Batticaloa District:

Batticaloa, Thwaites C.P. 3561 (K, PDA).

Distribution. Mediterranean Region, Tropical Africa, India, Indo-China,

southeastern U.S.A. to Central and South America. More frequent in the Neotropics.

It is rather interesting that this species has never been collected since Trimen’s

time.

7. Cyperus pangorei Rottboell, Descr. Icon. Rar. Nov. Pl. 31, ¢. 7 f. 3, 1773;

Thawaites, Enum. Pl. Zeyl. 344, 1864; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 76,

1936. Fig. 2.

Cyperus tegetum Roxburgh, Fl. Indica ed. 1, 1: 208, 1820; C. B. Clarke, FI.

Brit. India 6: 613, 1893, & Illustr. ¢. 17, f. J, 1909. Cyperus dehiscens

(Nees) Nees, Linnaea 9: 286, 1835; Kunth, Enum. Pl. 2: 56, 1837; Trimen & Hook.

f. Handb. Fl. Ceylon 5: 30, 1900. Papyrus pangorei (Rottb.) Nees in Wight,

Contrib. Bot. India 88, 1834, in major part, concerning basionym. Papyrus

dehiscens Nees in Wight, l.c. 89, 1834.

132 Gardens’ Bulletin, Singapore — XXX (1977)

Anuradhapura District: ca. 25 miles W of Trincomalee, along Rd. A 12,

T. Koyama et al. 14056 (NY, PDA, US); 2 km N of Yakalla, T. Koyama 13595

(NY). Monaragala District: road between Maha Oya and Mullajama at Mile 14,

T. Koyama et al. 13981 (NY, PDA, US); Kotagoda, Siyambaladuwa to Inginiya-

gala, Clayton 5825 (K, PDA). Colombo District: Ja-al on Gampaha Rd., Comanor

1026 (US).

Distribution. Ceylon, India, Nepal and Burma. According to Kiikenthal (op.

cit.) also cultivated in Mauritius for mat-making.

Like C. malaccensis and C. corymbosus this species provides material for

making mats. Trimen (op. cit.) mentions that in the Dambara area along Mahaweli

Ganga this species is exclusively used for this purpose, and the mats are named

after the locality.

8. Cyperus bulbosus Vahl, Enum. Pl. 2: 342, 1806; C. B. Clarke, Fl. Brit. India

6: 611, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 22, 1900; C. E. C. Fischer

in Gamble, Fl. Madras 9: 1641, 1931; Kukenthal, Pflanzenr. 4 (20), 101 Heft: 125,

f. [5 C-E, 1936; Kern, FI. Males. 1, 7 (3): 605, 1974.

Puttalam District: Karapitiya, Kundu & Balakrishnan 363 (US); Chilaw,

Ferguson, Nov. 1881 (PDA). Trincomalee District: beach S of Elizabeth Point,

sea level, Davidse 7523 (MO, NY, PDA, US). Hambantota District: Ruhuna

National Park, Patangala, beach, Cooray 16 Nov. 1969 (NY); Block III, 2 miles E.

of Vaddangewardiya, N of Kataragama, Wirawan 658 (NY).

Distribution. Tropical Africa, southern Asia, Malesia and northern Australia.

Sandy beach; in Ceylon rather scarce.

In southern Asia the bulbs of this species is sometimes eaten by natives.

9. Cyperus stoloniferus Retzius, Obs. Bot. 4: 10, 1786; C. B. Clarke, Fl. Brit.

India 6: 615, 1893 & Illustr. Cyper. t. 79 f. 1-3, 1909; Trimen & Hook. f., Handb.

Fl. Ceylon 5: 36, 1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 106, 1935; Blake,

Univ. Queens]. Papers 2 (2): 9, ¢. 7, 1942; Kern, Fl. Males. I, 7 (3): 606, f. 50,

1974.

Jaffna District: Punkudutivu, lagoon edge, Kundu & Balakrishnan 670 (US).

Mannar District: End of the Causeway at Mile 145, T. Koyama et al. 13923 (NY,

PDA, US). Puttalam District: Puttalam Lagoon, ca. 6 miles N of Mampuri, T.

Koyama et al, 13912 (NY, PDA, US); Pallugaturai. W of Wilpattu National Park,

Davidse et al, 8217 (NY, PDA, US). Trincomalee District: Irrakkakandi, N of

lagoon, ca. 12 miles NNW of Trincomalee, T. Koyama et al. 14067 (NY, PDA,

US). Colombo District: Negombo, Simpson 7926 (PDA); Mount Lavinia, Trimen

in 1981 (PDA); Kalagedihena, Amaratunga 1406 (PDA).

Distribution. Madagascar, Mauritius, India, Indo-China, Malesia and Australia.

In Ceylon rather frequent at margins of brackish lagoons, often forming a large

community.

10. Cyperus rotundus L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 343,

1864; C. B. Clarke, Fl. Brit. India 6: 614, 1893; Kiikenthal, Pflanzenr. 4 (20), 101

Heft: 107, f. 13, 1935; S. T. Blake, Univ. Queensl. Papers 2 (2): 8, 4. 4, 1942;

Kern, Fl. Males. I, 7 (3): 604, f. 49, 1974.

; Jaffna District: Jaffna, Gardner C.P. 804, ex p. (PDA). Anuradhapura

District: Minneriya Tank, Thwaites C.P. 804 ex p. (K, PDA); Anuradhapura,

Sinniah in 1926 (PDA). Polonnaruwa District: Kanniyai, Ramanathan 421

(PDA); Mile 79 on Rd. A 15, 1 m alt., Davidse 7568 (MO, NY). Kurunegala

District: Melsiripura, Amaratunga 1333 (PDA). Matale District: Nalande, Alston

640 (PDA). Kandy District: Peradeniya, Thwaites C.P. 804 in part (PDA);

Kadugannawa, Alston 1089 (PDA); between Balana and Alagalla, Comanor 1187

Ceylon Cypereae 133

(PDA, US); Haragama, Alston in 1926 (PDA); Gannoruwa, Alston 300 (PDA);

Katugastota, Amaratunga 1088 (PDA).

Distribution. A cosmopolitan weed seen in tropical, subtropical and temper-

ate regions of all the world.

11. Cyperus tuberosus Rottboell, Descr. Icon. Rar. Nov. Pl. 28, t. 7 f. 1, 1773;

C. B. Clarke, Fl. Brit. India 6: 616, 1893, in part.

Cyperus rotundus L. subsp. tuberosus (Rottb.) Kiikenthal, Pflanzenr. 4 (20),

101 Heft: 113, 1936.

Trincomalee District: Trincomalee, Ramanathan s.n. (PDA). Polonnaruwa

District: Polonnaruwa, Alston 292 (PDA). Mannar District: opposite to Mannar,

Mile 145 on A 14, end of the causeway, JT. Koyama et al, 13924 (NY, PDA, US);

along Rd. A 14, Mile 123, near the junction with Madhu Rd., 7. Koyama et al.

13918 (NY, PDA, US). Vavuniya District: Palayanalankulan, Mile 120 on Rd.

A 14, T. Koyama et al. 13936 (NY, PDA, US). Matale District: between Dambula

and Habarane, at Mile 96 on Rd. A 6, T. Koyama & Herat 13563 (NY, PDA, US);

vicinity of Mile 40, between Naula and Dambula, Jayasuria 59 (PDA).

Distribution. Eurasia,

12. Cyperus retzii Nees in Wight, Contrib. Bot. India 82, 1834.

Cyperus rotundus L. subsp. retzii (Nees) Kiikenthal, Pflanzenr. 4 (20), 101

Heft: 114, 1935.

Central Province: without definite locality, Thwaites C.P. 3750 (PDA).

Distribution. So far known from India, Ceylon and Australia; rare.

Thwaites’ C.P. 3750 cited above is the only Ceylonese collection of this species.

In this specimen relatively broad spikelets ranging from 2.8 to 3.25 mm in width

bear lance-ovate glumes which have two closely situated parallel veins on both

sides of the mid-nerve and attain as much as 4 mm in length. Its achenes, average

being 1.25 mm long, tend to be slightly larger than those of C. rotundus. Because

of these characters the identity of C.P. 3750 seems certain though I have not seen

as yet the type of C. retzii.

13. Cyperus scariosus R. Brown, Prodr. 216, 1810; C. B. Clarke, Fl. Brit. India 6:

612, 1893 & Illustr. Cyper. t. 75 f. 2-3, 1909; S. T. Blake, Univ. Queensl. Papers

2 (2): 9, t. 2, 1942: Kern, Reinwardtia 2: 103, f. 3, 1952 & Fl. Males. I, 7 (3):

607. 1974. j

Cyperus corymbosus Rottb. var. scariosus (R. Br.) Kiikenthal, Pflanzenr. 4

(20), 101 Heft: 83, 1935. Cyperus mitis Steudel, Synops. Pl. Glumac. 2: 316,

1855; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 103, 1935; Abeywickrama, Ceylon

Journ. Sci., Biol. Sci., 2 (2): 134, 1959.

Jaffna District: between Poonaryn and Elephant Pass, along Jaffna Lagoon,

T. Koyama et al. 14048 (NY, PDA, US); Jaffna, Silva in 1920 (PDA); near

Ampan, Clayton 5235 (K, PDA); Keerimalai, Point Pedro, Clayton 5203 (K, PDA);

Puttalam District: between Puttalam and Kurunegala, Trimen Aug. 1883 (PDA);

Madampe, Amaratunga 2577 (PDA). Matale District: between Dambula and

Habarane at Mile 96 on Rd. A 6, T. Koyama & Herat 13564 (NY, PDA, US);

Dambula, Thwaites C.P. 3966 (K, PDA). Hambantota District: Tissamaharama,

Trimen Dec. 1882 (PDA).

Distribution. Madagascar, India to Indo-China, Malesia, Australia.

Among very confusing nut-grasses C. scariosus can be easily recognizable by

its filiform, obtuse-tipped spikelets which are only 1.5 to 2 mm wide. In Ceylon

I saw this species growing together with C. tuberosus and C. rotundus respectively

in two habitats, where I have seen no evidence of hybrids with the latter species.

134 Gardens’ Bulletin, Singapore — XXX (1977}

Fig. 3. Cyperus tenuiculmis Steudel. A, habit. B, spikelet. C, prophyll at the base

of ame D, glume. E, lateral view of glume showing its hyaline extension of the base.

F, achene

Cyperus zollingeri steudel. G, cluster of spikelets. H, spikelet. J, portion of spikelet

showing glumes and achenes. K, prophyll. L, glume, M, achene. = | mm.

Ceylon Cypereae 135

14. Cyperus sphacelatus Rottboell, Descr. Pl. Rar. Progr. 21, 1772 & Descr.

Icon. Rar. Nov. Pl. 26, 1773. Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 129, 1935;

Kern, Reinwardtia 2: 107, f. 5, 1952, & Fl. Males. I, 7 (3): 609, 1974.

Kandy District: Peradeniya, Univ. of Ceylon Campus, 500 m alt., Comanor

321 (NY, US); ca. 5 miles SE of Gampaha, Mile 18/6 on Rd. A 5, 720 m alt.,

Davidse et al. 7922 (MO, NY, PDA, US). Colombo District: Horana, Alston

976 (PDA).

Distribution. Tropical America and tropical Africa; introduced to Ceylon,

Malesia, Queensland and Tahiti.

The introduction of this species into Ceylon seems pretty old as it was already

collected by Alston in the beginning of 1900.

15. Cyperus tenuiculmis Bockeler, Linnaea 36: 286, 1870; Kern, Reinwardtia 3:

30, 1954, & Fl. Males. I, 7 (3): 608, 1974. Fig. 3 A-F

“Cyperus zollingeri Steudel”’ sensu C. B. Clarke, Fl. Brit. India 6: 613, 1893,

& Illustr. Cyper. t. 8 f. 1-2, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 35,

1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 133, 1935.

Kandy District: Hantana, Mueller-Dombois et al. June 18, 1968 (PDA).

Nuwara-Eliya District: Ramboda, 1180 m alt., T. Koyama 13620 (NY); Hakgala,

Mueller-Dombois et al. Jan. 1968 (PDA, US). Badulla District: ca. 8 miles E of

Mahiyangana, 3.5 miles N of Mile 53 on Mahiyangana-Pediyatalawe Rd., T.

Koyama et al. 13977 (NY, PDA, US); Diyatalawa, Mueller-Dombois et al. Jan.

14, 1968 (PDA). Monaragala District: E of the south end of Jayanthiwewa Tank,

T. Koyama et al. 13993 (NY, PDA, US).

Distribution. Throughout the tropics of the Old World with the range extending

northwards to southern Japan in eastern Asia.

16. Cyperus zollingeri Steudel [in Zollinger, Syst. Verz 1: 62, 1854, name only, &]

Synops. Pl. Glumac. 2: 17, 1855: Kern, Reinwardtia 3: 28, 1954 & Fl. Males. I, 7

(3): 608, 1974. Fig. 3 G-M

Cyperus ramosii Kiikenthal in Fedde, Repert. Sp. Nov. 21: 326, 1925, &

Pflanzenr. 4 (20), 101 Heft: 136, 1935; Kern, Reinwardtia 2: 109, f. 7, 1952.

Hambantota District, Ruhuna National Park: Yala Dunes, Ecology Project

Plot R26, Cooray 8 Dec. 1969 (NY, PDA, US); Block I, at Rugamtota, Mueller-

Dombois et al. 7 Mar. 1969 (NY, US); Kumbukkan Oya, ca. 2 miles above mouth,

at Magabakanda Meda Duwa, Block II, Fosberg et al 51107 (NY, PDA, US).

Distribution. Tropical Africa, Madagascar, Malesia, northern Australia. In

Ceylon rare in open grassy places on sandy soil.

New to the flora of Ceylon. This species differs from the better known

C. tenuiculmis by the softer leaves and bracts, annual tufted habit without stolons,

and smaller floral parts as illustrated in Fig. 3.

17. Cyperus distans L. f., Suppl. Sp. Pl. 103, 1781; Thwaites, Enum. Pl. Zeyl. 344,

1864; C. B. Clarke, Fl. Brit. India 6: 607, 1893; Trimen & Hook. f., Handb. FI.

Ceylon 5: 30, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 137, 1935; Kern, FI.

Males. I, 7 (3): 610, 1974.

Polonnaruwa District: W of Siva Devale in Sacred Area Section 2A, Ripley

369 (PDA). Kegalle District: Kadugannawa, Grupe 2/3 (PDA, US). Nuwara

Eliya District: Hakgala Botanic Garden, Clayton 5784 (K, PDA); Maturata,

Ferguson in 1906 (PDA). Kandy District: Peradeniya, Thwaites C.P. 810 ex p.

(K, PDA); Gannoruwa, Alston 319 (PDA); ca. 2 miles E. of Maddakele, slope

N.W. of Kunckles Peak, 1440 m alt., Davidse 8320 (MO, NY, PDA, US).

Batticaloa District: Batticaloa, Thwaites C.P. 810 in part (PDA).

Distribution. Pantropic.

136 Gardens’ Bulletin, Singapore — XXX (1977)

18. Cyperus nutans Vahl, Enum. PI. 2: 363, 1806; C. B Clarke, Fl. Brit. India 6:

607, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 31, 1900; C. E. C. Fischer in

Gamble, Fl. Madras 9: 1640; 1931; Kukenthal, Pflanzenr. 4 (20), 101 Heft: 144,

f. 5 A-D, 1935; Kern, Fl. Males. I, 7 (3): 609, 1974.

Cyperus distans L. var. major Thwaites, Enum, Pl. Zeyl. 432, 1864.

18a. Subsp. nutans.

Anuradhapura District: Anuradhapura, Thwaites in 1881 (PDA); Wilpattu

National Park, ca. 2.7 miles to Ranger’s Office along pass from southern entrance,

T. Koyama & Jayasuria 13967 (NY, PDA, US). Trincomalee District: Trinco-

malee, Brodie C.P. 3844 (PDA). Matale District: Dambulla, Thwaites C.P. 3966 in

part (PDA). Colombo District: Galapitamada, Amaratunga 1148 (PDA). Monara-

gala District: E of south end of Jayanthiwewa Tank, T. Koyama et al. 13992 (NY,

PDA, US). Hambantota District: Ruhuna National Park, Yala Plain, Comanor

863A (NY, US).

Distribution. India. southern China and Malesia.

18b. Subsp. eleusinoides (Kunth) T. Koyama, stat. nov.

Cyperus eleusinoides Kunth, Enum. Pl, 2: 29, 1937; C. B. Clarke, FI. Brit.

India 6: 608, 1893; Kiukenthal, Pflanzenr. 4 (20), 101 Heft: 144, 1936.

Cyperus nutans Vahl var. eleusinoides (Kunth) Haines, Bot. Bihar Orissa 5: 898,

1924: Kern, Fl. Malesiana I, 7 (3): 610, 1974.

Kandy District: Haragama, Trimen C.P. 3044 (PDA). Hambantota District:

Ruhuna National Park, Mueller-Dombois et al. Jul. 1967 (PDA).

Distribution. Tropical Africa, India to Malesia and northern Australia, also

extending northwards to the Ryukyus.

Cyperus nutans and C. eleusinoides are very similar to one another, and the

only reliable difference between the two is the more densely disposed, longer

spikelets in the latter. But, this difference becomes obscure as ssp. subprolixus of

the Ryukyus and Taiwan comes between the two as to the density and the number

of florets in the spikelet. I, therefore, regard C. eleusinoides as a subspecies of

C. nutans.

19. Cyperus procerus Rottboell, Descr. Icon. Rar. Nov. Pl. 29, ¢. 5 f. 3, 1773;

Thwaites, Enum. PJ. Zeyl. 343, 1864; C. B. Clarke, Fl. Brit. India 6: 610, 1893;

Trimen & Hook. f., Handb. Fl. Ceylon 5: 34, 1900; Kiikenthal, Pflanzenr. 4 (20),

101 Heft: 91, 1935; Kern, Fl. Males. I, 7 (3): 611, f. 51, 1974. Fig. 4

Jaffna District: Murunkan, Silva in 1917 (PDA). Vavunia District: Vavunia,

Clayton 5306 (K, PDA); near Kokkavil, Clayton 5292 (K, PDA). Anuradhapura

District: Anuradhapura, Trimen Dec. 1884 (PDA); Alankulam, Trimen Oct. 1883

(PDA); Mile 123 on Rd. A 14, E of Medachchiya, T. Koyama et al. 13921 (NY,

PDA, US). Wilpattu National Park: Kuda Pathessa, T. Koyama & Herat 13394

(NY, PDA, US); Periya Naga Villu, Wirawan et al. 903 (US); Manikepola Uttu,

T. Koyama 13461 (NY, PDA, US); Wilpattu West Sanctuary, Mari Villu, 7.

Koyama 13972 (NY, PDA, US). Batticaloa District: Batticaloa, Thwaites C.P.

3752 (K, PDA); Ottawady, opposite to Valaichchenai, T. Koyama & Herat 13582

(NY, PDA, US). Mannar District: Murunkan, Silva in 1917 (PDA); Illupadichcha-

nai, Lord s.n. (PDA). Puttalam District: Madampe, Amaratunga 2575 (PDA).

Matale District: Dambulla, Thwaites C.P. 3752 in part (K, PDA); between Naula

and Dambula, vicinity of Mile 40, Jayasuria 60 (PDA). Kandy District: Siyam-

balatenna, Alston 390 (PDA); Peradeniya, Thwaites C.P. 802 (PDA), Alston

118I (PDA). Kurunegala District: Kurunegala, Thwaites C.P. 3752 in part (PDA).

Colombo District: Colombo, Ferguson s.n. (PDA). Hambantota District: Tissa-

maharama, Alston 1179 (PDA); Ruhuna National Park, Block II, 7 m alt.

Ceylon Cypereae 137

Sra 44

Fig. 4. Cyperus procerus Rottboell. AA, habit. B, spikelet. C, prophyll. D, glume.

E, achene. Scales for floral parts = | mm.

138 Gardens’ Bulletin, Singapore — XXX (1977)

Comanor 1037 (PDA), US). Amparai District: Helawe Eliya, ca. 7 miles S. of

Panama, E. of Helawe Lagoon, 7. Koyama et al. 14024 (NY, PDA, US).

Distribution, India, Ceylon, Indo-China, Taiwan, Malesia, Australia (Queens-

land). In Ceylon abundant in marshy places at low altitude.

20. Cyperus pilosus Vahl, Enum. Pl. 2: 354, 1806; Thwaites, Enum. Pl. Zeyl. 344,

1864; C. B. Clarke, Fl. Brit. India 6: 609, 1893; Trimen & Hook. f., Handb. FI.

Ceylon 5: 32, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 92, 1935; Kern, FI.

Malesi I, 7 (3): 611, f. 52, 1974.

Cyperus marginellus Nees in Wight, Contrib. Bot. India 83, 1834.

Cyperus obliquus Nees in Wight, I.c. 86, 1834.

Nuwara Eliya District: between Pussellawa and Ramboda, Mile 32/8 on

Nuwara-Eliya-Kandy Rd., 1000 m alt., 7. Koyama 13617 (NY, PDA, US). Badulla

District: Haptale, Silva in 1906 (PDA). Kandy District: Rangala, Ferguson Sept.

1885 (PDA). Kalutara District: Korosduwa, Amaratunga 2541 (PDA); Kande

Kanda, Amaratunga 2523 (PDA). Kegalle, District: Kegalle, Amaratunga 1651

(PDA); Kattiyakumbura, Amaratunga 1589 (PDA); Nayinankada, 700 m alt.,

T. Koyama & Samarakoon 13561 (NY, PDA, US). Kurunegala District: Nara-

munala, Amaratunga in 1966 (PDA); Malsiripura, Amaratunga 1330 (PDA).

Ratnapura District: 11 miles E of Deniyaya at Mile 62 on Rd. A 17, 750 m alt.,

Davidse 7876 (MO, NY, PDA, US). Galle District: Akeemana, Alston 1187

(PDA); Moragoda, Amaratunga 2359 (PDA). Monaragala District: Inginiyagala

National Park, between Baduluwila and ““Westminster Abbey”, T. Koyama et al.

13994 (NY, PDA, US); E of south end of Jayanthiwewa, T. Koyama et al. 13988

(NY, PDA, US).

Distribution. Rather widely distributed in tropical and subtropical regions of

the Old World from tropical West Africa through Central Asia and Indian Sub-

Continent eastwards to Malesia and northern Australia, In eastern Asia the range

extending northwards as far as to warm-temperate Japan.

Although this species is highly variable in the dimension of leaves and inflo-

rescences, it is well demarcated by the subdensely hispid rhachis of spikes.

21. Cyperus compressus L., Sp. Pl. ed. 1, 46. 1753; Thwaites, Enum. Pl. Zeyl. 342,

1864; C. B. Clarke, Fl. Brit. India 6: 605, 1893; Trimen & Hook. f., Handb. FI.

Ceylon 5: 33. 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 156, f. 4 A-D, 1935;

Kern, Fl. Males. I, 7 (3): 617,1974.

21a. Subsp. compressus.

Jaffna District: Jaffna, Trimen Feb. 1890 (PDA), Silva in 1920 (PDA).

Vavunia District: S of Mullaittivu, Fosberg & Balakrishnan 53527 (NY, US).

Polonnaruwa District: Gal Oya Reservoir, near the spilway, 270 m alt., Comanor

561b (US). Trincomalee District: 7 miles S of Dambulla on Kandy Rd., Clayton

5078 (K, PDA); between Naula and Dambulla, vicinity of Mile 40, Jayasuria 57

(PDA). Kurunegala District: Kurunegala, Thwaites C.P. 812 (K, PDA). Kandy

District: Peradeniya, Trimen C.P. 812 (PDA); Kandy, Alston in 1926 (PDA).

Colombo District: Danowita, Amaratunga 1450 (PDA). Hambantota District:

Ruhuna National Park, Gonalabbe, Block I, Cooray & Balakrishnan 20 Jan. 1969 —

(US). Monaragala District: E of the south end of Jayanthiwewa, T. Koyama et al.

13991 (NY, PDA, US); way to Nilgola, Trimen Jan. 1888 (PDA); Inginiyagala,

Amaratunga 1706 (PDA).

Distribution, Cosmopolitan; tropical, subtropical and temperate regions of all

world; weed of cultivated and waste places, more frequent along sea coast.

Ceylon Cypereae 139

21b. Subsp. micranthus [. Koyama, subsp. nov.

Planta omnibus partibus minor; inflorescentia semper in capitulum unicum

congesta; glumae 2 mm longae; achaenia obovato-orbicularia, basi valde attenuantia,

vix 1 mm longa, 0.75 mm lata; caeteroquin sicut ssp. compressus.

Trincomalee District: exposed eastern bottom of Kantalai Tank at Mile 134

on Habarana-Trincomalee Rd., Davidse 7561 (NY, holotype; MO, isotype).

Distribution. Thus far known only by the collection cited above.

Subspecies micranthus differs from the typical phase primarily by the smaller

glumes (2 mm vs. ca. 3.5 mm in length) and achenes (1 by 0.75 mm ys. 1.5 by

1 mm in size) in addition to its comparatively smaller habit with always congested

inflorescences. Besides the fertile achenes every detail of floral parts shows no

evidence of the possible hybrid nature of this subspecies, though it has been known

that C. compressus rather freely hybridize with several species. So far I have not

come across as yet any specimen of C. compressus bearing such smaller glumes

and achenes from its total range.

22. Cyperus iria L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 344, 1864;

C. B. Clarke, Fl. Brit. India 6: 606, 1893; Trimen & Hook. f., Handb. Fl. Ceylon

5: 18, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 150, 1935; Kern, Fl. Males. I,

7 (3): 616, 1974.

Jaffna District: Jaffna, Silva Nov. 1920 (PDA). Trincomalee District: Trinco-

malee, Ramanathan Jul. 1926 (PDA). Anuradhapura District: Mile 123 on Rd.

A 14, T. Koyama et al. 13920 (NY, PDA, US); Wilpattu National Park, Manike-

pola Uttu, T. Koyama & Jayasuria 13951 (NY, PDA, US). Matale District: between

Mirisgoniyawa and Dambulla, Silva Nov. 1926 (PDA); between Naula and Dam-

bulla, vicinity of Mile 40, Jayasuria 55 (PDA); Matale North, Jayasuria 53 (PDA).

Kandy District: Peradeniya, Trimen C.P. 811 (K, PDA); Pundaluoya, Hughes

Dec. 1902 (PDA). Kurunegala District: Kurunegala, Thwaites C.P. 811 in part

(PDA). Colombo District: Giriulla, Amaratunga 998 (PDA); Veyangoga, Amara-

tunga 1946 (PDA). Kalutara District: Bolgoda, Amaratunga 2524 (PDA). Ruhuna

National Park: Buttawa Plain, near Ecology Project Plot R35, Cooray 12 Dec.

1969 (US); Yala Plain, Patanagala Beach, 3 m alt., Comanor 863B (NY, US).

Distribution. Widely distributed in Asia from Iran and Afghanistan through

Indian Subcontinent northeastwards to China and Japan, and eastwards to Malesia

and Australia; also in East Africa. Apparently introduced to southeastern U.S.A.,

West Indies and South America. Common weed in cultivated ground and rice field.

23. Cyperus diffusus Vahl, Enum. Pl. 2: 321, 1806; C. B. Clarke, Fl. Brit. India

6: 603, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 28, 1900 Kikenthal,

Pflanzenr. 4 (20), 101 Heft: 208, 1936; Kern, Fl. Males. I, 7 (3): 619, 1974.

Cyperus nigro-viridis Thwaites, Enum. Pl. Zeyl. 344, 1864.

23a. Subsp. diffusus.

Kurunegala District: Kurunegala, Thwaites C.P. 2879 (PDA).

Distribution. India, Indo-China, southern China and Malesia.

23b. Subsp. macrostachyus (Boéckleler) T. Koyama, stat. nov.

Cyperus diffusus Vahl var. macrostachyus Béckeler, Linnaea 35: 534, 1868;

Trimen & Hook. f., Handb. Fl. Ceylon 5: 28, 1900; Kiikenthal, Pflanzenr. 4 (20),

101 Heft: 209, 1936; Kern, Fl. Males. I, 7 (3): 619, f. 54, 1974. Cyperus

pubisquama Steudel [in Zollinger, Syst. Verz. 1: 62, 1854, nomen,] Synops. PI.

Glumac. 2: 20, 1855.

140 Gardens’ Bulletin, Singapore — XXX (1977)

Kurunegala District: Kurunegala, Thwaites C.P. 3931 (PDA); Mallawapitiya,

Amaratunga 1101 (PDA); Athagala, Amaratunga 678 (PDA). Matale District:

Matale, Ferguson Oct. 1884 (PDA).

Distribution. From India through Indo-China eastwards to Malesia.

Subspecies macrostachyus differs from the typical phase in its generally larger

habit, far more copious inflorescence and longer spikelets as mentioned in the key

to species. In Ceylon subsp. diffusus is very rare, and has so far been documented

only by Thawaites C.P. 2879.

24. Cyperus platystylis R. Brown, Prodr. Fl. Nov. Holl. 214, 1810; C. B. Clarke,

Fl. Brit. India 6: 598, 1893; Trimen & Hook, f., Handb. Fl. Ceylon 5: 24, 1900;

C. E. C. Fischer in Gamble, Fl. Madras 9: 1639, 1931; Kiikenthal, Pflanzenr.

4 (20), 101 Heft: 185, f. 27, 1936; Kern, Fl. Males. I, 7 (3): 618, 1974.

Fig. 5

Cyperus pallidus Nees [Linnaea 9: 284, 1835, nomen nudum, &] in Wight,

Contrib. Bot, India 79, 1834; Thwaites, Enum. Pl. Zeyl. 34, 1864. Not of

Willdenow ex Link, 1820, nor of Savi, 1830. Anosporum pallidum (Nees)

Bockeler, Linnaea 36: 412, 1870.

Kurunegala District: Kurunegala, Trimen C.P. 3559 in part (PDA); Wariya-

pola, Amaratunga 1104 (PDA). Batticaloa District: Kalmunai, Ahmed Aug. 1940

(PDA); Batticaloa Trimen C.P. 3559 in part (K, PDA). Amparai District: Lahu-

gala Tank, T. Koyama et al. 14030 (NY, PDA, US). Colombo District: Mutu-

rajawela, Amaratunga 144 (PDA).

Distribution. India and Ceylon to Taiwan, and through Malesia eastwards to

northern and eastern Australia. In marshes and ponds usually growing in large

communities,

This rare species is easily discernible in its achenes with spongy thickened

angles and densely spiculose hemisphaerical umbel. The rhizomes are often more

or less elongated and obliquely ascending, but do not emit stolons.

25. Cyperus alternifolius L., Mant. 2: 28, 1771.

This species, known from East Africa and Madagascar, is represented in

Ceylon by the following.

Subsp. flabelliformis (Rottboell) Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 193, 1936.

Cyperus flabelliformis Rottboell, Descr. Pl. Rar. Progr. 22, 1772, & Descr.

Icon. Rar. Nov, Pl..-42, f° 92°." 2) 14.773.

Badulla District: Rawanaella Waterfall, ca. 2 miles SE of Ella, Mile 13,

780 m alt., Davidse et al. 8858 (MO, NY, PDA, US). Monaragala District: 4

miles N of Wellawaya, Wheeler 12689 (PDA, US).

Distribution. A native of tropical Africa and Arabia. Widely cultivated as an

ornamental; in Ceylon escaped and established in grassy places.

26. Cyperus radians Nees & Meyen [ex Nees, Linnaea 9: 285, 1835, nomen

nudum,] ex Kunth, Enum. Pl. 2: 95, 1837, as “‘radicans”; C. B, Clarke, Fl. Brit.

India 6: 605, 1893; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 214, 1936; Kern, FI.

Males. I, 7 (30): 623, 1974.

Trincomalee District: seashore several miles E of Toppur, ca. 15 miles SE of

Trincomalee, 7. Koyama 14060b. (NY).

Distribution. Indo-China, Malay Peninsula, southeastern China, Borneo.

Sporadically noted in sandy sea coast.

New to the flora of Ceylon. The above collection marks a considerable range

extension of this southeastern Asian species into the Indian Subcontinent. This is a

Ceylon Cypereae 141

SSS Es ©, je

44 Se es KA AT

\ 2s es —_ <a

-—

ad i

ive d

~ ~ a

SIRENS

LAN G Q

]

Fig. 5. Cyperus platystylis R. Brown. AA, habit. B, spikelet. C, glume. D, dorsal view

of achene. E, ventral side of achene. Scales = | mm,

142 Gardens’ Bulletin, Singapore — XXX (1977)

peculiar looking species growing in sand. Since the culms are normally extremely

abbreviated and hidden in the leaf tuft, the elongated umbel rays look like tufted

culms.

27. Cyperus pulcherrimus Willdenow ex Kunth, Enum. Pl. 2: 1837; C. B. Clarke,

Fl. Brit. India 6: 600, 1893, & Illustr. Cyper. t. 10 f. 1-3, 1909; Trimen & Hook.

f., Handb. Fl. Ceylon 5: 27, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 242,

1936; Kern, Fl. Males. I, 7 (3): 624, 1974.

“Cyperus silletensis Nees’ sensu Thwaites, Enum. Pl. Zeyl. 343, 1864.

Wilpattu National Park: between Kattankandal Kulam and Occapu Junction,

Wirawan et al. 977 (US); Manikepola Uttu, T. Koyama et al. 13462 (NY, PDA,

US); between Kunbuk Wila and Kokkare Villu, T. Koyama & Jayasuria 13946b

(NY); Malimaduwa Guards Quater, Wirawan & Cooray 1141 (NY, PDA).

Batticaloa District: Batticaloa, Trimen C.P. 3558 (K, PDA). Kurunegala District:

Kurunegala, Rasanayaka in 1927 (PDA). Matale District: 8 miles E, of Habarane,

T. Koyama & Herat 13567 (NY, PDA, US). Colombo District: Kotugoda,

Amaratunga 2027 & 2262 (PDA).

Distribution. India, Indo-China, Malesia.

28. Cyperus haspan L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 343,

1864; C. B. Clarke, Fl. Brit. India 6: 600, 1893; Trimen & Hook. f., Handb. FI.

Ceylon 5: 26, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 247, f. 28 E-G, 1936;

Kern, Fl. Males. I, 7 (3): 624, ff. 56 & 57, 1974.

Anuradhapura District: Anuradhapura, Brodie C.P. 799 (K, PDA); Wilpattu

National Park, Manikepola Uttu, T. Koyama & Jayasuria 13953 (NY, PDA, US).

Trincomalee District: Trincomalee, Glenie C.P. 799 in part (PDA); Periya Kulam,

ca. 7 miles NW of Trincomalee, 7, Koyama et al. 14066 (NY, PDA, US). Puttalam

District: Wilpattu West Sanctuary, Marai Villu, 7. Koyama 13973 (NY, PDA,

US); Madape, Amaratunga 2569 (PDA). Vavunia District: Kokkavil, Clayton

5294 (K, PDA). Nuwara Eliya District: Rangalla to Corbett’s Gap, Ballard 1416

(K, NY). Kurunegala District: Kurunegala, Rasanayake in 1927 (PDA); Ibbaga-

muwa, Amaratunga 1581 (PDA). Kandy District: Peradeniya, Alston 311 (PDA);

Upper Hantana Rd., Comanor 314 (PDA, US); Hantane, Thwaites C.P. 965 in

part (K, PDA). Kegalle District: Alapalawala, along river Watura Oya, ca. 500 m

alt.. T. Koyama & Samarakoon 13558 (NY, PDA, US); Mawanella, Amaratunga

1199 (PDA). Badulla District: Rd. A 4, ca. 3 miles W of Koslanda, ca. 900 m alt.,

T. Kvyama et al, 14032 (NY, PDA, US). Ratnapura District: 11 miles E of

Deniyana at Mile 62 on Rd. A 17, 750 m alt., Davidse 7885 (MO, NY, PDA, US);

Kuruwita, Trimen in 1895 (PDA). Colombo District: Muthurajawela, Amaratunga

135 (PDA). Kalutara District: Nugegoda, Bolgoda Lake Scheme, Amaratunga

2538 (PDA). Galle District: Galle, Thwaites C.P. 965 (PDA): Bentota Ganga,

Amaratunga 2332 (PDA); Ambalangoda, Amaratunga 2642 (PDA). Monaragala

District: ca. 3 miles W. of Wellawaya at Mile 135/10 on Rd. A 4, 990 ft. alt.,

Davidse 7731 (MO, NY).

Distribution. Tropical and subtropical regions of all the world with its range

extending northwards into Japan in East Asia. Abundant in wet places; frequent

in rice fields.

This species is sometimes divided into two subspecies, ssp. haspan and ssp.

junciformis Kiikenthal, the latter differing from the typical phase in having long-

creeping rhizome along which the culms are disposed in a row in a more or less

spaced manner. In plants from Asia these two subspecies seem quite well circum-

scribed, and consequently, appear to be recognizable. Typical ssp. haspan, with

the culms tufted without conspicuous rhizome, is often confused with closely related

C. tenuispica, The only reliable character to separate these two are floral glumes.

In C. tenuispica the floral glumes are spaced exposing a considerable part of the

Ceylon Cypereae 143

subtending achene between two glumes, while in C. haspan the achenes are com-

pletely hidden by the closely disposed glumes.

29. Cyperus tenuispica Steudel, Synops. Pl. Glumac. 2: 11, 1855: Kiikenthal,

Pilanzenr. 4 (20), 101 Heft: 245, f. 28 A-D, 1936; Kern, Fl. Males. I, 7 (3): 625,

f. 58, 1974.

“Cyperus flavidus Retzius” sensu C. B. Clarke, Fl. Brit. India 6: 600, 1893;

Trimen & Hook. f., Handb. Fl. Ceylon 5: 27, 1900.

Jaffna District: Mullaitiva, Ferguson s.n. (PDA); south shore of Jaffna Lagoon,

ca. 8 miles SE of Poonaryn, T. Koyama et al. 14046 (NY, PDA, US). Wilpattu

National Park; Kuda Pathessa, 7. Koyama 13387 (NY). Matale District: Matale

north, Jayasuria 54 (PDA). Kandy District: Paradeniya, Alston in 1926 (PDA),

Trimen C.P. 805 (PDA); Gannoruwa, Alston 315 (PDA). Colombo District:

Gampaha, Simpson 8602 (PDA); Makawita, Amaratunga 1779 (PDA). Hamban-

tota District: Ruhuna National Park, Block I, Rakinawala, Cooray 7 Dec. 1969

(US).

Distribution. Tropical Africa, India, Nepal, Indo-China, Malesia, south-

eastern China, Japan. Wet places; often as a weed in rice field.

30. Cyperus difformis L., Cent. Pl. 2: 6, 1756; C. B. Clarke, Fl. Brit. India 6: 599,

1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 25, 1900; Kiikenthal Pflanzenr.

4 (20), 101 Heft: 237, f. 27 F-H, 1936; Kern, Fl. Males. I, 7 (3): 629, 1974.

Trincomalee District: Trincomalee, Ramanathan in 1926 (PDA). Polonaruwa

District: ca. 1 mile E of Kolakanaweli, at Mile 56/7 on Rd. A 11, 7. Koyama

13577 (NY). Batticaloa District: Batticaloa, Thwaites C.P. 3042 (K, PDA). Kandy

District: Gannoruwa, Alston 316 (PDA). Matale District: Ereula Tank, ca. 5

miles SE of Dambulla, 650 ft. alt., Davidse 7390 (MO, NY). Colombo District:

Danowita, Amaratunga 192 (PDA). Amparai District: Tandiadi Kalapu Lagoon,

S of Mile 215 on Rd. A 4, T. Koyama et al. 14015 (NY, PDA, US). Ruhuna

National Park: Komawa Wewa, Cooray 10 Dec. 1969 (NY), US); Uraniya,

Cooray 22 Mar. 1970 (NY).

Distribution, Widely distributed in Eurasia from southern Europe through

India and China to Japan and Malesia, also in Pacific Islands and Australia;

introduced to Central America and South Africa possibly with rice.

31. Cyperus castaneus Willdenow, Sp. Pl. 1: 278, 1797; Thwaites, Enum. Pl. Zeyl.

343, 1864; C. B. Clarke, FI. Brit. India 6: 598, 1893; Trimen & Hooks. f., Handb.

FI. Ceylon 5: 25, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 264, 1936; Kern, FI.

Males. I, 7 (3): 630, 1974.

Anuradhapura District: Wilpattu National Park, East Intermediate Zone,

along Wildlife Dept. Path, ca. 2 km from its barrier gate. T. Koyama & Jayasuria

13966 (NY, PDA, US). Polonnaruwa District: SE of Giritale Wewa near Circuit

Bangalow, Fosberg et al. 51955 (PDA, US); Polonnaruwa, Clayton 5115 (K,

PDA). Matale District: Dambulla Rock, Trimen in 1896 (PDA). Kegalle District:

Simpson 8358 (PDA). Badulla District: between Ratupahana and Haldunmulla,

Ormiston in 1909 (PDA). Ruhuna National Park: Block I, Cooray 7 Dec, 1969

(US); between Buttawa and Karangaswela, Cooray 3 Dec. 1969 (NY, US).

Distribution. From India through Indo-China to Malesia and northern

Australia.

32. Cyperus cuspidatus Kunth in Humb., Bonpl. & Kunth, Nov. Gen. et Sp.

Pl. 1: 204. 1815: C. B. Clarke, FI. Brit. India 6: 598, 1893; Trimen & Hook. f.,

Handb. Fl. Ceylon 5: 26, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 261,

f. 29 E, 1936; Kern, Fl. Males. I, 7 (3): 629, f. 59, 1974.

Cyperus angustifolius Nees in Wight, Contrib. Bot. India 79, 1834.

144 Gardens’ Bulletin, Singapore — XXX (1977)

Anuradhapura District: Wilpattu National Park NE of Kuruttu Pondi Villu,

T. Koyama 13413 (NY). Polonnaruwa District: 7 miles NW of Polonnaruwa, SE

of Giritale Wewa, Fosberg & Ripley 51955 (US). Nuwara Eliya District: ca. 15

miles NW of Nuwara Eliya, at Mile 32/8 on Road A 5, 1020 m alt., Davidse

et al. 7946 (MO, NY, PDA). Monaragala District: Inginiyagala National Park,

between Baduluwila and ‘‘Westminster Abbey,” 7. Koyama et al. 13997 (NY,

PDA, US); ca. 25 miles W of Pottuvil on road to Wellawaya, near Mile 76,

Davidse et al. 8934 (MO, NY, PDA). Hambantota District: Ruhuna National

Park, Karasugaswela, Cooray 12 Dec. 1969 (NY, PDA).

Distribution. Pantropic, with its range extending north to southern China and

Formosa.

33. Cyperus arenarius Retzius, Obs. Bot. 4: 9, 1786; Nees in Wight, Contrib. Bot.

India 77, 1834; Thwaites, Enum. Pl. Zeyl. 342, 1864; C. B. Clarke, FI. Brit. India

6: 602, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 23, 1900; C. E. C. Fischer

in Gamble, Fl. Madras 9: 1640, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 270,

1936. Fig. 6

Bobartia indica L., Fl. Zeyl. 17, 1747. — Cyperus conglomeratus Rottb.

var. arenarius (Retzius) Cosson, Explor. Algir. Bot. 2: 245, 1855.

Mannar District: Talaimannar, Silva in 1916 (PDA). Trincomalee District:

Trincomalee, Glenie C.P. 798 (K, PDA); Puttalam District: Kalpitiya, Trimen in

1883 (PDA); Chilaw, Trimen in 1880 (PDA); Wilpattu National Park, West

Intermediate Zone, Pallugaturai Beach, Wheeler 12104 (PDA). Batticaloa District:

Passikudah, Mueller-Dombois in 1968 (PDA); Batticaloa, Gardener C.P. 798

(PDA). Colombo District: Colombo, Ferguson C.P. 794 (PDA); Uswetakeiyawa,

Amaratunga 126 (PDA). Amparai District: Pottuvil, Rest House beach, Bala-

krishnan 389 (PDA); Arugam Bay, T. Koyama et al. 14029 (NY, PDA, US),

Fosberg & Sachet 53043 (NY, US). Hambantota District: Bentota, beach behind

rest house, Ballard 1510 (K, PDA); Ruhuna National Park, beach E of Buttawa

Modera, 2-3 m alt., Fosberg 50315 (US); Patanagala Beach, Cooray 17 Nov. 1969

(PDA, US).

Distribution. Southern Iran, Pakistan, India, Ceylon, Cochinchina.

A coastal species easily reconginable by the whitish head with broad spikelets

and extensive rhizome system, with which it usually forms a large pure community.

34. Cyperus conglomeratus Rottboell, Descr. Icon. Rar. Nov. Pl. 21, t J5 f.7,

1773; Thwaites, Enum. Pl. Zeyl. 343, 1864; C. B. Clarke, Fl. Brit. India 6: 602,

1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 23, 1900; Kiikenthal, Pflanzenr.

4 (20), 101 Hen: 27 2..3c Fos leo.

Cyperus pachyrrhizus Nees ex Bockeler, Linnaea 35: 545, 1868; C. B. Clarke,

Fl. Brit. India 6: 603, 1893; C. E. C. Fischer in Gamble, Fl. Madras 9: 1640,

193 t: Cyperus conglomeratus Rottb. var. pachy[r]rhizus (Nees ex

Bockeler) Trimen in Trimen & Hook. f, Handb. Fl. Ceylon 5: 23, 1900.

Puttalam District, Wilpattu National Park: Kollankanatta Beach, Cooray 28

Sept. 1969 (US); Pallugaturai, 7. Koyama & Jayasuria 13960 (NY, PDA, US),

Fosberg et al. 50917 (NY, US), Wheeler 12105 (PDA).

Distribution. Mediterranean Region, North Africa, East Africa, Iran, Arabia,

Madagascar, southern India and Ceylon.

This is one of the western Asian element reaching Ceylon, Having compared a

good series of Indian and Ceylonese specimens with those from western Asia I am

unable to separate Indian plants from western Asian ones.

35. Cyperus cephalotes Vahl, Enum. PI. 2: 311, 1806; C. B. Clarke, FI. Brit. India

6: 597, 1893, & Illustr. Cyper. t. 6 f. I-7, 1909; Trimen & Hook. f., Handb, FI.

Ceylon 5: 17, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 292, 1936; Kern, FI.

Males. I, 7(3): 633, ff. 61-62, 1974.

ee -

145

Ceylon Cypereae

Fig. 6. Cyperus arenarius Retzius. A, habit. B, spikelet. C, prophyll. D, glume. E, pistil

F, achene. Scales = 1 mm.

146 Gardens’ Bulletin, Singapore — XXX (1977)

Cyperus hookerianus Thwaites, Enum. Pl. Zeyl. 342, 1864. Anos-

porum cephalotes (Vahl) Kurz, Journ. Asiat. Soc. Bengal 45 (2): 159, 1876.

Kurunegala District: Ibbagamuwa, Amaratunga in 1967 (PDA).

Distribution. From India through Indo-China northeastwards to southern

China, and eastwards to Malesia and northern Australia.

Trimen (op. cit.) reported this species from Galle, Lunugala and Uva.

Currently in the Peradeniya Herbarium the specimen cited above is the only sheet.

36. Cyperus alopecuroides Rottboell, Descr. Pl. Rar. Progr. 20, 1772, & Descr.

[con. Rar. Nov. Pl. 38, ¢. 8 f. 2, 1773; Thwaites, Enum. Pl. Zeyl. 342, 1864;

Trimen & Hook. f., Handb. Fl. Ceylon 5: 38, 1900; Kiikenthal, Pflanzenr. 4 (20),

101 Heft: 71, 1935; Kern, Fl. Males. I, 7 (3): 603, 1974

Juncellus alopecuroides (Rottb.) C. B. Clarke, Fl. Brit. India 6: 595, 1893.

Anuradhapura District: between Ratmale and Talawa, ca. 7 miles SSW of

Anuradhapura, T. Koyama & Herat 13608 (NY, PDA, US). Polonnaruwa District:

Minnariya Tank, Trimen in 1884 (PDA). Batticaloa District: Batticaloa, Trimen

C.P. 3560 (K, PDA). Hambantota District: Tissamaharama, Trimen in 1882

(PDA).

Distribution. North and Tropical Africa, Madagascar, India, Indo-China,

Malesia and northern Australia.

37. Cyperus pygmaeus Rottboell, Descr. Icon. Rar. Nov. Pl. 20, ¢. 14 f. 4-5

1773; Nees in Wight, Contrib. Bot. India 72, 1834; Trimen & Hook. f., Handb.

Fl. Ceylon 5: 18, 1900; Kern, Fl. Males. I, 7 (3): 634, 1974.

Juncellus pygmaeus (Rottb.) C. B. Clarke, Fl. Brit. India 6: 596, 1983.

Cyperus michelianus (L.) Delile subsp. pygmaeus (Rottb.) Aschers, &

Graebn., Synops. Mitteleur. Fl. 2 (2): 273, 1903; Kiikenthal, Pflanzenr. 4 (20), 101

Heft: 312, f. 35 F-G, 1936.

Polonnaruwa District: Gal Oya Reservoir, near spilway, 270 m alt., Comanor,

56la & 561b (NY, US). Puttalam District: Palavi Atta Villu, Cooray 6 Oct. 1969

(NY); Puttalam, Trimen Aug. 1883 (PDA). Trincomalee District: Kantalai Tank,

Mile 134 on Rd. A 6, Davidse 7518 (MO, NY). Mannar District: Aruvi Aru, Mile

125 on Rd. A 4, T. Koyama et al. 13934B (NY); Palavi, Cooray in 1969 (PDA).

Anuradhapura District: Anuradhapura, Trimen Oct, 1883 (PDA). Matale District:

Ereua Tank, ca. 5 miles ESE of Dambulla, 650 ft. alt., Davidse 7391 (MO, NY,

PDA, US). Colombo District: Colombo, Ferguson C.P. 3947 (K, PDA). Hamban-

tota District: Ruhuna National Park, Attaville, Cooray 6 Oct. 1969 (NY).

Distribution. Widely spread in Eurasia, from Mediterranean Region and East

Africa through Asia Minor and India to southeastern Asia, Malesia and Australia.

PYCREUS P. Beauvois

1. Achenes smoothish to puncticulate with isodiametrical or hexagonal epidermal

cells

2. Culms few- to several-nodose below the middle, the lower part decumbent

or obliquely ascending, branching and rooting at lower nodes; glumes

furrowed on both sides of costa ..:...0..s..scsesevesceevee 39 P. sanguinolentus

2. Culms not nodose above the base, erect from very base; glumes not

furrowed

Ceylon Cypereae 147

3. Culms 20-90 cm tall; glumes acute to subobtuse at apex, never

emarginate nor cuspidate

4. Glumes orbicular-obovate, ca. 1 mm wide in half view, the margins

very broadly whitish-hyaline, rounded to apex, spikelets 3 mm wide;

culms 2—3 mm thick, mostly solitary ............ 40 P. puncticulatus

4. Glumes ovate to lance-ovate, ca. 0.5 mm wide in half view, hardly

or very narrowly hyaline on margins, acute at apex; spikelets ca.

1.5 mm wide; culms 0.7-2 mm thick, as a rule tufted

5. Lowest bract as long as to at most 2.5 times as long as the

umbel, when longer than that, then inflorescence congested in

a head; spikelets rusty brown to yellowish brown ..................

- eee RUM Sidtnn -adteon pbinwng- Eh cate redaea, 41 P. polystachyos

5. Lowest bract 3 to 5 times as long as the umbel; spikelets dark

purplish-brown to chestnut-brown .................. 42 P. flavidus

3. Culms 1-8 cm tall; glumes truncate to emarginate at apex, the midvein

excurrent beyond the glume apex into a recurved mucro

6. Glumes ovate, close together not exposing achenes; rays normally

Porry tre veloped y. 660. Sete LS 43a P. pumilus ssp. pumilus

6. Glumes oblong to lance-oblong, spaced, so that achenes exposed

between glumes; rays well developed ......................cccccecee eee

Mes See sone snvoXe eas enc <5 dtoh es 43b P. pumilus ssp. membranaceus

38. Pycreus stramineus (Nees) C. B. Clarke, Fl. Brit. Ind. 6: 589, 1893; Alston

in Trimen & Hook. f., Handb. Fl. Ceylon 6: 306, 1931.

Cyperus stramineus Nees in Wight, Contrib. Bot. India 74, 1834; Trimen,

Handb. FI. Ceylon 5: 19, 1900. Not of Desf. ex Link, 1820. Cyperus sub-

stramineus Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 398, 1936; Kern, Fl. Males.

Ser. 1, 7 (3): 653, 1974.

Kurunegala District: Kurunegala, Trimen C.P. 3776 (PDA).

Distribution. India, Ceylon, Indo-China and Malay Peninsula.

Thus far known from Ceylon by a single collection only. This species closely

resembles P. flavescens of much wider distribution, from which it differs primarily

in its longer spikelets that are 8 to 35 mm long bearing many acute-tipped glumes

in contrast to shorter spikelets (6 to 10 mm in length) with several obtuse

glumes in the latter, .

39. Pycreus sanguinolentus (Vahl) Nees [Linnaea 9: 283, 1835. Invalid combina-

tion] ex C. B. Clarke, FI. Brit. India 6: 590, 1893; C. E. C. Fischer in Gamble, FI.

Presid. Madras 9: 1627, 1931; Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6:

307, 1931.

Cyperus sanguinolentus Vahl, Enum. Pl. 2: 351, 1806; Nees in Wight,

Contrib. Bot. India 75, 1834; Thwaites, Enum. Pl. Zeyl. 342, 1864; Trimen &

Hook. f., Handb. Fl. Ceylon 5: 20, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft:

385, 1936; Kern, Fl. Males. Ser. 1, 7 (3): 646, 1974. Cyperus eragrostis

Vahl, Enum. Pl. 2: 322, 1806; Trimen, Syst. Cat. Flow. Pl. & Ferns Ceylon 99,

1885. Not of Lamarck, 1791.

Wilpattu National Park: Manikepola Uttu, 7. Koyama et al. 13460 (NY,

PDA, US). Kandy District: Peradeniya, near water fall above University campus,

Comanor 690 (NY): ca. 5 miles S.E. of Gampola, at Mile 18/6 on Gampola-

Nuwara Eliya Rd., 720 m alt., Davidse et al. 7924 (MO, NY). Ratnapura District:

11 miles E. of Diniyaya on Rd. A17, 750 m alt. Davidse 7586 (MO, NY).

Amparai District: Senanaike Samudra, Padagoda, T. Koyama et al. 13983 (NY.

PDA, US).

Gardens’ Bulletin, Singapore — XXX (1977)

148

ne.

G, lateral view of ache

, habit. B & C, two views of prophyll.

F, glume.

ikelet. E, portion of rhachilla with two flowers.

spi

Scales = 1 mm.

Fig. 7. Pycreus puncticulatus (Vahl) Nees. A

Ceylon Cypereae 149

Rather widely distributed in the Old World from northeastern Africa through

Afghanistan, India and Indo-China northeastwards to China and Japan, eastwards

to Malesia.

40. Pycreus puncticulatus (Vahl) Nees, Fl. Brasil. 2 (1): 10, in note, 1842: C. B.

Clarke, Fl. Brit. India 6: 593, 1893; C. E. C. Fischer in Gamble, Fl. Presid. Madras

9: 1628, 1931; Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1931.

Fig. 7.

Cyperus puncticulatus Vahl, Enum. Pl. 2: 348, 1806; Thwaites, Enum. PI.

Zeyl. 342, 1864; Trimen & Hook. f., Handb. Fl. Ceylon 5: 21, 1900; Kiikenthal,

Pflanzenr. 4 (20), 101 Heft: 362, 1936. Cyperus baccha Kunth, Enum.

ee 2 BSS S357. Pycreus baccha (Kunth) Nees, [Linnaea 9: 283, 1835.

Invalid combination] Fl. Brasil. 2 (1): 10, 1842. In note.

Trincomalee District: Mavadichchenai, Alston 544 (PDA); Trincomalee,

Glenie C.P. 3751 in part (PDA). Matale District: Dambulla, Thwaites C.P. 3751

(K, PDA). Kurunegala District: Kurunegala, Ranasinghe 843 (PDA). Galle

District: Haburagala, Amaratunga 2315 (PDA). Amparai District: E of Kunukala

Kalapuwa Lagoon, ca. 4 miles S of Panama, sea level, T. Koyama et al. 14027

(NY, PDA, US); Helawe Eliya, ca. 7 miles S of Panama, E of Helawe Lagoon,

sea level, T. Koyama et al. 14025 (NY, PDA, US). Ruhuna National Park:

Uraniyawala, ca. 1 mile W of Buttawa, Fosberg et al. 51018 (US); Palugaswela,

Ecology Project Block I, Cooray 23 Mar. 1970 (NY).

Distribution. Rather sporadically noted from southern India (Bombay, Mysore

and Madras), Malay Peninsula and Cochinchina.

In the genus Pycreus this species is well demarcated by its relatively loose,

large umbel with broad, red-brownish spikelets, in which it resembles superficially

Cyperus procerus, a stoloniferous perennial with trigonous achenes.

41. Pycreus polystachyos (Rottboell) P. Beauvois, Fl. d;Oware 2: 48. ¢. 8&6, f. 2,

1807; C. B. Clarke in Hook. f., Fl. Brit. India 6: 592, 1893: Alston in Trimen

& Hook. f., Handb. Fl. Ceylon 6: 307, 1931.

Cyperus polystachyos Rottboell, Descr. Icon. Rar. Nov. Pl. [llustr. 39,

t. 11, f. 1, 1773; Nees in Wight, Contrib. Bot. India 75, 1834; Thwaites, Enum.

Pl. Zeyl. 342, 1864; Trimen & Hook. f., Handb. Fl. Ceylon 5: 20, 1900; Kuken-

thal, Pflanzenr. 4 (20), 101 Heft: 367, 1936; Kern, Fl. Males. Ser. 1, 7 (3): 649,

1974. Cyperus paniculatus Rottboell, Descr. Icon. Rar. Nov. Pl. Illustr.

40, 1773. Pycreus paniculatus (Rottb.) Nees [Linnaea 9: 283, 1835. Invalid

combination] ex Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1931.

Cyperus polystachyos Rottb. var. Jaxiflorus Bentham, Fl. Austral. 7: 261,

1878; Kiikenthal, Pflanzenr. 4 (20), Heft. 101: 370, 1936. Pycreus poly-

stachyos P. Beauv. var. laxiflorus (Bentham) C. B. Clarke, Fl. Brit. India 6: 592,

1893.

Jaffna District: Jaffna, Ferguson C.P. 800 (PDA). Anuradhapura District:

Anuradhapura, Alston 638 (PDA). Wilpattu National Park: Kuda Patessa,

Wirawan et al. 1035 (US); Kollankanatta, Mueller-Dombois et al. 27 Apr. 1969

(US); NE. of Kuruttu Pondi Villu, T. Koyama 7560 (NY). Trincomalee District:

Trincomalee, Glenie C.P. 800 (PDA); Ottawady, valley of downstream of Mahaweli

Ganga, ca. sea level, T. Koyama 13652 (NY, PDA, US); Mile 97 on Rd. A 15,

Davidse 7560 (MO, NY). Polonnaruwa District: ca. 1 mile NE of Elahera along

Amban Ganga, 400 ft. alt., Davidse 7357 (MO, NY). Batticaloa District: Batticaloa,

Thwaites C.P. 800, March 1868 (K, PDA). Nuwara Eliya District: Ramboda Rd.,

near Ramboda, 3200 ft., Ballard 1136 (K, NY); Maskeliya, Wright 11 Nov. 1926

(PDA). Matale District: Bata-Anduwa Gap, Sevastine, at marker 34/6, 1300 m

alt., Tirvengadum et al. 17 (PDA). Kandy District: Hantane, Silva 296 (PDA);

Peradeniya, Alston 280 (PDA): Gampola-Dolosbage Rd., Grupe 159 (PDA).

150 Gardens’ Bulletin, Singapore — XXX (1977)

Kurunegala District: Kurunegala, Thwaites C.P. 800 (PDA); Beddegama, Alston

1397 (PDA). Colombo District: Muturajawela, Amaratunga, 18 Jan, 1971 (PDA).

Galle District: Horawala, left bank of Bentota Ganga, Amaratunga 2495 (PDA),

Haburagola, Amaratunga 2381 (PDA). Amparai District: Tandiadi Kalapu

Lagoon, S of mile 215 on Road A4, T. Koyama et al. 14018 (NY, PDA, US);

3 miles NW of Maha Oya, 50 m alt., Davidse et al. 9008 (MO, NY.); Inginiyagala

National Park, between Baduluwela and ‘‘Westminster Abeey”’, 7. Koyama et al.

13995 (PDA). Ruhuna National Park: Uraniya, Ecology Project Block I, Cooray

22 Mar. 1970 (PDA, US).

Distribution. Cosmopolitan, tropical and subtropical regions, The range is

extending into warm-temperate regions in eastern Asia as far as to central Japan,

and into Mediterranean Region.

This species is extremely variable in the color of spikelets and in the

inflorescences, which vary from a rather open umbel with spicately disposed spike-

lets to a head through an intermediate state bearing fascicled spikelets on short

rays. In Ceylon plants with open inflorescences are common. Although Alston (in

Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1932) listed P. ferrugineus as

occurring in Ceylon without citation of specimens, I have so far been unable to

see any documentation of this essentially African species from Ceylon and India.

Pycreus ferrugineus distinctly differs from P. polystachyus at least in its spreading

spikelets that are much broader than those of the latter at 2 to 3 vs. 1.25 to

1.5 mm in width, and it can no way be confused with the latter. Hence, Alston’s

note under P. ferrugineus mentioning that “This is doubtfully distinct from P.

polystachyus,” and ‘‘Up to 5000 ft.; common” implies that his P. ferrugineus may

constitute a misidentification for P. polystachyos itself.

42. Pycreus flavidus (Retzius) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976.

Cyperus flavidus Retzius, Obs. Bot. 5: 13, 1788; Vahl, Enum. PI. 2: 334,

1806; Kern, Fl. Males. Ser. 1, 7 (3): 648, 1974. [Cyperus globosus Allioni,

Auctuar. Fl. Pedemont. 49, 1789; Trimen & Hook. f., Handb. Fl. Ceylon 5: 21,

1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 352, 1936. Not of Forskael, 1775.]

Cyperus strictus Roxburgh, Fl. Ind. ed. 1, 1: 203, 1820. Cyperus

capillaris Konig ex Roxburgh, l.c. 1: 198, 1820; Nees in Wight, Contrib. Bot.

Ind. 76, 1834. Pycreus globosus (Allioni) Reichenbach, Fl. Germ. Excurs.

140, 1830. Pycreus capillaris (Konig ex Roxb.) Nees [Linnaea 9: 283,

1834. Invalid combination] ex C. B. Clarke, FI. Brit. India 6: 591, 1893.

Pycreus strictus (Roxb.) Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307,

1931, “Cyperus flavescens L.” sensu Thwaites, Enum. Pl. Zeyl. 342, 1864.

Polonnaruwa District: ca. 1 mile N.E. of Amban Ganga, 400 ft. alt. Davidse

7357A (MO). Kandy District: ca. 5 miles SE of Gampola at mile 18/6 on Road

A5, 720 m alt., Davidse et al. 7925 (MO, NY); ca. 2 miles E, of Maddakele, slope

N.W. of Krinckles, 1440 m alt., Davidse 8332 (MO); Pallekale, Alston 278 (PDA);

Gordindihela, Willis, 27 Feb. 1906 (PDA); Upper Hantane Rd., above the University

Campus, Comanor 317 (PDA, US). Kurunegala District: Kurunegala, Gardner

C.P. 801 (PDA). Nuwara Eliya District: Nuwara Eliya, Trimen C.P. 801 (PDA),

Amaratunga 230 & 1305 (PDA); Pattipola, Lazarides 7291 (PDA, US); between

Hakgala and Nuwara Eliya, 5400 ft., Ballard 1269 (K, NY); Corbett’s Gap, 4000

ft., Ballard 1050 (K, NY); Moon Plains, 6000 ft., Ballard 1217A (K, NY). Ratna-

pura District: 11 miles E of Deniyaya at mile 62 on Rd. A17, 750 m alt., Davidse.

7884 (MO, NY); Ratnapura, Thwaites C.P. 801 (K, PDA). Colombo District:

Mount Lavinia, Trimen Oct. 1881 (PDA). Monaragala District: ca. 3 miles W. of

Wellawaya at mile 135/10 on Rd. A4, 990 ft., Davidse 7742 (MO, NY). Ruhuna

National Park: Kumbukkan Oya, Ecology Project Block II, Cooray 31 July 1969

(NY, US). Hambantota District: 2 miles E of Katuwana, Lazarides 7350

(PDA, US). ,

Ceylon Cypereae 151

Widely distributed from Mediterranean Region and tropical Africa eastwards

to Central and southern Asia, and northeastwards to Japan.

This widespread species is variable particularly in the width and the color

of spikelets. Ceylonese plants from up countries at altitudes of over 750 m generally

possess strongly compressed, dark purplish-brown spikelets that are mostly less

than 1.5 mm in width, a form resembling plants from Japan and temperate India.

A few specimens collected at low altitudes such as Cooray 31 July 1969 and

Davidse 7357 4A cited above bear less compressed, broader spikelets with stramineous-

brown scales.

43. Pycreus pumilus (L.) Nees [Linnaea 9: 283, 1935. Invalid combination] ex

C. B. Clarke, Fl. Brit. India 6: 591, 1893, concerning the nomenclature, but

excluding basionym; Domin, Bibl. Bot. Heft. 85: 417, 1915; C. E. C. Fischer in

Gable, Fl. Madras 9: 1627, 1931.

Cyperus pumilus L., Cent. Pl. 2: 6, 1756; Trimen & Hook. f., Handb. FI.

Ceylon 5: 19, 1900; Kiikenthal, Pflanzenr. 4 (20), Heft. 101: 375, f. 44 A-E, 1936;

Kern, Fl. Males. Ser. 1, 7 (3): 650, f. 66, 1974. Cyperus pluvinatus Nees

& Meyen in Wight, Contrib. Bot. India 74, 1834; Thwaites, Enum. Pl. Zeyl. 342,

1864. Pycreus nitens Nees, Nova Acta Nat. Cur. 19, Suppl. 1: 53, 1843;

C. B. Clarke, Fl. Brit. India 6: 591, 1893. Pycreus pulvinatus (Nees &

Meyen) Nees & Meyen ex Nees [Linnaea 9: 283, 1835, invalid combination, &]

Nova Acta Nat. Cur. 19, Suppl. 1: 53, 1843. Dichostylis nitens (Nees)

Palla, Bot. Jahrb. 10: 296, 1889. ““Pycreus patens (Vahl)” sensu Alston

in Trimen & Hook. f., Handb. Fl. Ceylon 6: 306, 1931. Invalid combination.

43a. Subsp. pumilus.

Jaffna District: Jaffna Lagoon, north shore at ca. 10 miles SE of Navatkuli,

T. Koyama et al. 14040 (NY, PDA, US); Jaffna, Gardner in 1846 (PDA).

Anuradhapura District: Maradaukadewela, Trimen 21 Jan. 1896 (PDA); Wilpattu

National Park, East Intermediate Zone, along Wildlife Dept. path at ca. 2 miles

from barrier gate, T. Koyama & Jayasuria 13966 (NY, PDA, US). Polonnaruwa

District: Tamankaduwa, Townsend 73/250 (K, US). Trincomalee District: Periya

Kulam, ca. 7 miles NW of Trincomalee, 7. Koyama et al. 14064 (NY, PDA, US):

Trincomalee, west side of Welcomble Hotel. Wheeler 1241] (PDA). Mannar

District: Aruvi Aru, mile 125 on Rd. Al4, T. Koyama et al. 13931 (NY, PDA,

US). Puttalam District: Chilaw, Trimen, Dec. 1880 (PDA). Kandy District: Pera-

deniya, Thwaites C.P. 806 in part (PDA); Gannoruwa, Alston 298 (PDA); Hara-

gama, Alston 3 Oct. 1926 (PDA).

Distribution. From India through Indo-China and southern China eastwards

to Malesia and Australia (rare in Queensland).

Alston (l.c., 1931) applied Cyperus patens Vahl to the Ceylonese plants then

passing as Cyperus pumilus, and proposed a combination, Pycreus patens (Vahl).

Typical C. patens from Africa closely resembles subsp. membranaceus, and it may

not be sufficiently different from the latter. However, C. patens without doubt does

not represent the common phase of Ceylonese C. pumilus.

43b. Subsp. membranaceus (Vahl) T. Koyama, stat. nov.

Cyperus membranaceus Vahl, Enum. PI. 2: 330, 1806. Cyperus nitens

Retz. var. membranaceus (Vahl) Bockeler, Linnaea 35: 484, 1868. Cyperus

pumilus L. var. membranaceus (Vahl) Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 376,

1935. Cyperus pumilus L. forma membranaceus (Vahl) C. B. Clarke, Journ.

Linn. Soc. 21: 44, 1884.

Hambantota District: Ruhuna National Park, between the Entrance and

Ecology Project Plot R9, Cooray s.n., 11 Dec. 1969 (NY, US); Uraniya, Cooray,

22 Mar. 1970 in part (NY).

152 Gardens’ Bulletin, Singapore — XXX (1977)

Distribution. India, Ceylon and Thailand; rather frequent in southern India,

but rare in Thailand.

Cooray’s record, cited above, makes a new entry of subsp. membranaceus to

the flora of Ceylon. The difference between subsp. membranaceus and the typical

phase exists mainly in the relatively minor details of floral glumes and achenes, In

subsp. membranaceus the oblanceolate to oblong-obovate glumes are only ca. 1/3 mm

wide in half view, being broadest much above the middle, and are considerably

loosely disposed on the rhachilla exposing the greater portion of achenes between

the glumes just as in Cyperus tenuispica, whereas in subsp. pumilus its broader

elliptic to ovate-elliptic glumes are 1/2 to 3/5 mm wide in half view being broadest

just below the middle, and are closely arranged, there being no space between two

glumes, In general, the achenes of subsp. membranaceus are truly obovate and

cannot be seen through the membranous glumes, while in subsp. pumilus its

elliptic to lance-elliptic achenes are usually seen through the hyaline glumes. The

spikelets of subsp. membranaceus are shorter and more laxly disposed on well

elongated rays in contrast to the usually more congested nearly head-like inflores-

cence in subsp. pumilus. Though minute, these differances are sufficiently consistent

as far as I have examined materials, and hence I regard subsp. membranaceus a

valid taxon.

MARISCUS Gaertner

1. Spikelets more or less flattened with acute edges, bearing 3 to many achenes;

glumes folded with a conspicuous keel, laxly holding achene

2. Plants perennial; culms more than 45O cm tall; leaves septate-nodulose;

glumes without recurved awn

3. Spikelets lanceolate, 2 mm wide, straw-coloured, disposed in a spike

scaseh desde de cild abs dome RhRe ENE oy ke ey Uieed DRDEIS Mecreee ase ae ean 44 M. javanicus

3. Spikelets linear, 1 mm wide, reddish-brown, densely congested in a

globose head .£.0 27S eet te Se ee 45 M. compactus

2. Plants annual; culms 4-20 cm tall; leaves not septate-nodulose; glumes with

recurved awn, heance spikelets squarrose .................. 46 M. squarrosus

1. Spikelets terete without conspicuous edges, bearing 1 or 2 achene(s) only;

glumes involute without distinct keel, tightly surrounding achene

4. Rhachilla slender, not at all spongy-thickened; spikelets 1- or 2-fruited

5. Base of culms hardly enlarged or slightly globose with a corm-like

enlargement; basal leaf sheaths purplish-brown

6. Spikelets mostly 2-fruited, greenish to greenish-straw-coloured,

patent to ascending at maturity, mostly 4-5 mm long; achenes

2 mm long

7. Spikes umbellate with elongated rays; spikelets spreading or

the lower ones weakly reflexed at maturity ........................

7. Spikes subsessile, nearly capitate; spikelets at least the lower

ones obliquely patent at maturity

8. Rhizome short, not stoloniferous; leaves 2-5 mm wide; -

umbel truly terminali(.9.... .VoOU yO a ee

£6 SUR TT i ee 48a M. cyperinus ssp. cyperinus

8. Rhizome emitting slender stolons; leaves as a rule

0.7-1 mm wide; umbel quasi-lateral with the lowest bract

continued down to the culmx..)..°2i0001. cai

Ceylon Cypereae 153

6. Spikelets 1-fruited, whitish-green, spreading at maturity, mostly

2-3 mm long; achenes ca. 1.3 mm long ............ 49 M. paniceus

5. Base of culm quasi-bulbose, i.e, with an ovoid to ovoid-oblong

thickening clothed with leaf sheaths; basal leaf sheaths straw-brown.

9. Rhizome emitting slender stolons; spikes sessile yet recognizable.

a as a cp ncn boacne «a mpen = deci use 50 M. clarkei

9. Rhizome loosely tufted without stolons; spikes congested in an

ovoid or conical head, not readily recognizable ........................

I Ps tka ones ane psec Pend Ras antipins spies S66 = «areal 51 M. dubius

4. Rhachilla spongy-thickened; achenes sunken in a depression of spongy

rhachilla-internodes; spikelets 1-fruited ............... 52 M. pedunculatus

44. Mariscus javanicus (Houttuyn) Merrill & Metcalfe, Lingnan Sci. Journ. 21:

4, 1945.

Cyperus javanicus Houttuyn, Nat. Hist. I], 13 Aanw. Pl. (1). ¢. 8&8 f. J,

1782; Kern, Fl. Males. I, 7 (3): 635, f. 63, 1974. Cyperus pennatus Lamarck,

Illustr. 1: 144, 1791; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 476, f. 53 A-G, 1936.

— Mariscus albescens Gaudichaud in Freyc., Boy. Bot. 415, 1826; C. B.

Clarke, Fl. Brit. India 6: 623, 1893; Trimen & Hook. f., Handb. Fl. Ceylon §: 40,

1900. ——- Mariscus pennatus (Lamarck) Domin, Bibl. Bot. Heft. 85: 440, 1915.

Wilpattu National Park: Manikepola Uttu, 7. Koyama 13456 (NY, PDA, US).

Anuradhapura District: Ritigala Strict Natural Reserve, approach to Weweltenna

along southern slope, 700 ft. alt., Jayasuria 1294 (PDA); Anuradhapura, Trimen

in 1881 (PDA). Trincomalee District: Mile 79 on Road A 4, Davidse 7558 (MO,

NY). Batticaloa District: Batticaloa, Trimen C.P. 678 p.p. (PDA, K). Monaragala

District: Crossing of Rd. A 2 and Kirindi Oya at Mile 183/2, ca. 15 miles S

of Wellawaya, 400 ft. alt., Davidse 7760 (MO, NY, PDA, US). Ratnapura District:

Raigam Korale, Thwaites C.P. 678 p.p. (PDA). Galle District: Bentota, Ballard

1507 (K, NY). Hambantota District: N of Kataragama, Wirawan 644 (PDA);

Ruhuna National Park: Block 3, Cooray 23 May 1968 (PDA, US).

Distribution. Tropical Africa, Madagascar through Indian Subcontinent to

southern China and the southern Ryukyus, as well as to Malesia, northern Australia

and the Pacific Islands.

45. Mariscus compactus (Retzius) Boldingh, Zakfi. Landb. Java 77, 1916; Druce,

Rep. Bot. Exch. Club Brit. Isls. 1916: 634, 1917; Fischer in Gamble, Fl. Madras

9: 1645, 1931. ————- Fig. 8

Cyperus compactus Retzius, Osb. Bot. 5: 10, 1789; Kiikenthal, Pflanzenr. 4

20), 101 Heft: 423, 1936; Kern, Fl. Males. I, 7 (3): 638, 1974. Cyperus

dilutus Vahl, Enum. Pl. 2: 357, 1806. —— Mariscus microcephalus Pres]. Reliq.

Haenk. 1: 182, 1828; C. B. Clarke, Fl. Brit. India 6: 624, 1893; Trimen & Hook. f.,

Handb. Fl. Ceylon 5: 40, 1900. —— Mariscus dilutus (Vahl) Nees in Wight,

Contrib. Bot. India 90, 1834.

Badulla District: ca. 8 miles E of Mahiyangana, 3.5 miles N of Mile 53 on

Mihiyangana-Padiyatalawe Rd., T. Koyama et al. 13979 (NY, PDA, US).

Distribution. India, southern continental China, Taiwan and Malesia.

This species, easily recognizable by its characteristic globose spikes, occurs

very sporadically in the low countries in Ceylon. I have not seen Gardner's

collection from Kornegalle, which was cited by both Thwaites and Trimen as

C.P. 815, and which seems to be the only other documentation of this species

from Ceylon.

46. Mariscus squarrosus (L.) C. B. Clarke, Fl. Brit. India 6: 623, 1893, concerning

the basionym.

154 Gardens’ Bulletin, Singapore — XXX (1977)

Cyperus squarrosus L., Cent. Pl. 2: 6, 1756; Kern, Fl. Males. I, 7 (3): 631,

1974. Cyperus aristatus Rottboell, Descr. Pl]. Rar. Progr. 22, 1772; C. B.

Clarke, Fl. Brit. India 6: 606, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 24,

1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 502, f. 55 F -J, 1936. Mariscus

aristatus (Rottboell) Chermezon, Bull. Soc. Bot. France 85: 366, 1938; Tang &

Wang, FI. Reipubl. Pop. Sin. 11: 178, pl. 59, 1961.

Jaffna District: north shore of Jaffna Lagoon, ca. 10 miles SE of Navatkuli,

sea level, T. Koyama et al. 14037 (NY, PDA, US); Keerimalai to Point Pedro,

Clayton 5205 (K). Wilpattu National Park: near Sadpuda Kallu, 2 miles E of

Kattankandal Kulam, 30 m alt., Fosberg et al. 50830 (NY, US). Trincomalee

District: Trincomalee, Glenie C.P. 966 ex p. (PDA). Puttalam District: Puttalam,

Ferguson in 1881 (PDA). Polonnaruwa District: Polonnaruwa, Clayton 5116 (K,

PDA); 13 miles E of Polonnaruwa on Trincomalee Rd., Clayton 5135 (K, PDA);

ca. 1 mile E of Kolakanaweli, Mile 56/7 on Batticaloa Rd., T. Koyama 13575

(NY). Mannar District: Aruvi Aru, Mile 125 on Rd. A 14, T. Koyama et al.

13934 (NY, PDA, US). Anuradhapura District: Galpitigala, a few miles from

Ritigala, Ballard 1451 (K, NY). Vavunia District: Palayanalankulan, Mile 120 on

Rd, A 14, T. Koyama et al, 13938 (NY, PDA, US). Kandy District: Hantane,

Thwaites C.P. 966 (K, PDA); Giragama, Amaratunga 359 (PDA). Hambantota

District: Tissamaharama, Trimen Dec. 1882 (PDA); Rhuhuna National Park,

Kohombagaswala, Block I, Cooray et al, 22 Jan. 1969 (US); Uraniya, Cooray

22 Mar. 1970.

Distribution. Widely distributed in tropical and subtropical regions of the

world, extending also into temperate regions in the Americas, where the range

covers from southern Canada south to Argentina and Chile. In Ceylon common on

wet sandy soil in open grasslands.

In the Linnaean Herbarium the specimen marked as “‘sguarrosus” in Linnaeus’

hand represents this taxon, while the second sheet with a single inflorescence is

Cyperus maderaspatanus. Consequently, Cyperus squarrosus is the correct name

of what has generally been called C. aristatus. Although I place this species in

Mariscus following Chermezon and Wang & Tang (locis cit.), as was already

pointed out by O’Neill (Rhodora 44: 47, 1942) and Kern (op. cit.) the caducity

of the rhachillas of this species is not constant. In Ceylonese specimens I noticed

in a few specimens that the glumes fall off apart from the rhachilla, which

eventually falls off the rhachis. This species, which is characteristically intermediate

between Cyperus and Mariscus, requires more detailed anatomical studies which

might elucidate the systematic link between the two genera.

47. Mariscus sumatrensis (Retzius) T. Koyama, comb. nova

Scirpus cyperoides L., Mant. 2: 181, 1771. [Kyllinga umbellata Rott-

boell, Descr. Icon. Rar. Nov. Pl. 15, 1773, excluding ¢. 4 f. 2. Illegitimate name.]

Kyllinga sumatrensis Retzius, Obs. Bot. 4: 13, 1786. Kyllinga

umbellata Rottb. var, sumatrensis (Retz.) Willdenow, Sp. Pl. 1: 258, 1797.

[Mariscus umbellata (Rottb.) Vahl, Enum. 2: 376, 1806. Combination based on

illegitimate name. ] Mariscus sieberianus Nees [Linnaea 9: 286, 1835, nomen

nudum] ex C. B. Clarke, Fl. Brit. India 6: 622, 1893, & Illustr. Cyper. t. 23 f. 5-6,

1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 42, 1900. Mariscus cyperoi-

des (L.) Urban, Symb. Antill. 2 (1): 164, 1900. Not of Dietrich, 1833.

Cyperus cyperoides (L.) O. Kuntze, Rev. Gen. Pl. 3 (2): 333, 1898; Kiikenthal, .

Pflanzenr. 4 (20), 101 Heft: 514, 1936; Kern, Fl. Males. I, 7 (3): 642, 1974.

Nuwara Eliya District: Hakgala, Alston 633 (PDA), Mueller-Dombois May

1968 (PDA, US); Talawakele, Bond 17 Feb. 1944 (PDA).

Distribution. Tropical Africa, tropical and subtropical Asia, Malesia, northern

Australia; also introduced to the West Indies. In Ceylon rarely found in up

countries.

Ceylon Cypereae

155

Fig. 8. Mariscus compactus (Retzius) Boldingh. A, habit. B, orifice of a cladoprophyll

at the base of umbel ray. C, spikelet. D, prophyll. E, lateral view of two glumes showing

a base forming the wings on the rhachilla. F, glume. G, fruiting pistil. H, achene. Scales

= | mm.

156 Gardens’ Bulletin, Singapore — XXX (1977)

Fig. 9. Mariscus clarkei (T. Cook) T. Koyama. A, habit. B & C, two views of spikelet.

D. eewrone subtending a spikelet. E, prophyll. F, fruiting glume. G, empty glume. H, achene.

es = 1 mm.

Ceylon Cypereae 157

When this taxon is placed in Mariscus, its earliest valid name, Scirpus

cyperoides, cannot be transfered because of Dietrich’s homonym. Next earliest name,

Kyllinga umbellata, is illegitimate since Rottboell cited Scirpus cyperoides in the

synonymy. Kyllinga sumatrensis, therefore, needs to be transferred to Mariscus to

form the correct name of this species under the genus.

48. Mariscus cyperinus (Retzius) Vahl, Enum. Pl. 2: 377, 1806; C. B. Clarke, FI.

Brit. India 6: 621, 1893, & Illustr. Cyper t 12 f. 3-4, 1909: Trimen & Hook. f.,

Handb. Fl. Ceylon 5: 42, 1900; C. E. C. Fischer in Gamble, Fl. Madras 9: 1644,

1931.

Kyllinga cyperina Retzius, Obs. Bot. 6: 21, 1791. Cyperus cyperinus

(Retzius) Suringar, Het. Gesl. Cyper. Mal. Archip. 154, ¢. 6 f. 10, 1898; Kiikenthal,

Pflanzenr. 4 (20), 101 Heft: 518, 1936; Kern, Fl. Maels. L 7 (3): 641, 1974.

48a. Subsp. cyperinus.

Ceylon: without definite locality, Trimen C.P. 816 (PDA). Kandy District:

ca. 2 miles N of Hunnasgiriya, near Mile 23, 380 m alt., Davidse 8457 (MO, NY);

Pusselawa, Alston 1170 (PDA); Peradeniya, Thwaites CP. 816 (K, PDA), Amara-

tunga 643 (PDA), Alston 593 (PDA); Morakande Estate, Alston 594 (PDA);

Gannoruwa, Alston 851 (PDA); Gampola, Alston 592 (PDA). Matale District:

33/7 Midlands, 1200 m alt., Tirvengadum et al. 14 (US); Matale, Tirvengadum et al.

16 (PDA). Nuwara Eliya District: Ramboda Pass, 3200 ft. alt.. Ballard 1142A

(K, NY). Badulla District: Rawanaella Waterfall, ca. 2 miles SE of Ella, 750 m

alt., Davidse et al. 8868 (NY, PDA, US). Colombo District: Dewalgama, Amara-

tunga 1162 (PDA): Ekala, Alston 2389 (PDA).

Distribution. From India to Malesia and northern Australia, also in Eastern

Asia extending northwards to southern China and the Ryukyus. Open grassy places

in hilly countries.

48b. Subsp. laxatus (C. B. Clarke), T. Koyama, stat. nov.

Cyperus umbellatus C. B. Clarke var. laxatus C. B. Clarke, Journ. Linn. Soc.

21: 201, 1888. Cyperus cyperinus Suringar forma curvata Suringar, Het

Gesl. Cyper. Mal. Archip. 156, 1898; Kern, Fl. Males. I, 7 (3): 624, 1974.

Mariscus tenuifolius Schrader ex Nees, Fl. Brasil. 2 (1): 46, 1842: C. B. Clarke,

Fl. Brit. India 6: 622, 1893; Trimen & Hook. f., Handb. Fi. Ceylon 5: 43, 1900;

C. E. C. Fischer in Gamble, Fl. Madras 9: 1644, 1931.

Kandy District: Hunnasgiriya, collector unknown 590 (PDA); Peradeniya,

Alston 387 (PDA), Thwaites C.P. 817, ex p. (PDA); Maturata, Thwaites CP.

817 ex p. (PD). Matara District: Weligama, Alston 1172 (PDA). Monaragala

District: ca. 3 miles W of Wellawaya, near Mile 135/10, 990 ft. alt., Davidse 7741

(MO, NY). Ruhuna National Park: Ecology Project Prot R19, Cooray 11 Dec.

1969 (US).

Distribution. India to Malesia; not as common as the typical phase.

Subspecies /axatus differs from the typical phase in its narrow habit with long-

creeping slender stolons and the few-spiked contracted umbels that tend to become

pseudo-lateral being subtended by the lowest leafy bract continued down to the

culm. As far as the Ceylonese plants are concerned this combination of characters

appears to be rather constant, on which basis I regard this slender phase to be a

subspecies of M. cyperinus.

49. Mariscus paniceus (Rottboell) Vahl, Enum. Pl. 2: 373, 1806; C. B. Clarke, FI.

Brit. India 6: 620, 1893, incl. var. roxburghianus, & Illustr. Cyper. t. 22 f. 1-2,

1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 41, 1900, incl. var. roxburghianus.

158 Gardens’ Bulletin, Singapore — XXX (1977)

[Schaenoides paniceus Rottboell, Descr. Pl. Rar. Progr. 15, 1772, provisional

name. | Kyllinga panicea Rottboell, Descr. Icon. Rar. Nov. Pl. 15, ¢. 4 f. J,

1773. Cyperus paniceus (Rottboell) Bockeler, Linnaea 36: 381, 1870, in

part, incl. basionym; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 526, 1936; Kern, Fl.

Males. I, 7 (3): 643, 1974.

Anuradhapura District: Ritigala Nature Reserve, 2400 ft. alt., Jayasuria 1710

(NY, PDA). Mannar District: Aruvi Aru, Mile 125 on Rd. A 4, T. Koyama et al.

13928 (NY, PDA, US). Polonnaruwa District: SE of Giritale Wewa, 7 miles N.W.

of Polonnaruwa, Fosberg & Ripley 51935 (US); Polonnaruwa, Ripley 339 (PDA);

Galoya Reservoir, near spilway, 270 m alt., Commanor 568 (NY). Nuwara Eliya

District: Ramboda Pass, 3200 ft., Ballard 1142A (K, NY); Hakgala Botanic

Gardens, Clayton 5783 (K, PDA); between Pusselawa and Ramboda, at Mile 32/8

on N. Eliya Rd., 1100 m alt. 7. Koyama & Herat 13613 (NY, PDA, US). Matale

District: ca. 8 miles ESE of Dambulla, ca. 900 ft. alt., Davidse 7433 (MO, NY).

Kandy District: Hantane, Thwaites C.P. 814 (K, PDA); Hewaheta, Thwaites C.P.

2878 (PDA); Maragala, Alston 1629 (PDA); Haragama, Alston 306 (PDA);

Aladeniya, Amaratunga 644 (PDA); between Alagalla and Balana Rd., Comanor

440 (PDA, US). Kurunegala District: Kurunegala, Thwaites C.P. 2878 ex p.

(PDA); Kurunegala Rock, Alston 699 (PDA); Wariyapola, Trimen Aug. 1883

(PDA); Siyambalatenna, Alston 596 (PDA). Ratnapura District: ca. 13 miles

N.E. of Deniyaya at Mile 64 on Rd. A 17, 1050 m alt., Davidse 7899 (MO, NY,

PDA, US). Galle District: Galle, Alston 289 (PDA). Ruhuna National Park:

Rugamtota, Cooray 16 Nov. 1969 (NY, US).

While in Ceylon I was able to investigate the variation ranges of leaf width

and the intensity of the umbels in this very common species. My conclusion

is that var. roxburghianus should not be retained. It was claimed to differ from

the typical phase in the relatively broader leaves and larger umbels.

50. Mariscus clarkei (T. Cook) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976.

Fig. 9

Cyperus clarkei T. Cook, Fl. Presid. Bombay 2: 873, 1908; Kikenthal,

Pflanzenr. 4 (20), 101 Heft: 540, f. 58, 1936. [Mariscus pictus Nees in

Wight, Contrib. Bot. India 90, 1834, in part excluding type.] [Mariscus

bulbosus C. B. Clarke, Fl. Brit. India 6: 620, 1893; C. E. C. Fischer in Gamble,

Fl. Madras 9: 1644, 1931. Not. of Steudel, 1855.] [Kyllinga bulbosa Vahl,

Enum. Pl. 2: 376, 1806, invalid name mentioned in note; ROmer & Schultes, Syst.

Veg. 2: 247, 1817, in note.]

Monaragala District ca. 11 miles W of Tanamalwila, 125 m alt., Davidse &

Sumithraarachchi 8812 (MO, NY, PDA, US). Hambantota District: Ruhuna

National Park, Kohombagaswala, Cooray 23 Nov. 1969 (NY, PDA, US).

Distribution, Confined to southern India and Ceylon.

This species, newly found in Ceylon, rarely grows on wet shallow soil over

rock outcrops. It is well demarcated by the peculiar bulb-shaped base, which

emits a few slender stolons. The umbel of this species resembles that of M.

ala and hence it is never confused with M. dubius, another bulbose Mariscus

in Ceylon.

ob 4 Maem dubius (Rottboell) Kiikenthal ex Fischer in Gamble, Fl. Madras 9:

Cyperus dubius Rottboell, Descr. Icon. Rar. Pl. 20, t. 4 f. 5, 1773; Kiikenthal,

Pflanzenr. 4 (20), 101 Heft: 563, 1936; Kern, Fl. Males. I, 7 (3): 643, 1974.

Cyperus kyllingaeoides Vahl, Enum. Pl. 2: 312, 1806. Mariscus dregeanus

Kunth, Enum. Pl. 2: 120, 1837; C. B. Clarke, Fl. Brit. India 6: 620, 1893, &

ney Cyper. t. 2] f. 1-6, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 39,

Ceylon Cypereae 159

Anuradhapura District: Ritigala Natural Reserve, on summit rock, 2500 ft.

alt., Jayasuria 905 (PDA). Polonnaruwa District: Lankatilaka, Silva 182 (PDA).

Kurunegala District: Kurunegala, Thwaites C.P. 2942 (K, PDA) & C.P. 855 (K,

PDA). Kandy District: Aladeniya, Amaratunga 642 (PDA); between Balane and

Alagalla, 440 m alt. Comanor 1194 (NY, US). Badulla District: Rawanaella Water-

fall, ca. 2 miles SE of Ella, 750 m alt., Davidse et al. 8866 (MO, NY, PDA, US):

Lunugala, Trimen Jan. 1888 (PDA). Monaragala District: ca. 11 miles W of

Tanamalwila, 125 m alt., Davidse et al. 8802 (MO, NY, US, PDA). Galle District:

Galle, Gardner C.P. 855 (K, PDA). Ruhuna National Park: Bambowa, Cooray

17 Nov. 1969 (NY, US); Rugamtota, Cooray 16 Nov. 1969 (NY, US).

Distribution. Tropical Africa through India and Indo-China eastwards to

Malesia.

52. Mariscus pedunculatus (R. Brown) T. Koyama, comb. nov.

Remirea maritima Aublet, Hist. Pl. Guian. Franc. 1: 45, ¢. 16, 1775. Not

Mariscus maritimus Miquel, 1860, nor of C. B. Clarke, 1896. Remirea

pedunculata R. Brown, Prodr. Fl. Nov. Holl. 236, 1810. Cyperus peduncu-

latus (R. Brown) Kern, Act. Bot. Neerl. 7: 798, 1958.

Jaffna District: Jaffna, Thwaites C.P. 3227 (K, PDA). Colombo District:

Colombo, Ferguson C.P. 3227 (PDA); Mt. Lavinia, Trimen Oct. 1881. (PDA).

Distribution. Southern Asia, Malesia and northern Australia; also in tropical

America from eastern Venezuela through the Guianas south to Brazilian Parana

Sand dunes and sandy shores of sea coast.

The cyperoid morphology exhibited by this species was fully discussed by

Kern (op. cit.), with whom I concur. When Cyperus sensu lato is divided into a

few genera, this species properly belongs to Mariscus because of its continuous

rhachilla jointed at its base.

KYLLINGA Rottb.

1. Inflorescence open, umbelliform with elongated rays ............... 53 K. hyalina

1. Inflorescences congested in a head

2. Glumes not winged

3. Rhizome short; culms tufted

4. Central spike cylindrical or oblong, 8-15 mm long, ca. 5 mm wide;

spikelets broadly ovate, 2 mm long; culms slightly or hardly

enlatmery at. Gase rt). £5.66 0265.5.0.5..5- 54 K. odorata ssp. cylindrica

4. Central spike globose or ovoidal, 5-6 mm long, 3-4 mm across;

spikelets oblong-elliptical, 1.7-2 mm long; culms bulbose-thickened

(Ts RES \ 6 See Os ae epee eee a Oe 55 K. triceps

3. Rhizome horizontally creeping or stoloniferous; culms distantly or close-

ly arranged in one row along rhizome

5. Culms 30-50 cm tall, close together

Gem tier Practs 2 06:3 Ges eee. ieee eae 56 K. melanosperma

i Fe Ode ee 57 K. polyphylla

5. Culms as a rule 7-30 cm tall, more or less spaced ...................

Te as J cemataele ihe inion 58 K. brevifolia

SS ee eee pats. bt alti ea 59 K. nemoralis

160

’ Bulletin, Singapore — XXX (1977)

Gardens

iA Z

Afe a

ti Lk

KY 44

YT:

“fp i

/. F.

Fig. 10. Kyllinga hyalina (Vahl) T. Koyama. A, habit. B, portion of rhachis with

bracteoles and prophylls remaining after spikelets falling off. C, spikelet. D, prophyll. E,

lateral view of two glumes. F, lateral view of achene. Scales = 1 mm,

Ceylon Cypereae 161

33: as “tage yep (Vahl) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976.

ig.

Cyperus hyalinus Vahl, Enum. Pl. 2: 329, 1806; Trimen & Hook. f. Handb.

Fl. Ceylon 5: 19, 1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 498, 1936: Kern,

Fl. Males. I, 7 (3): 655, t. 68, 1974. “Cyperus pumilus L.”’: sensu Nees in

Wight, Contrib. Bot. Ind. 74, 1834. “Pycreus pumilus’’ C. B. Clarke, FI.

Brit. Ind. 6: 591, 1893, concerning description. Pycreus hyalinus (Vahl)

Domin, Bibl. Bot. Heft 85: 417, 1915, in obs. Queenslandiella mira Domin,

Bibl. Bot. Heft 85: 416, 4 JJ f. 7-13, 1915. Mariscopsis suaveolens Cher-

mezon, Bull. Mus. Paris 25: 60, 1919. Mariscopsis hyalinus (Vahl) Ballard,

Kew Bull. 1932: 457, 1932. Queenslandiella hyalina (Vahl) Ballard in

Hook., Icon. Pl. 33: t. 3208, 1933.

Anuradhapura District: Anuradhapura, Trimen 18 Dec. 1881 (PDA). Kuru-

negala District: Kurunegala, Trimen C.P. 3787 (K, PDA); Kuliyapitiya, Simpson

9159 (PDA). Ruhuna National Park; near turn off to Andunoruwa Wewa, Cooray

10 Dec, 1969 (NY, PDA, US); Ecology Project Block I, Plot R 13, Cooray 7

Dec. 1969 (US).

Distribution, Tropical East Africa, Mascarene Is., India, Indo-China, Malesia

and northern Australia. Sporadically occurring in dry grasslands.

When we split the genus Cyperus of broad meaning into several genera of

smaller size, the systematic attribution of Cyperus hyalinus deserves special mention.

Morphologically this species doubtlessly falls within the generic criterion of

Kyllinga, from which it slightly deviates in having open inflorescences and in the

spikelets of which nearly all the glumes are fruit-bearing. However, in the genus

Kyllinga, umbel rays really develop as seen in K. transitoria for instance, and

further where in all the cyperoid genera both open and head-like inflorescences

occur without any clear border, thus showing that these conditions of inflorescences

do not warrant generic segregation. Similarly the number of fruit-bearing glumes

in a spikelets also does not serve as a distinguishing character between genera. In

Mariscus, the genus most closely r€élated to Kyllinga, both multi- and few-flowered

spikelets coexist (cfr. M. cyperinus vs. M. javanicus, for example). It is therefore

permissible to include Cyperus hyalinus in Kyllinga in spite of the multi-flowered

spikelets. In reality Cyperus hyalinus strikingly resembles Kyllinga ssp. not only

in its bilaterally flattened oblong-elliptic achenes but also in its thinly membranous

glumes with several prominent veins on both sides of the midrib.

54. Kyllinga odorata Vahl, Enum. Pl. 2: 382, 1906.

This species, occurring in tropical America and Africa, is represented in

Ceylon by:

subsp. cylindrica (Nees ex Wight) T. Koyama, comb. nov.

Kyllinga cylindrica Nees ex Wight, Contrib. Bot. Ind. 91, 1834; C. B. Clarke,

Fl. Brit. Ind. 6: 588, 1893, incl. var. subtriceps Nees; Trimen & Hook. f., Handb.

Fl. Ceylon 5: 44, 1900. Kyllinga odorata Vahl var. cylindrica (Nees)

Kiikenthal ex Merrill, Journ. Str. Branch Roy. As. Soc. 76: 80, 1917.

Cyperus sesquiflorus (Torrey) Mattfeld & Kiikenthal var. cylindricus (Nees)

Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 593, 1936. Cyperus sesquiflorus

Mattfeld & Kiikenthal subsp. cylindricus (Nees) T. Koyama, Bot. Mag. Tokyo

83: 187, 1970. Cyperus sesquiflorus Mattfeld & Kiikenthal var. subtriceps

(Nees) T. Koyama, Quart. Journ. Taiwan Mus. 14: 191, 1961; Kern, FI. Males.

I, 7 (3): 659, 1974.

Kandy District: Pusselawa, Alston 19 Sept. 1926 (PDA). Badulla District:

Haldummulla, Thwaites C.P. 3754 (K, PDA). Nuwara Eliya District: Ohiya,

Mueller-Dombois 18 May 1968 (PDA, US); Hakgala, Simpson 9041 (PDA);

162 Gardens’ Bulletin, Singapore — XXX (1977)

Ramboda, along N. Eliya Rd., 1180 m alt., T. Koyama & Herat 13619 (NY, PDA,

US). Hambantota District: Ruhuna National Park, Cooray 30 November 1967

(PDA, US).

Distribution. Tropical Africa, tropical and subtropical Asia.

The morphological differences between subsp. cylindrica and the typical phase

were discussed in my previous paper (Koyama, op. cit. 1970).

55. Kyllinga triceps Rottboell, Descr. Icon. 14, t. 4 f. 6, 1773; C. B. Clarke, FI.

Brit. Ind. 6: 587, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 45, 1900.

[Schaenoides triceps Rottboell, Descr. Pl. Rar. Progr. 15, 1772, invalid name.]

Cyperus triceps (Rottb.) Endlicher, Cat. Hort. Acad. Vindb. 1: 94, 1842;

Kikenthal, Pflanzenr. 4 (20), 101 Heft: 578, 1936; Kern, Fl. Males. I, 7 (3): 659,

1974.

Mannar District: Aruvi Aru, Mile 125 on Rd. A 14, T. Koyama et al. 13932

(NY, PDA, US). Polonnaruwa District: Polonnaruwa Sacred Area, Sect. 3A,

61 m alt., Ripley 334 (US); Polonnaruwa-Trincomalee Rd., Clayton 5136 (K,

PDA). Matale District: 33/7 Midlands, Tilyengadum et al. 12 (US). Puttalam

District: Anamaduwa, Trimen Aug, 1883 (PDA); Puttalam, Brodie C.P. 3234 in

part (PDA). Colombo District: Colombo, Thwaites C.P. 3234 (K, PDA). Mona-

ragala District: ca. 11 miles W of Tanamalwila, 125 m alt., Davidse et al. 8810

(MO, NY, PDA). Ruhuna National Park: Komawa Wewa, Cooray 16 Nov. 1969

(PDA, US); Gonalabbe, Cooray 24 Nov. 1969 (NY, PDA); Yala, Andunoruwa

Wewa, Comanor 673 (NY, US).

Distribution. From tropical Africa through India and Indo-China north-

eastwards to southern China, and eastwards to Malesia and northern Australia.

56. Kyllinga melanosperma Nees in Wight, Contrib. Bot. India 91, 1834; C. B.

Clarke, Fl. Brit. India 6: 588, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 45,

1900.

Cyperus melanospermus (Nees) Suringar, Het. Gesl. Cyperus Mal. Archip.

50, t. 2 f. 8, 1898; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 583, 1936; Kern, FI.

Males. I, 7 (3): 655, 1974.

Kandy District: Peradeniya, Lower Hantana Rd., near the waterfall, 535 m

alt., Comanor 692 (NY, US); between Pusselawa and Ramboda, Comanor 950

(PDA); between Laxapana and Maskeliya Rd., Kostermanns 24079 (PDA);

Ambagamuwa, Amaratunga 2014 (PDA), between Udahentenna and Nawalapitiya,

Amaratunga 909 (PDA); Aladeniya, Amaratunga 1643 (PDA). Nuwara Eliya

District: Hakgala, Wheeler 12371 (PDA); Nuwara Eliya, Thwaites C.P. 818 (K,

PDA); Horton Plains, forest opposite to the Horton Plains Farm, along Pattipola

Rd., 7200 ft. alt., 7. Koyama 13506 (NY, PDA, US); base of Hakgala Mtn. 1870 m

alt., Davidse et al. 7981 (MO, NY, PDA, US). Badulla District: ca. 3 miles W of

Koslanda, along Rd. A 4, ca. 900 m alt., T. Koyama et al. 14033 (NY, PDA, US).

Colombo District: Wattala, Comanor 1153 (PDA, US); Danowita; Amaratunga

193 (PDA); Kimbulapitiya, Amaratunga 2157 (PDA). Ratnapura District: 11

miles E of Diniyaya, near Mile 62 on Rd. A 17, 750 m alt., Davidse 7875 (MO,

NY, PDA, US). Galle District: Corbett’s Gap. 4500 ft. alt., Ballard 1052 (K, NY).

__ Distribution. Tropical and subtropical Africa, southern Asia, Fiji and Malane-

sia. In Ceylon common on the grassy beds of montane forests.

This species, well characterized by its relatively thick leafless culms and

large heads, can be easily located in the field because of its strong odor of

essential oil, which is contained in the thick rhizome.

ee ee

Ceylon Cypereae 3 163

57. Kyllinga polyphylla Willdenow ex Kunth, Enum. Pl. 2: 134, 1937.

Kyllinga aromatica Ridley, Trans. Linn. Soc. II, Bot. 2: 146, 1884.

Cyperus aromaticus (Ridley) Mattfeld & Kiikenthal, Pflanzenr. 4 (20), 101 Heft:

581, 1936; T. Koyama, Bot. Mag. Tokyo 83: 186, 1970; Kern, Fl. Males. I, 7 (3):

656, 1974.

Nuwara Eliya District: between Pussellawa and Ramboda, at Mile 28/10 along

Kandy-N. Eliya Rd., 950 m alt., Comanor 328 (NY, PDA, US).

Distribution, E. Africa. Also eoHEeS to Singapore, Ceylon, Solomon Islands,

Samoa and Fiji.

This African species can be separated from the closely allied indigenous

species, K. melanosperma, by its head subtended by 6 to 8 leafy bracts in contrast

to 2 or 3 in the latter. In Ceylon it is thus far known only by the collection cited

above, and is apparently of rather recent introduction.

58. Kyllinga brevifolia Rottboell, Descr. Icon. Rar. Nov. Pl. 13, t. 4 f. 3, 1773;

C. B. Clarke, Fl. Brit. India 6: 588, 1893, & Illustr. Cyp. ¢. J f. 1-4, 1909; Trimen

& Hook. f., Handb. Fl. Ceylon 5: 45, 1900.

[Schaenoides brevifolius Rottb., Descr. Pl. Rar. Progr. 15, 1772, invalid

name. |] Cyperus brevifolius (Rottb.) Hasskarl, Cat. Hort. Bogor. 24, 1884;

Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 600, 1936; Kern, Fl. Males. I, 7 (3): 656,

f. 70, 1974.

Wilpattu National Park: Kali Villu, Wirawan et al. 990 (US). Mannar

District: Aruvi Aru, Mile 125 on Rd. A 4, T. Koyama et al. 13926 (NY, PDA,

US). Kandy District: Peradeniya, Upper Hantana Rd., 550 m alt., Comanor 315

(NY, US); Ambewela, Mueller-Dombois 12 Jan. 1968 (PDA, US). Nuwara Eliya

District: Kande Ela Reservoir, Mueller-Dombois 18 May 1968 (PDA). Colombo

District: Dompe, Amaratunga 2182 (PDA); Ranmuthugal, Amaratunga 2221

(PDA). Galle District: Galle, Trimen C.P. 3755 (K, PDA).

Distribution. Pantropic; in Asia extending northwards into the warm regions

of continental China and the temperate region of Japan.

This cosmopolitan weedy sedge is rather poorly documented in Ceylon. Since

it frequently grows in and around rice fields, it is quite possible that this sedge

extended its range with the introduction of rice as assumed for Cyperus iria and

Fimbristylis miliacea.

59. Kyllinga nemoralis Gg. R. & G. Forster) Dandy ex Hutchinson & Dalziel, FI.

W. Trop. Africa 2: 486, in Key, & 487, 1936.

[Kyllinga monocephala Rottboell, Descr. Icon. Rar. Nov. Pl. 13, t. 4 f. 4,

1773, invalid name; C. B. Clarke, Fl. Brit. India 6: 588, 1893 & Illustr. Cyper.

t, 2 f. 1-2, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 44, 1900.] Cyperus

kyllingia Endlicher, Cat. Hort. Acad. Vindb. 1: 94, 1842; Kikenthal, Pflanzenr.

4 (20), 101 Heft: 606, f. 64, 1936; Kern, Fl. Males. I, 7 (3): 659, 1974.

Thryocephalon nemoralis J. R. & G. Forster, Char. Gen. Pl. 130, 1776.

Trincomalee District: Mutur, Wheeler 12425 (PDA). Kandy District:

Roseneath, 2101 ft. alt., Ballard 1014 (K, NY). Monaragala District: ca. 3 miles

W of Wellawaya at Mile 135/10 on Rd. A 4, 990 ft. alt. Davidse 7728 & 7732

(MO, NY, PDA, US). Ruhuna National Park: Rugantota, Block I, Cooray 16

Nov. 1969 (NY, US); Ecology Project Block I, Rugantota on Minik Ganga,

Fosberg et al. 50191 (NY, US).

Distribution. Pantropic, relatively scarce in tropical America.

It is unfortunate that to this common species, which has long been known as

Kyllinga monocephala, must be assigned an unfamiliar epithet as its correct name.

Rottboell’s Kyllinga monocephala is invalid as the name was published with an

earlier legitimate name, Schoenus coloratus L., cited in the synonymy.

164 Gardens’ Bulletin, Singapore — XXX (1977)

Literature Cited outside Taxonomic Enumerations

Kern, J. H. 1974. Cyperaceae. Flora Malesiana Ser. I, 7 (3): 435-753.

Koyama, T. 1970. Beitrage zur Cyperaceenflora von Ceylon. Bot. Mag. Tokyo

83: 183-192.

1974. Additional notes on Ceylonese Cyperaceae. Bull. National

Sci. Mus. Tokyo 17: 65-73.

Kiikenthal, G. 1935-36. Cyperaceae-Scirpoideae-Cypereae. Pflanzenr. 4 (20),

101 Heft: 1-671.

Trimen, H. & J. D. Hooker. 1900. A Handbook to the Flora of Ceylon 5: 1-477.

The Limestone Hill Flora of Malaya I

S. C. CHIN

University of Malaya, Kuala Lumpur

Malaysia

CONTENTS

‘PAGE

Abstract : - - . - - : - - - 165

Preface and Acknowledgements’ - - - - > - : 166

SECTION | — GENERAL

1. Introduction’ - : - - - - - - 167

2. Geology, origin and distribution of the limestone hills - - 172

3. Vegetation - - - ‘ = i > , 174

4. Phytography and size of the flora - . . - : 182

5. Affinities of the floristic components to limestone’ - - - 189

SECTION 2— The Flora

Pteridophytes : re : * - - - 198

Gymnosperms - - - - - - - - 218

Abstract

Limestone habitats, chiefly karst towers, in the Malayan Peninsula, support a rich flora

of about 1216 species of vascular plants, in 582 genera and 124 families. Phanerogam

families total 119, representing 61.6% of the total number of phanerogam families recorded

for the whole Malayan flora; only 72 families are not represented on limestone, and these

are mostly aquatics or small rare groups. Specific endemism among the limestone plants

is 21.4% (261 species), and of these 10.7% (130 species) are found only on limestone. There

are 335 species “characteristic” of the limestone flora, and 254 of these (20.8%) are restricted

to limestone.

The limestone vegetation is described and classified into “types” and secondary vegetation

and succession is discussed. Pioneer species on limestone include those found on other

disturbed terrestrial habitats in Malaya. Some plants found on limestone are found elsewhere

in Malaya only at significantly higher elevations. A discussion of the geological origin and

distribution of the Malayan limestone areas is also included.

165

166 Gardens’ Bulletin, Singapore — XXX (1977)

PREFACE AND ACKNOWLEDGEMENTS

This publication is based on the dissertation accepted for the Master of

Science Degree in 1973 by the School of Biological Sciences, University of Malaya.

I am grateful to Professor van Steenis, external examiner, for his critical comments

and helpful suggestions.

My work was supervised by Dr. B. C. Stone who introduced me to this topic,

provided an unfailing source of guidance, and who also permitted me free access

to his most remarkable collection of botanical literature which has been an

inspiring and invaluable source of reference. I am also grateful for his encourage-

ment to publish this work.

The Department of Botany, University of Singapore allowed me the use of

their herbarium and Dr. H. Keng helped with the identification of several Labiatae.

He also generously provided me with the records of the collection made by the

UNESCO 1962 limestone expedition to Ulu Kelantan. These consisted of a map,

several note books and a set of duplicate labels to the collection.

The Director of the Botanic Gardens, Singapore, gave me permission to work

and the staff helped me in the herbarium on several occasions.

Dr. T. C. Whitmore and subsequently Dr. Francis Ng allowed me to use

freely the facilities of the herbarium at the Forest Research Institute, Kepong, and

manuscripts of their “Tree Flora of Malaya’, vol. 2 then in press. Dr. Whitmore

also helped with the identification of some Garcinias, Euphorbiaceae and Palms;

Dr. Ng with some Ebenaceae and other members of the staff with some general

collections.

Dr. K. U. Kramer of the State University, Utrecht, Netherlands, identified

several fern specimens and Dr. T. Shimizu of Shinshu University, Japan, provided

literature and helped with the identification of a specimen of Impatiens.

With Dr. P. R. Wycherley, formerly of the Rubber Research Institute, Kuala

Lumpur, I had interesting discussions and he also introduced me to the Johore

limestone.

Mr. W. Swinson, Project Manager for the South East Johore Project, then

engaged in a Master Plan Study of South East and Central Johore for the Govern-

ment of Malaysia and State Government of Johore and his staff provided maps,

transport, guides and equipment for two trips into the Gunong Sumalayang

limestone outcrops in Johore. Mr. Swinson also provided unpublished data on the

locality. 7 |

Dato Haji Wan Hassan bin Abdul Halim, the State Forest Officer of Johore

gave permission to enter and botanize in the Johore limestone area.

Incik Mahmud bin Sidek and Incik Badaruddin, field investigator and

herbarium assistant respectively, assisted in several excursions. Friends, particularly

J. Boey, provided help and interesting company on numerous trips to the limestone.

To all these helpful persons. J express my most grateful thanks.

I must also state my appreciation for the comments of Dr Chang Kiaw Lan

editor. | |

Most of this work was carried out during the tenure of a tutorship in the

School of Biological Sciencés, Universiti Malaya.

Malayan Limestone Flora 167

SECTION 1— General

I. INTRODUCTION |

Background to the flora

The limestone landscape in Malaya is very distinctive. Typically the hills are

tower-like, rising from the surrounding scenery, with sheer rock walls and often

jagged summits. These ‘tower karst’ formations occur as isolated crags or are

grouped together into small massifs — all products of erosion of calcareous rock.

Erosion too is responsible for the presence of caves that often characterize limestone

hills. These caves have, since prehistoric times, often been used as human dwellings,

and as much as the vegetation on the hills provide clues to the history and evolution

of the Malayan flora, provide evidence for an insight into prehistoric Malayan

culture.

Most of the past work on the Malayan limestone concerned the geology,

origin and age of the rock, e.g. Scrivenor (1931), Paton (1961, 1964), Gobbett

(1965) and Hutchison (1963, 1968). Other investigators have concerned them-

selves with cave archaeology, e.g. Evans (1920), Tweedie (1940) and Peacock

(1965), or cave ecology and fauna, e.g. Bullock (1963, 1965), Dunn (1965),

McClure (1965) and Tweedie (1947).

The only separate work on the flora is that of Henderson (1939). This

contains an introduction giving the collecting localities, with notes on the vegetation

and the characteristics of the flora. The localities are introduced state by state; the

chief hills are named and the major collectors mentioned. There is however,

no map of the distribution of the hills mentioned. The vegetation is described;

there is a division of the hills into two main groups, the ‘wet’ and ‘dry’ hills. The

frequency of the occurrence of the plant families (restricted to the spermatophytes)

is briefly summarized.

A total of about 745 species is recorded from the limestone by Henderson.

The species are presented in a check-list with short distributional notes; the

arrangement of the families follows Ridley (1922-25), while the genera and

species are arranged in alphabetical order. Of the 745 species recorded about 195

(26%) in the Peninsula are known only from limestone. The species endemic and

confined to the limestone number about 130.

Apart from this study, species distribution on limestone is sometimes men-

tioned in floristic or monographic works on the local flora. Thus this well known

habitat is, botanically, relatively unworked.

The history of botanical exploration on the Malayan limestone hills began in

August 1880 when Kunstler, collector for Sir George King, made a collecting trip

to the Gopeng limestone in Perak (the exact locality is unknown). Other early

collectors (before 1900) include Curtis, Fox, Kelsall, Ridley and Wooldridge. Up

to the present day more than 60 collectors have left their mark on this history.

Their efforts have accumulated about 4,500 numbers (an estimate based on my

having seen about 4000 numbers and assuming that I have not seen about 10%

of the total); of this the writer contributed 1550 numbers.

The total area of limestone in Malaya, considering the average height to be

243.8 m (800 ft), is about 260 sq. km. (an estimate based on Scrivenor, in Burkill,

1935). Therefore per unit area the limestone may seem to have been very intensively

botanised, with about 1962 numbers per 100 sq. km. (compared with the average

of 175 numbers per sq. km. for the whole of Malaya, Steenis-Kruseman, 1973).

These figures are, however, deceptive and misleading when the richness of the

limestone flora is taken into account. The total number of collections from the

Malay Peninsula is about 232,000 (a rough estimate based on Steenis-Kruseman,

1973) and the total species present in Malaya is estimated to be between

8000-8500 (Keng, 1970). This total number of species is an estimation only and

is more than the number of species presently known. Thus this shows that on the

168 Gardens’ Bulletin, Singapore — XXX (1977)

average, each species is represented by 26-29 numbers, On the other hand, 1216

species are recorded from the limestone (present study) which are represented by

about 4500 numbers, thus giving an average of about 4 numbers per species. Even

if only the characteristic species (with affinities for the limestone habitat) were

considered (they number 335 and the total numbers collected would also drop

far below 4500) and the total numbers collected be retained at 4500, the average

number of specimens per species will be 13, which is still low.

Besides low total numbers collected with respect to the extremely rich flora,

many of the limestone species have a very local distribution which intensifies the

problem of insufficient collection. Though most of the limestone localities have

been visited by botanists, the total number of hills actually climbed and collected

on is only about 50, and the number of islands (in Langkawi) visited is about 25

(the writer has visited 25 hills and 12 islands, some repeatedly). There are no

estimates of the total number of limestone hills in Malaya, but they probably

number well over 200, and Langkawi consists of 99 islands (though not all are

limestone, a large percentage are). Thus we have a situation where many hills,

islands and localities have been repeatedly visited while others have not been

botanised at all. The “‘popular” localities include Bukit Takun, Gunong Rapat,

Gua Musang, Pulau Dayang Bunting (around the lake), Pulau Jerkom and Pulau

Timun. However, this is not to say that these relatively frequently visited spots are

sufficiently botanised. For instance, recently I discovered a clump of Oncosperma

horridum on the summit of Gua Batu (which rates high on the list of popular

localities), a species which has not been recorded on the limestone before. Con-

sidering that Gua Batu is less than 2.59 sq. km (1 sq. mile), in area, this clearly

shows that the many rugged faces of a limestone hill can conceal even the most

conspicuous plant.

Therefore in relation to its rich flora, extremely varied habitat and the fact

of uneven local exploration, the limestone vegetation is probably the least botanised,

and hence botanically known, of all the vegetation types in Malaya.

The collectors who have contributed to our knowledge of the limestone flora

are listed below with a brief mention of collection dates and localities.

Collectors

Allen, B.E.G.M. Dec. 15th. 1950; Gunong Keriang (Kedah), a specimen of Chirita viola

Ridl. 1958-1960; widely in Perak, visited Gunong Idong, Gunong Kanthan, Rotan Segar.

Almost 100 numbers.

Allen, E.F. May 1946; collected on ‘Kinta limestone’. April 1955-Jan. 1957; collected on

several occasions from Gua Batu and Bukit Takun in Selangor.

Alphonso, A.G. & A. Samsuri. Nov.—Dec. 1960; several numbers from Langkawi (Pulau

Bumbon Besar and Bumbon Kechil).

Batten Pooll, A.LH. A poorly labelled Dendrocalamus elegans (Ridl.) Holtt. from the Langkawi

limestone.

Bra Oct. 29th-Nov, 3rd, 1929; miscellaneous, about 80 numbers from Gunong Baling,

edah. |

Boey, H.Y. July 1970 — Sept 1971; collected from Bukit Takun on four occasions and once

from Bukit Anak Takun, Selangor. August 1971; collected from Gua Musang, Gua Batu

Boh, Batu Neng and Batu Tapah (all in Kelantan) and Gua Layang, Pahang. November

1971; visited Langkawi and collected from a number of limestone islands. About 350

numbers in all.

Burkill, ILH. 1916-1920; collected from Gua Batu, Selangor and around Ipoh, Perak. Some-

times with Haniff.

Burtt, B.L. & Woods, P.J.B.; Langkawi, at Sungei Kisap and Pulau Dayang Bunting; Perlis

at Bukit Chupeng and Bukit Bintang; Perak, around Ipoh. About 146 numbers in all.

Carr, C.E. 1928-1930; visited Kota Glanggi, Gunong Senyum, Gua Tipus and Tembeling in

Pahang. Collected mainly orchids; some ferns also. |

Chan, Y.C. July 1970; collected from around Lenggong, Perak. Several assorted numbers.

Chew, W.L. April-May 1957; collected on Bukit Hantu, Bukit Kalong, Gunong Baling and

around Dayang Bunting, Langkawi; all in Kedah. In the same period also visited Bukit

Manek and Kaki Bukit in Perlis.

Malayan Limestone Flora 169

Cockburn, P.F. June 10th 1968; some specimens from Batu Biwa in Trengganu.

Corner, E.J.H. Nov. 16th—25th, 1929 (with Henderson); collected on Bukit Lagi, Perlis.

1936-1937; visited Ipoh and Bukit Takun. Nov. 13th-22nd 1941 (sometimes with Nauen);

collected from all over Langkawi, resulting in some 200 numbers. 25th Nov. 1941 (with

Nauen); collected on Gunong Baling, Kedah.

Curtis, C. 1888-1902; visited Langkawi (Kedah), Gua Batu in Selangor, Kinta district, around

Ipoh and Sungei Siput in Perak. Most of the specimens are very poorly labelled.

Durant, C.L. Oct. 30th 1940; a specimen of Maxburretia rupicola (Ridl.) Fur. from Bukit

Takun (Selangor).

Dransfield, J. July 3lst 1968; several orchids from Bukit Anak Takun (Selangor),

Enoch, I. Sept. 25th 1954; a specimen of Impatiens mirabilis Hk.f. from Pahang.

Evans, G.B. Early 1966; Ipoh Temple limestone (Perak) and Bukit Takun (Selangor);

several ferns.

Everett, B. July 1970; collected from around Lenggong, Perak. Eighteen numbers.

Fox, W. 1899-1904; several plants from Langkawi (Kedah) and around Ipoh (Perak).

Furtado. C.X. June 4th 1937; several palms and two aroids from Gunong Baling, Kedah.

Haniff, Mohamed. 1900-1921; collected occasionally from limestone, visited Bukit Wang,

Gunong Keriang and Langkawi (Kedah), and Gunong Pondok (Perak). After 1918

usually with Nur. See also Burkill.

Hashim, Ja’afar. Collected a Wikstroemia indica (L.) C.A. Mey from Langkawi.

Henderson, M.R. 1923-1935; visited most of the limestone localities. Langkawi, usually on

both sides of Selat Panchor (straits), and around Pulau Dayang Bunting. Gua Lambok,

Gua Musang, Gua Panjang and Gua Teja (Kelantan). Bukit Cheras, Bukit Chintamani,

Bukit Serdam, Gua Tipus, Gunong Senyum and Kota Glanggi (Pahang). Gua Lanno,

Gunong Pondok, Ipoh, Lenggong and Pulai (Perak). Besih Hangat, Bukit Chupeng,

Bukit Lagi, Bukit Wang Tangga and Tebing Tinggi (Perlis). More than 700 numbers,

Holttum, R.E. Oct. 3rd 1922; Gua Batu, Selangor, several ferns. Aug. 22nd-26th 1925;

Langkawi, around Pulau Dayang Bunting and Pulau Timun. About 25 numbers.

Johnson, A. Nov. 17th 1968; several ferns from Gua Batu, Selangor.

Kadim, Tassim. Aug. 13th 1959 (with Allen, B.E.G.M.); several numbers from Gunong

Tempurong, Perak.

Kassim, Rajab. Aug. 17th 1962; several numbers from Gua Batu, Selangor.

Kelsall, H. Jan. 1891; several plants from Gua Batu, Selangor; all poorly labelled.

Keng, H. Sept. 25th 1954; at Kuala Trengganu, several ferns. Mar. 28th 1962 (with

Mrs. Keng); a specimen of Monophyllaea horsfieldii R. Br. Nov. 3rd—7th. 1968; Lang-

kawi islands (sometimes with others); some assorted specimens.

Kerr, A-F.G. June 20th 1932; Langkawi, around Kuah; several numbers.

Kiah, Hj.M.S. April 11th-May 9th 1938: collected on Gunong Baling, Kedah and Kaki

Bukit, Perlis. More than 150 numbers.

King, Sir G. See under Kunstler. .

Kunstler, H.H. (collector for Sir G. King, as King’s collector) 1880-1885; collected around

Gopeng and Larut, also visited Gunong Pondok (all in Perak). Almost 100 numbers.

Loh, H.S. July 1970; Gua Peningat, Pahang. Recorded 101 numbers. This is the first known

occasion that this outcrop (at 713 m the tallest in Malaya) has been climbed. February

1972; visited Gua Musang, Kelantan. Sixteen numbers.

Mahmud, S. 1970-1971; Bukit Anak Takun (Selangor), and Ipoh (Perak). Several numbers

See also Haniff. About 160 numbers.

170 Gardens’ Bulletin, Singapore — XXX (1977)

Ogata, K. Feb. 14th-15th, 1968; Gunong Gajah and Gunong Tempurong in Perak. Some

assorted specimens.

Phang, C.I. Sept. 15th 1960; several ferns from Bukit Anak Takun, Selangor.

Poore, M.E.D. Sept. 12th 1960 & July 16th 1961; visited Gua Batu, Selangor. Several

numbers.

Reid, J. Jan. 29th 1950; three numbers from Bukit Takun, Selangor.

Ridley, H.N. 1896-1897; visited Kuala Dipang (Perak) and Gua Batu, Selangor 1898-1920;

visited Gua Batu about five times, also Gunong Keriang, Kedah and Bukit Lagi, Perlis.

About 150 numbers.

Samat, A. 1961-1969; visited Langkawi (Kedah), Bukit Anak Takun and Bukit Takun

(Selangor). Mainly ferns; about 15 numbers.

Samsuri, A. Mar. 8th-14th, 1971 (with Mahmud); collected in Perak, around Gopeng, on

Gua Putri, Gunong Pondok, Gunong Rapat, around Ipoh, and in Tambun district. About

120 numbers.

Shimizu, T. Oct-Nov. 1967 (with Stone); visited Gua Musang (Kelantan) and Bukit Takun

(Selangor). Some ferns.

Sinclair, J. 1958-1962; visited hills around Ipoh (Perak) and Gua Batu (Selangor). About

30 numbers.

Singh, Hardial. Jan. 23rd 1966; visited Gua Batu (Selangor). Several numbers.

Smith, G. One Chirita caliginosa Cl. from Perak Cave Temple.

Smith, J.W. July 19th 1956; one Vitex siamica Will. from Gunong Baling, Kedah.

Soepadmo, E. Sept. 1968; one Carallia brachiata (Lour.) Merr. from Ipoh, Perak.

Spare, G.H. Oct. Ist 1939; Gunong Pondok, Perak. Several numbers.

Start, A. Jan. 23rd 1972; one Mucuna biplicata Teysm. et Binn. from Bukit Anak Takun,

Selangor.

Stone, B.C. 1965-1972; visited Langkawi (many of the islands), Kedah, Gua Musang

(Kelantan), Gunong Pulai (Perak), Bukit Anak Takun, Bukit Takun and Gua Batu

(Selangor). Chintamani (Pahang). About 450 numbers.

Students (University of Malaya 1967) May 1967; some numbers from Langkawi (Kedah).

Symington, C.F. 1934-1938; visited Langkawi (Kedah) and Bukit Takun. About a dozen

numbers.

Teruya, Z. Jan. 20th 1929; one Monophyllaea horsfieldii R. Br. from Gua Batu, Selangor.

Tomlinson, P.B. Sept. 1955; one Monophyllaea hirticalyx pr. from Ipoh Perak.

Turnau, E.A. Aug. Sth 1962; two numbers from Langkawi, Kedah.

UNESCO 1962 July 27th-Aug. 13th, 1962; visited the Kelantan limestone around Gua

Musang and Bertam, collected on Batu Bayan, Batu Hayan, Batu Ner, Batu Pinta, Gua

Batu Boh, Gua Musang, Gua Serai, Gua Seri and Gua Panjang. About 700 numbers.

Led by Dr. H. Keng.

Whitmore, T.C. 1966-1970; visited Langkawi (Kedah), Gua Batu Boh, Gua Musang, Kuala

Jenera, Sungei Nenggiri (Kelantan), and Bukit Anak Takun, Bukit Takun, and Gua

Batu (Selangor). About 120 numbers.

Wong, Y.K. May 13th 1962; a Maxburretia rupicola (Ridl.) Furt. from Bukit Takun,

Selangor.

Wooldridge, T.A. June 1896; a Dracaena curtisii Ridl. from Langkawi (Kedah).

Wray, L. 1894; two specimens from Perak limestone; labels mutilated, one undated.

Wyatt-Smith, J. June 11th 1960; one Salacia grandiflora Kurz. from Gua Batu, Selangor.

Wycherley, P.R. Dec. 28th 1969 (with Stone); Gua Batu, Selangor. Some assorted specimens.

__ For the purpose of this study most of the limestone areas in Malaya were

Mim and numerous hills explored and botanised. About 1550 numbers were

collected.

The plants treated include the terrestrial as well as the epiphytic members. »

Included also are species from hill bases where the soil is definitely of limestone

Origin; this is usually a rusty-red loam and very distinctive. Characteristically there

are numerous boulders strewn about. Admittedly it is often difficult to decide what

does or does not constitute the limestone vegetation. Certainly the vegetation at

hill bases (with limestone derived soil) is different from that on hill tops, but

equally it also differs from that of the surrounding lowland forest. This hill base

Malayan Limestone Flora 171

vegetation has arisen as a result of the interaction between the presence of the

hill and the surrounding forest. Accordingly I have included the species found

here in this treatment.

In the case of plants from the small outcrops recently discovered in Johore

(Rajah, 1970) great care was exercised to see that only those actually growing on

the rocks were recorded. This is because the outcrops here are very limited and

low, from boulder size protrusions to several of about 20 m tall and as broad.

Most of the cracks and crevices are filled with soil derived from the surrounding

sandstone formation. (This publication also records the first botanical exploration

on the Johore limestone.)

All specimens from ‘near the base of hill’, or from ‘hill bases’ as recorded

on the labels of herbarium sheets are excluded unless the labels also specify that

the particular plant was from limestone rock or soil.

In order to obtain as complete a record as possible of the limestone flora,

every sheet in the herbaria at the Singapore Botanic Gardens, the University of

Singapore and the University of Malaya was scrutinized. A start was also made

at the herbarium of the Forest Research Institute but as the search was not

showing success it was stopped. Practically all the old and most of the new lime-

stone records there (at Kepong) are duplicated in the Singapore Botanic Gardens

herbarium. Subsequently the examination of specimens at Kepong was restricted

to the known limestone genera and to recent (post 1960) records from limestone

districts (with the help of collector’s field notes).

Many labels are unclear and to decide whether a specimen was recorded from

limestone or not is not always straightforward. Sometimes the town, village or

district bears the same name as the limestone hill (or vice versa), e.g. Baling,

Gua Batu and Gua Musang. Specimens in such cases were excluded unless the

record states explicitly that it was from the hill.

By searching the herbaria and literature I have managed, I think, to include

all the species (possibly with few exceptions) so far recorded from the Malayan

limestone.

Purpose and Scope

This work attempts to present a comprehensive account of all pteridophytes

(true ferns only) and spermatophytes ever found growing on limestone hills and

sites in the Malayan Peninsula. Included also are the non-indigenous species

(introduced, inadvertently or otherwise) which have become naturalized and now

form part of the wild-growing population.

Sect. 1 discusses the geology, origin and distribution of the limestone hills, the

vegetation, phytogeography and size of the flora, and affinities of the floristic

components to limestone.

Sect. 2 the flora proper, starts with the Pteridophytes (true ferns only), going

on to the Gymnosperms and ending with the Angiosperms (Dictoyledons and

Monocotyledons). Under these four main groups, the families are arranged in

alphabetical order and thereunder the genera and species. Dichotomous keys

leading to the identification of almost all the species are included. For the

exceptionally large groups, the Pteridophytes, Euphorbiaceae, Orchidaceae and

Rubiaceae, an introductory key with leads to the main key is provided.

172 Gardens’ Bulletin, Singapore — XXX (1977)

Every species is provided with its approved name and the original place of

publication. Synonyms, if any, especially those published in major works relevant

to the local flora, are cited. References are also made to publications on the local

flora, and reference to Henderson (1939) is cited for every species included in

his work.

The names of characteristic species (those that come under my ‘affinity’

grouping I and IJ in ch. 5) are printed in bold face type. In addition, brief notes

on the distribution, and frequency on the Malayan limestone are often included for

these species.

All literature cited in the discussion are listed in full in the bibliography at

the end of this work.

Specimens are cited only in special cases, such as rarity, or doubtful locality,

or in relation to certain taxonomic problems. However, full lists of specimens are

to be found in the dissertation paper, and this may be consulted in the Library

of the University of Malaya or in the University Herbarium, Kuala Lumpur.

Lastly Bryophyte collections were made during this study but are not included

here because of the difficulty in obtaining identifications. A further study of this

group would be desirable.

2. GEOLOGY, ORIGIN AND DISTRIBUTION OF THE LIMESTONE

HILLS.

Geology

Most of the limestone hills in Malaya rise from flat or undulating plains. They

form conspicuous and often spectacular hills up to 2342 ft. (713 m.), (Gua

Peningat, Pahang), often with sheer, vertical or overhanging cliffs.

They range in age from Ordovician to Triassic. The estimation of age has

been a difficult problem since are recrystallised, unfossiliferous (Hutchison, 1968),

and as all studied samples are recrystallised, thermoluminescence cannot be used

to differentiate limestone stratigraphically in Malaya (Hutchison, 1968). Up to

as late as the 1950’s anything calcareous in Malaya has been assumed to be

carboniferous and was ascribed to the calcareous formation. Since then many

studies have been made, providing a more accurate picture of the age of our

limestone, e.g. Paton (1961), Ingham and Bradford (1960), Gobbett (1965a),

Hutchison (1968) and Suntharalingam (1968), but the ages of many still remain

indicated by only a very rough estimation.

The Kedah/Perlis(including Langkawi) limestone areas are mainly Ordovi-

cian-Silurian. The Selangor limestone appears to be confined to the Silurian

(Gobbett, 1964), and the recently discovered Johore limestone is Permian (Rajah,

1970). Most of the others are Permian (Paton, 1961), but further studies will

probably show a variation in age from the Silurian to Permian (as is the Selangor

limestone Paton included under Permian). However no hill has been found to be

Triassic which so far is limited to small lenticular beds in Pahang (Paton, 1961).

It is generally thought that the older limestone gives rise to more or less

continuous ranges of hills without marked vertical cliffs whereas on the other hand

the younger limestone produces more isolated hills with high, vertical or even

overhanging cliffs (Paton, 1961). This weathered-look is shown by the Kedah/-

Perlis (including Langkawi) limestone of Ordovician-Silurian age. This genera-

lisation was. made before the Selangor limestone was known to be Silurian,

and now no longer holds. This is because the latter shows marked cliff development

(except for Bukit Anak Takun). Thus it would seem that the age of the actual

limestone rock itself has little bearing on the development of the hills, and there is

no difference in form between hills from Permian and Ordovician-Silurian rock.

Malayan Limestone Flora 173

Our limestones are generally very pure. Most are white, pale grey or slightly

yellowish; some are dark grey to almost black because of carbonaceous or

argillaceous impurity. Rarely it is red because of hematite inclusion or by iron

staining. Hutchison (1968), who analysed 182 very random samples chemically

(which therefore should give a rough indication of the true nature of the Malayan

limestone) recorded that most of them have about 2.5% insoluble residue with an

arithmetic mean of about 4.4% insoluble residue. He further notes that 81% of

his samples are limstone (CaCO,) and 19% dolomite (CaMg (COs) :).

Origin

Originally sedimentary, the Malayan limestone is, as mentioned earlier,

recrystallised. It is thus in fact marble. This happened when peninsular Malaya

underwent an orogeny (an episode of mountain-building during which the rocks,

including the Malaysian limestone were compressed, folded and heated) in the

Triassic period (about 200 million years ago). Towards the end of the Triassic

orogeny the rocks were uplifted into hills and mountains. The botanical stage was

set, since then most of the Malayan limestone was never submerged again, and

they were ready to receive their first plant. However it is not certain how long it

took before the limestone topography as we know came into being.

Various theories have been put forward to explain the existence of the lime-

stone in the form of spectacular karst-tower hills. These are reviewed by Paton

(1964), and include block faulting, sub-aerial erosion, folding, marine erosion and

plastic flow. The most plausible explanation seems to be simple sub-aerial denuda-

tion modified in certain cases by marine erosion.

The theory of sub-aerial erosion takes into account the high solubility of

calcium carbonate compared to the majority of the silicates, of which most of all

other rocks are composed. This is an important point, for any theory seeking to

explain why the limestone hills are in the form they are must also explain why

formations of other rocks are not existing in the form the limestone hills are.

Weathering of silicate rocks produces more or less rounded topography; on lime-

stone deep channels are often cut.

Streams and swamps at the bases of limestone (assuming that they were once

well-jointed and continuous) are highly effective in eroding it. The pH of swamp

and stream water in low-lying areas is commonly as low as 3.5 (Paton, 1964).

This acidic water gradually erodes the rock, forming channels; floors and walls

are enlarged, and from above, downward erosion cuts deep channels that

eventually reach the base. The underground channels continue to expand laterally,

vote resulting in the collapse of the roofs, producing steep-sided and isolated

The initial stages of erosion by water are evident in the form of grooves or

nips, characteristic of many coastal limestone areas in Langkawi and at the bases

of hills on the mainland. Some grooves on hills in the Kinta and Kedah/ Perlis

area are marine in origin, formed in the post-glacial periods (post Pleistocene) when

the sea level, at least 50 ft. higher than now (Scrivenor, 1949; Walker, 1956),

flooded the plains.

Distribution

Until recently, all the known limestone outcrops were from north of Kuala

Lumpur. However in 1965-66, officials of the Geological Survey of Malaysia

discovered limestone outcrops in Johore while carrying out geological mapping of

that area (Rajah, 1970). The outcrops are in an area 500 by 1200 m on the south-

eastern slope of Gunong Sumalayang at an elevation of 250 to 400 m. There are

about 30 outcrops here varying in size from boulders 1-2 m across to the largest

about 20 m tall and as wide.

174 Gardens’ Bulletin, Singapore — XXX (1977)

Elsewhere in Malaya the limestone hills are in their most spectacular and

prolific development in south-west Kelantan and north-west Pahang, in the Kinta

valley from Kampar northwards to Ipoh, Tambun and Tanjong Rambutan, and

in north-west Perlis.

Hills are also found around Alor Star, Kedah, elsewhere in Pahang, near

Kuala Lumpur, Selangor and rarely in Trengganu. They are very prominent in

the Langkawi group of islands off the extreme north-west coast of Malaya. Not

all the islands are limestone, and some are limestone only in part, but generally

limestone is abundant, especially in the eastern and southern aspects of Langkawi.

This is the only coastal limestone in Malaya.

The distribution of the Malayan limestone hills is shown in Maps 1, 2 & 3.

All the hills botanised are included in the maps except some of the smaller hills

which are not named even in regional 1 inch to 1 mile maps. It has also not been

possible to include all existing outcrops, but all the limestone localities are

represented and most of the popuarly and less popularly known have been included.

3. VEGETATION

The limestone vegetation forms a characteristic and easily recognised forest

type in Malaya. It is a distinct Edaphic Climax Formation (Symington, 1943). In

fact, Corner (1960) considered the limestone forest as one of the three constituent

forest types (the other two being the lowland and the mountain forest).

The vegetation supports an extremely diverse and rich flora which is a

reflection of the diversity of the habitat. Henderson (1939) classified the Malayan

limestone hills into three broad groups, viz.:

1. Very dry hills with little soil and much exposed rock. Trees usually

rather stunted; mosses and herbaceous plants scarce.

2. Wet hills often with gullies filled with rich soil. Trees fair-sized and not

stunted; mosses and herbaceous plants present.

3. Soil-rich hills, well covered with soil (usually a stiff red clay), little rock

exposed. Trees tall; mosses and herbaceous plants few.

This is in effect, as Henderson further discussed, dividing the limestone hills

into two general categories, ‘dry hills’ (1) and ‘wet hills’ (2 and 3). This classifi-

cation however does not do justice in summing up the diverse limestone habitat.

‘Dry hills’ often have sheltered gullies with soil supporting fair-sized trees, mosses

and herbaceous plants. Similarly ‘wet hills’ often have exposed rocky ridges with

stunted vegetation.

Anderson (1965) subdivides the Sarawak limestone habitat into eight groups,

viz.: 1. Alluvial soils at the base of hills. 2. Base of hills and ravines between hills.

3. Slopes of hills 4. Cliffs. 5. Scree slopes on hills. 6. Summits of hills at low

altitudes. 7. Submontane limestone. 8. Secondary vegetation on hills.

In Malaya there are no submontane limestone hills and this classification

does not seem to effectively describe the Malayan limestone habitat. Thus

another classification is proposed. This is, like the other schemes, only provisional

and there is a great deal of overlap between the subdivisions. This subdivision

of the habitat takes into account the geology and topography of the hill, the

physiognomy of the vegetation and the floristic composition. Succession is also

considered and disturbed or secondary vegetation is therefore made a subdivision.

Malayan Limestone Flora 175

PULAU

TIOMAN

cea Ss (

JOHORE

GUNONG®

SUMALAYANG

MAP 1. MALAYA IN GENERAL

176

6° 20S

Gardens’ Bulletin, Singapore — XXX (1977)

199° 50°E

PULAU LANGKAWI

199° 50'E

MAP 2. LANGKAWI, SHOWING THE MAJOR LIMESTONE LOCALITIES

Key to the numbers (in a clockwise direction starting from the north).

1. Pulau Dangli 11.

2. Pulau Gasing

3. Pulau Pasir 12.

4. Tanjong Batu Kulat 13.

5. Tanjong Ru 14.

6. Gua Cherita 15.

7. Pulau Langgun 16.

(with lake) 17.

8. Pulau Tanjong Dendang 18.

9. Tanjong Dagu 19.

10. Tanjong Dagu Forest 20.

Reserve 21.

Selat Panchor Forest

Reserve

Selat Panchor (straits)

Pulau Timun

Pulau Lading

Pulau Chupak

Pulau Bumbon Besar

Pulau Bumbon Kechil

Telok Sambar

Tanjong Tirai

Pulau Dayang Bunting

Pulau Jerkom Kechil

22.

Pulau Jerkom Besar

Pulau Singa Kechil

Pulau Singa Besar

Tasek Dayang Bunting

(lake)

Pulau Beras Basah

Pulau Rebak Besar

Tanjong Belua

Tanjong Chinchin

Tanjong Tok Manap

Malayan Limestone Flora 177

The subdivisions are:

Base of hills

Talus slopes

Hill slopes to about 60° steepness

Gullies and valleys

Cliffs and near-vertical slopes

Summits with considerable soil cover

Summits with none or very little soil cover

Coastal limestone

Disturbed areas

CHONAARWN

Although the nature of the habitat necessitates a classification taking into

account all the characteristics present to separate the subdivisions effectively, this

classification is more physiognomic than floristic in kind. The physiognomy of the

vegetation is in turn dependent on the topography of the hill, the abundance or

scarcity of soil, moisture and shelter. The floristic composition however varies a

great deal from hill to hill and although it would be possible to subdivide a hill

floristically, this pattern of floristic distribution would not hold for other hills.

Species characterising a dry rocky summit like Bukit Takun (Buxus malayana and

Wikstroemia androsaemifolia) are not found on a similarly dry and rocky part

of, for example, Batu Neng. Sycopsis dunnii is common on Bukit Serdam summit

(with thin soil) but on Gunong Rapat summit (also with thin soil) a common

plant is Murraya paniculata, and Sycopsis dunnii is absent, and the converse

applies.

Floristically also, the limestone north of Alor Star, including those in

Langkawi, mainland Kedah and Perlis differs from that further south. They form

part of the Thai flora which begins north of Alor Star, in Kedah (Ridley, 1911)

or north of the mouth of Kedah river which is near Alor Star (Kloss, 1922). (This

difference in the floristic composition is due mainly to the regular dry season from

December to February which is confined to the extreme north of Malaya).

However in this classification, the Langkawi limestone is treated in the same

manner as the rest and not as a distinct floristic entity. This is in order to provide

a coherent picture of the Malayan limestone in which floristic composition is one

of the criteria used and not the deciding factor.

The examples of plants cited in the following discussion are representative of

the subdivisions of limestone habitats in which they are found. However a con-

siderable number of such examples do not have rigid habitat preferences and may

also be found in the other subdivisions. Though they are representative of the

various subdivisions they are not necessarily found on all hills with similar

habitats; in fact many species form very local populations on one or several hills.

1. Base of hills

The bases of hills are usually covered by a mixture of alluvial soil not derived

from the limestone but from the surrounding geological formation and also soil

derived from weathering of the hills. The vegetation here varies with the locality

of the hill and probably with the nature of the derived alluvial soil (from the

surrounding geological formation). Protected valleys (wangs) may occur; these

are surrounded, often entirely, by abrupt limestone walls. The floor is usually the

typical Terra Rossa soil. Such areas are often cultivated (as in the hills around

Ipoh).

The vegetation is generally tall and closed, with the crowns of the plants

touching or overlapping. Typical species include Villebrunea sylvatica, Gmelina

asiatica, G. villosa, Dendrocnide stimulans, Diospyros cauliflora, Mallotus

philippensis, Melanolepis multiglandulosa, Orophea hirsuta and Pisonia umbelli-

fera. Some climbers are also found e.g.; Jodes cirrhosa and Mucuna biplicata.

178 Gardens’ Bulletin, Singapore — XXX (1977)

Sometimes where water drips down from steep rocky slopes or from over-

hanging cliffs, a distinct herb community develops. Species include Alocasia lowii,

Chirita caliginosa (which is also found on dry situations) Epithema saxatile and

Monophyllaea horsfieldii. Bryophytes are usually luxuriant.

2. Talus slopes

At and near the base of many hills are accumulations of boulders, organic

matter and general debris from the hill. Rocks from this scree have evidently

fallen from the hill. Often the bases of hills surrounding amphitheatres or wangs

have talus slopes at their bases. These are probably remnants of the roofs of large

caverns that have suffered a cave-in. The vegetation here is tall and closed,

physiognomically not unlike that on cliff bases (Gp. 1.). The composition is

however somewhat different; and perhaps the distinguishing character is the herb

and shrub vegetation characteristing such a shady, rocky terrain. The trees and

shrubs include Atalantia roxburghiana (peculiar to Bukit Takun), Cleidion

javanicum, Morinda elliptica, Streblus ilicifolius and Trivalvaria macrophylla. The

herbs include Heterogonium pinnatum, H., alderwereltii, Impatiens spp. (1. mirabilis

dominates the scenery on Pulau Langgun, Langkawi, north of the lake), Pilea

and Procris.

3. Hill slopes to about 60° steep

These more gentle slopes are nearly always covered with soil and support a

dense, close vegetation 5-10 m or more tall. The soil may be thin or to about

1 m deep at places, and boulders are usually present emerging from the soil.

Ground vegetation is usually spares and is often associated with the boulder out-

crops. These include Antrophyum parvulum (on rocks), Ophiorrhiza, Piper, and

in the more sheltered niches Gymnostachyum decurrens and Scutellaria discolor.

The trees and shrubs include Cladogynus orientalis, Croton cascarilloides, Mallotus

dispar, Microdesmis casearifolia, Orophea cuneiformis, Polyalthia brunneifolia,

Saraca declinata, S. thaipingensis and Rinorea bengalensis. On Gua Kechil, Raub,

the rare tree Diospyros adenophora is fairly common.

4. Gullies and valleys

These are often the most sheltered spots on hills. This group includes

pot-holes, depressions, gullies, valleys, ravines and other concavities which may

be narrow and small or broad and extensive, extending between two peaks on the

same hill. The ground is usually deep or shallow soil with a covering of litter and

other organic debris. The vegetation is fairly dense, 5-10 m tall with occasional

emergents, and physiognomically not unlike that on soil-covered slopes and talus

slopes. The floristic composition varies somewhat and its location and topography

is usually distinctive. (Sometimes however gullies and valleys may emerge into

hill slopes.) Gullies may even be present on the so-called ‘dry hills’ (the most

outstanding example is Bukit Takun), harbouring mosses and delicate herbaceous

plants like Corybas mucronatus and Goodyera hispida. The ground vegetation is

usually quite sparse, no doubt because of the closed canopy. Bryophytes are

common and herbs like Epithema saxatile and Monophyllaea are usually associated —

with boulder outcrops.

The trees and shrubs include A grostistachys gaudichaudii, Canthium didymum,

Fagraea curtisti, Randia densiflora, Sauropus suberosus and Sterculia rubiginosa.

Climbers like Connarus and Lasiobema are found and the palms present include

Aaa westerhoutii (also from talus slopes), A. hookeriana and Iguanura poly-

mor pha.

Malayan Limestone Flora 179

_ EN

MILES

———— International boundary

———— Stote boundary

seer Railway

ot ere cies aby

“i .

Bertam

Batu Pintg

- Kuala @ 8

J Bati

Gunong ‘s NGua Panjang

Pondck

{ ae i

7 ao Pes

be bel RAMBUTAN -—— ;

4 eT, N>\ v j rere ue Layers Ve”

.Gue Peningat

coh

yap

exRnt? x

-KUALA

LUMPUR

100° 101° 102° 103°

MAP 3. MALAYA (EXCLUDING LANGKAWI!) SHOWING THE MAJOR LIMESTONE OUTCROPS

Key to the numbers indicating position of outcrops.

1. Bukit Besih Hangat 5. IPOH — ‘including Perak Cave Temple

Bukit Wang Tangga 6. GOPENG — including Gunong Lanno

Tebing Tinggi Gunong Rapat

2. KODIANG — including Batu Hantu 7. Kuala Dipang

Batu Kalong Sungei Siput

3. LENGGONG — including Batu Tukang g GUA MUSANG — including Batu Machang

Gua Badak Batu Papan

Gua Pipit Batu Tapah

Gua Putri Gua Serai

4. TAMBUN — including Gunong Ginting

Rotan Segar

180 Gardens’ Bulletin, Singapore — XXX (1977)

5. Cliffs and near-vertical slopes

Vertical cliffs, overhanging cliffs and near vertical slopes present a unique

habitat and support a distinct vegetation which depends on the degree of exposure

of the cliffs, presence or absence of soil pockets and availability of moisture. This

is a very easily recognised subdivision topographically and the vegetation varies

from very sparse and open (or sometimes absent) to a fairly dense herb and scrub

cover. This denser vegetation cover is found along ledges where there is accumu-

lated soil and debris and in other soil pockets. Where there is seepage or water

dripping down a characteristic vegetation develops along this line of moisture.

This may be on overhangs or clefts on cliff faces. If on clefts this vegetation

decends to the base of the cliff and is very similar to that described under sub-

division 1 (where water is available). This includes Chirita, Epithema saxatile,

Monophyllaea and the fern Adiantum malesianum (which is also from dry

localities) and bryophytes.

Cliff faces with no cracks or crevices do not support any macro-vegetation,

but those with cracks and crevices which invariably collect soil and other debris

support a distinctive flora. The species include the very characteristic Boea,

Paraboea and Chirita. Boea spp. especially are often very prominent high up on

cliff faces.

Near vertical slopes and cliffs with ledges and other soil pockets often support

a herbaceous or scrubby vegetation. The species include Canscora pentanthera,

Cymbopogon calcicola, Callicarpa angustifolia, Cheilanthes farinosa (only on

Bukit Chintamani), Jasminum, Pyrrosia stigmosa and Zizyphus oenoplia.

6. Summits with considerable soil cover.

Most hill summits are covered with soil (at least in part). Some have

considerable to extensive rock outcrops with sparse vegetation (these rocky spots

are included under subdivision 7.) In places scattered boulders and pot-holes are

common; generally the topography is uneven to very rugged. Ridges with some

soil supporting a closed vegetation are included here. The vegetation is usually

scrubby, 2-7 m tall and the canopy is closed. In places the plants are spindly.

Generally these shrubs and trees are fairly well spaced, and though the canopy

is closed there usually is a rich ground flora. Occasional emergent trees are present,

eg; Mangifera sp. on Gunong Pondok, Madhuca ridleyi on Bukit Serdam and

Garcinia murdochii on Gua Kechil.

The trees (they are usually somewhat stunted, though not extremely so) and

shrubs include Cleistanthus gracilis, Decaspermum fruticosum, Dehaasia curtisii,

Eriobotrya bengalensis, Erythroxylum cuneatum, Eugenia pendens, Garcinia

minutiflora, Mallotus brevipetiolatus, Mallotus miquelianus, Murraya paniculata,

Neolitsea zeylanica and Sycopsis dunnii (especially on Bukit Serdam). The pandan,

Pandanus irregularis is common and conspicuous on ridges on Batu Neng.

Climbers and scramblers are usually scarce and include Maytenus curtisii and

Lygodium polystachyum.

The ground vegetation is fairly rich in bryophytes especially in the more

sheltered spots on rocks and at the basal parts of tree trunks. The herbs include

the lycopod, Lycopodium squarrosum; ferns are common and in places from the

dominant ground cover; they include Asplenium adiantoides, A. macrophyllum,

Loxogramme avenia, Photinopteris speciosa, Phymatodes scolopendria and

Pyrresia stigmosa. Ground orchids are equally common and include Agrostophyl-

lum bicuspidatum (which may also be a low epiphyte), Coelogyne asperata

(which on parts of Batu Neng and Batu Tapah forms almost pure mats),

Dendrobium luxurians, Eria rigida and Renanthera histrionica. Epiphytes are

usually not conspicuous though occasionally they are found; these include orchids,

e.g. Appendicula torta and Bulbophyllum apodum and ferns, e.g. Pwrosia.

Malayan Limestone Flora 181

7. Summits with no or very little soil cover

Most hills have some exposed rocky parts, while a very few have extensive

rocky summits. Soil is scanty and found in little hollows, cracks and clefts in the

rock but is never completely absent. Even extensively rocky summits have gullies

and valleys where soil and litter are accumulated and the vegetation supported is

different (included under subdivision 4.)

The vegetation on these rocky summits is sparse and open but fair-sized

trees rooted in clefts and hollows are present. Most are, however, stunted and

gnarled though occasionally large ones occur. An example are the figs, the roots

of which ramify all over the rocks, into every crack and crevice and often

extending for many meters over cliff faces in an intermeshed network.

Bukit Takun in Selangor is the classic example of a hill with an extensive

rocky summit, but other examples occur, e.g. in Langkawi. The plants on these

hills include some herbs usually from soil pockets and from amongst the surface

roots of trees where some debris have accumulated. These include Adenia nico-

barica, Amorphophallus spp., Arisaema fimbriatum, A. roxburghii, Boea spp.,

Chirita spp. and Dichanthium mucronulatum. It is interesting to note that of these,

Adenia nicobarica, Amorphophallus spp. and Arisaema spp. have fleshy tubers

which often grow into cracks and crevices. The possession of tubers could be an

advantage under such dry and exposed conditions. Climbers present include

Dischidia hirsuta, Morinda umbellata and Secamone micrantha, and the pandan,

Pandanus alticola (commonly an epiphyte outside the limestone field) which is

often present growing from rocks (it is abundant on Bukit Takun).

The trees and shrubs present include Buxus malayana, Diospyros ferrea

(commonly on hills around Gua Musang), Ficus calcicola, Glycosmis calcicola,

Memecylon laevigatum, Pistacia malayana, Planchonella obovata, Podocarpus

polystachyus, Tarenna curtisii, Terminalia triptera (only Langkawi), Vaccinium

littoreum (only on Bukit Takun and Anak Takun), and Vitex siamica.

8. Coastal limestone

In Malaya this only occurs in Langkawi where part of the coast of the main

island and many of the smaller islands are entirely limestone. Hills and islets

which descend gradually to the sea are often fringed on the seaward edge by

narrow sandy beaches (non-limestone, and derived from other geological forma-

tions as weathered limestone produce hardly any ‘sand’) on which develops a

typical strand vegetation. It is these low limestone areas that are mostly influenced

by the sea and are invaded by some species from the strand vegetation. Hills with

tall abrupt cliffs do not show any littoral influence. However, it must be remember-

ed that many species from the Langkawi limestone are not found on limestone

further south on the mainland, but this is due to the northerly position of

Langkawi principally, and not due to littoral influence (which is coincidental and

at the most contributary). These species include Bombax anceps, Cycas siamensis,

Colona merguensis, Euphorbia antiquorum, Grewia viminea, Hopea ferrea,

Pentacme siamensis, Sterculia lancaviensis and Vatica cinerea.

The coastal limestone therefore is restricted to that part of the limestone in

close proximity to the sea, and this is usually near sea-level. The seaward edge is

fringed with sand which supports a typical Indo-Malaysian littoral vegetation

(sensu Schimper, 1903). The vegetation on this limestone is sparse to dense, short

or tall, depending on the availability of soil and the topography; but littoral

species are present. The species include some herbaceous members like Davallia

solida, Humata pectinata, Eurycles sylvestris and Tacca leontopetaloides. Trees

and shrubs include Barringtonia asiatica, Caesalpinia crista, Guettarda speciosa,

182 Gardens’ Bulletin, Singapore — XXX (1977)

Heritiera littoralis, Pandanus odoratissimus, Peltophorum pterocarpum, Thespesia

populnea and Xylocarpus granatum. Together with the species mentioned above

this makes up a distinctive coastal limestone vegetation (probably distinctive even

if there were other coastal limestone areas further south).

9. Disturbed areas

Disturbed areas usually occur on the summits, lower slopes and bases of hills.

These disturbances could have been brought about by a number of causes, viz:

(i) Fire, occurring naturally or artificially, e.g. Bukit Takun in August 1970

and Gua Musang in mid-1969.

(ii) Mining and quarrying activities. This is seen on a large number of hills

in all the major limestone districts.

(iii) Agriculture, affecting only hill bases, e.g. some hills around Ipoh and

in limited spots on many other hills.

(iv) Erection of communication transmitters and trignometrical stations, e.g.

on Gunong Lanno, G. Layang-Layang and G. Tasek, also on other hills.

(v) Frequent trampling by visitors, e.g. on Bukit Takun and Gua Musang.

Whatever the cause, disturbances result in the partial or complete destruction

of the original vegetation. It is interesting to note that Cratoxylum maingayi,

Podocarpus polystachyus and Radermachera lobbii survived the 1969 fire on

Gua Musang that destroyed all the other vegetation affected. Cratoxylum and

Podocarpus probably survived because they were the largest trees there and con-

sequently had thick bark. Several plants of Radermachera (they were slender and

probably not more than 4 m tall) were burnt almost to the ground. By August

1971 the sucker-shoots sent out were already 2 m tall and flowering.

Though the complete development from sere to high forest has never been

observed, a composite and representative picture (at least of the initial stages)

can be formed by observation of a number of hills. One of the earliest colonisers

is the moss Bryum coronatum Schwaegr. which was abundant on Gua Musang

in August 1970 and also in August 1971 on burnt earth and on rocks. This moss

is followed (or accompanied) by a large number of ubiquitous weeds, truly

opportunistic and very adaptable and probably completely indifferent to the lime-

stone influence. The species include ChArysopogon aciculatus, Colocasia gigantea

(prominent on Gua Musang), Fleusine indica, Erechtites valerianifolia, Eupato-

rium odoratum, Macaranga tanarius, Muntingia calabura (prominent on _ the

quarried slopes of Bukit Chintamani), Nephrolepis biserrata, Neyraudia reynau-

diana (conspicuously gracing the summit of Gua Musang in 1971), Paspalum

conjugatum, Pityrogramma calomelanos, Pteridium caudatum var. yarrabense,

Pteris vittata, Rhynchelytrum repens and Solanum ferox. Limestone elements

present at this early stage include Boea spp., Chirita spp. and Schizaea inopinata.

Trees like Macaranga tanarius and Trema spp. are probably quite persistent,

though the latter stages of this succession have not been observed. Presumably

seedlings of other limestone elements get established and eventually from the

dominant vegetation once again, shading out the colonisers.

4. PHYTOGEOGRAPHY AND SIZE OF THE FLORA

The total number of species recorded for the limestone flora is 1216 distributed

in 582 genera and 124 families (This includes the true ferns, Gymnosperms and

Angiosperms). The introduced elements (as weeds and escapes from cultivation)

number 39 species or 3.2% ‘of the total flora.

Malayan Limestone Flora 183

The breakdown of the flora is as follows:

Dubious

Genera Species

P Records

PTERIDOPHYTES

Adiantaceae 8 16 eee

Dennstaedtiaceae 17 45 =

Grammitidaceae l - ees

Hymenophyllaceae l 4 alt

Polypodiaceae ll 9% ary

Schizaeaceae 2 3 =

Thelypteridaceae 3 6 ra!

TOTAL

GY MNOSPERMS

Araucariaceae l

Cycadaceae l

Gnetaceae l

Podocarpaceae l

TOTAL

HS. aD: me

Il pe

ANGIOSPERMS

DICOTYLEDONS

Acanthaceae 16 fs

Amaranthaceae 2 2 l

Anacardiaceae 8

Annonaceae 19

Apocynaceae 10

Aquifoliaceae

Araliaceae

Aristolochiaceae

Asclepiadaceae

Balanophoraceae

Balsaminaceae

Begoniaceae

Bignoniaceae

Bombacaceae

Boraginaceae

Burseraceae

Buxaceae

Caprifoliaceae

Capparidaceae

Cardiopteridaceae

Celastraceae

Chloranthaceae

Combretaceae

Compositae

Connaraceae

Convolvulaceae

Cucurbitaceae

Datiscaceae

Dilleniaceae

Dipterocarpaceae

Ebenaceae

Elaeocarpaceae

— WwW

“IO ©

lon

—

ES eal aati AR ik ee ft, el Seals Seen ha A ties a ee AEN |. a oe

_—_— —

184 Gardens’ Bulletin, Singapore — XXX (1977)

Dicotyledons (cont.)

Ericaceae

Erythroxylaceae

Flacourtiaceac

Fagaceae

Eacourtiaceae

Gentianaceae

Gesneriaceae

Guttiferae

Hamamelidaceae

Hernandiaceae

Hypericaceae

Icacinaceae

Labiatae

Lauraceae

Lecythidaceae

Leguminosae

Lentibulariaceae

Loganiaceae

Loranthaceae

Lythraceae

Malvaceae

Melastomaceae

Meliaceae

Menispermaceae

Monimiaceae

Moraceae

Myristicaceae

Myrsihaceae

Myrtaceae

Nepenthaceae

Nyctaginaceae

Ochnaceae

Oleaceae

Onagraceae

Oxalidaceae

Passifloraceae

Piperaceae

Pittosporaceae

Polygalaceae

Polygonaceae

Primulaceae

Ranunculaceae

Rhamnaceae

Rhizophoraceae

Rosaceae

Rubiaceae

Rutaceae

Santalaceae

Sapindaceae

Sapotaceae

Scrophulariaceae

Solanaceae

Staphyleaceae

Sterculiaceae

Thymelaeaceae

Tiliaceae

Ulmaceae

Urticaceae

Verbenaceae

Violaceae

Vitaceae

TOTAL

Om nOON==NhemewUe Tb

—

NS)

— i We Om Oe Ao oo Rh Noe

—

SNR RR KK WK WN KH WK HNK RPRANWHK DAA ot —=ARwWe

—_

— ON

We AR OO RANK UNS — NRK RNS =

—

CO Ar wmONR— OK NWA RL — CO

— —

OhANWIW

805 11

Malayan Limestone Flora 185

MONOCOTYLEDONS

Agavaceae

Alismataceae

Amaryllidaceae

Araceae

Burmanniaceae

Commelinaceae

Cyperaceae

Dioscoreaceae

Flagellariaceae

Gramineae

Hypoxidaceae

Liliaceae

Lowiaceae

Marantaceae

Musaceae

Orchidaceae

Palmae

Pandanaceae

Stemonaceae

Taccaceae

Triuridaceae

Zingiberaceae

TOTAL 22

tr —_

Ne oe br et eer eer Sip il =

— (NN

—_—

GRAND TOTAL 126

les}

“

The total number of families presented here include Aristolochiaceae and

Alismataceae which have only been dubiously recorded. Thus the number of

families actually recorded from the limestone is 124. The total number of plants

recorded from limestone is 1216 and excluding the ferns this give a total of 1119

species of spermatophytes. Keng (1970) estimated that between 8000-8500 species

of spermatophytes occur in the Malay Peninsula. Thus the total number of species

of spermatophytes on the limestone is between 13.0-13.9% of the total number

of species found in the Peninsula.

Henderson (1939) compared the number of species on limestone and the area

of limestone with the total number of species in Malaya and the total area of

Malaya. Based on Scrivenor, in Burkill (1935), Henderson concluded that the

total area of limestone, which is about 260 sq. km. (100 sq. miles), is 0.2-3% of

the total area of the Peninsula. The total number of species from limestone

recognised then was 745 which was about 8-9% of the total number of flowering

plants (then known) in the Peninsula. These figures seem to indicate a much richer

flora per unit area on the limestone than on the other formations. The present

figure of 13.0-13.9% would suggest an even richer flora.

This method of comparison assumes that there is a direct relationship between

the number of species and the area of land, and that the number of species increases

linearly with the increase in land area, This seems incorrect. For instance, Gua

Batu supports about 170 species of plants. This is 14.5% of the total number

recorded from limestone. It has an area of only about 2.59 sq. km. (1 sq. mile),

which is 1% of the total area of limestone in Malaya. It would appear that Gua

Batu were floristically 14.5 times richer than the other limestone hills.

To obtain a correct picture, only the ‘minimal area’ of a community should

be used in comparison with the ‘minimal area’ of another community. This

concept of ‘minimal area’ (Braun-Blanquet, 1932; Poore, 1955) has been defined

186 Gardens’ Bulletin, Singapore — XXX (1977)

as the smallest area which can contain an adequate representation of an associa-

tion. However, the ‘minimal area’ for both the limestone and the lowland forest

in Malaya have not been worked out. One alternative, therefore, is to select an

area with the same or about the same area as the limestone and whose flora has

been worked out. This flora could be used to compare the limestone flora with.

The flora of Penang island which is about 285 sq. km. (110 sq. mi.) in area

has been worked out by Curtis (1894). His list of flowering plants total 1805

species. Similarly the flora of Singapore (Ridley, 1900) which is about 582 sq. km.

(225 sq. mi.) included 1952 species. The two floras give an idea of the richness

of the Malayan flora but as far as a comparison with the limestone flora go, they

do not give an accurate picture. The limestone flora is from an edaphic climax

formation whereas areas like Singapore and Penang support (originally) a climatic

climax formation which includes a number of other climaxes, e.g.; mangrove

swamp forest, beach forest, hill dipterocarp forest (in Penang), Adinandra forest

(in Singapore), riparian fringes and seasonal swamp forest (at least in Singapore)

apart from the lowland dipterocarp forest.

The other alternative therefore is to compare a “‘minimal area”’ of the lime-

stone flora with the “‘minimal area” of the flora of another edaphic climax forma-

tion. This comparison will probably give the most accurate picture of the relative

size and richness of the limestone flora. Unfortunately no flora of any other

edaphic climax in Malaya has been worked out that could be conviniently used.

However, Anderson (1963) published an account of the past swamp forest

of Sarawak and Brunei, which under the present circumstances should be the most

relevant to compare the limestone flora with. The coastal and deltaic peat swamps

cover a total of 15644 sq. km. (6040 sq. mi.) of Sarawak and Brunei. The total

number of flowering plants recorded in Anderson’s list is 393 species. Thus in

comparison even the vastness of the Sarawak and Brunei peat swamps have not

support a flora as rich as that on the Malayan limestone. The limestone vegetation

of Sarawak (surface area much less than the peat swamp) is also exceptionally

rich in plant species. A preliminary report on the Sarawak limestone (Anderson,

1965) records over 600 species, and this number is bound to increase as more

limestone areas are explored.

Thus one can conclude that the limestone flora of Malaya (and Sarawak) is

extremely rich in species, intrinsically and also relatively, in comparison with the

peat swamp flora of Sarawak and Brunei, and most probably with floras of other

edaphic climaxes.

The total number of families recorded from limestone is 124; of these 117 are

families of spermatophytes. The total number of families of flowering plants

recorded from Malaya (both native and introduced) is 188 (figure obtained by

totalling the list in Keng, 1969). The delimitation of families in this study is the

same as in Keng except for Icacinaceae, which is split into Cardiopteridaceae and

Icacinaceae, and Tiliaceae which has been treated as Elaeocarpaceae and Tiliaceae.

With the addition of these two families the total number of plant families in Malaya

becomes 190; the total recorded from limestone amounts to 61.6% of this.

This rich flora differs from that of the rest of Malaya only in degree and not

in kind (Henderson 1939). No one family or group of families predominates or is

confined to the limestone except for two small families represented by very few

species in Malaya, Buxaceae (three species) and Primulaceae (one species). The

representation of a family on the limestone also more or less reflects the size of

the family as represented in Malaya. Table 1 shows the twelve largest families in

Malaya (from Keng, 1969) which is based on Ridley, 1925, with the total genera

and species from limestone compared beside them.

Malayan Limestone Flora 187

TABLE 1. GENERA AND SPECIES IN THE TWELVE LARGEST (SPECIESWISE)

FAMILIES IN MALAYA AND THEIR OCCURRENCE ON LIMESTONE

Families Genera Species

Malaya Limestone Malaya Limestone

Orchidaceae 104 52 669 136

Rubiaceae 67 27 368 66

Euphorbiaceae 71 37 343 81

Leguminosae 66 19 266 31

Gramineae 83 26 205 32

Annonaceae 30 19 184 39

Palmae 30 12 181 18

Melastomaceae 25 8 174 18

Lauraceae 16 8 174 13

Acanthaceae 36 16 168 32

Gesneriaceae 21 11 161 39

Zingiberaceae 20 9 157 16

Averages: Ratio z : 1 6 : 1

On limestone, Orchidaceae still tops the list but there are proportionally more

members of the Euphorbiaceae than Rubiaceae. The limestone seems a little poor

in Leguminosae, Palmae, Melastomaceae, Lauraceae and Zingiberaceae though

the other families are more or less proportionally represented. The reasons for this

difference may be various and not conjectured as knowledge on the requirements

of these plants is almost non-existent.

The families recorded from Malaya but not represented on the limestone are:

GYMNOSPERMS Te hemtstie eae Simaroubaceae

. onystylaceae Stylidiaceae

as 8 aie gn oe pi Shae

£ Hydrophyllaceae ymplocaceae

Dicotyledons Illiciaceae Th sdeeak

Aceraceae Juglandaceae Turneraceae

Actinidiaceae Linaceae Umbelliferae

Aizoaceae 2 25 iE ON

Ancistrocladaceae Malpighiaceae

Basellaceae Monotropaceae Monocotyledons

Bixaceae Moringaceae Bromeliaceae

Cactaceae Myricaceae Butomaceae

Campanulaceae Nymphaeaceae Cannaceae

Caricaceae Olacaceae Eriocaulaceae

Caryophyllaceae Opiliaceae Hydrocharitaceae

Casuarinaceae Orobanchaceae Iridaceae

Chenopodiaceae Pedaliaceae Juncaceae

Clethraceae Plantaginaceae Lemnaceae

Cornaceae Plumbaginaceae Najadaceae

Crassulaceae Portulacaceae Philydraceae

Cruciferae Proteaceae Pontamogetonaceae

Cunoniaceae Rafflesiaceae Pontederiaceae

Daphniphyllaceae Sabiaceae Restionaceae

Dichapetalaceae Salicaceae Smilacaceae

Droseraceae Saxifragaceae Typhaceae

Elaeagnaceae Schisandraceae Xyridaceae

These 71 families can be divided into six groups.

1. Aquatic or marsh plants. These are represented by 12 families, and include

most of the monocots. They are, Nymphaeaceae, Stylidiaceae, Butomaceae,

Eriocaulaceae, Hydrocharitaceae, Lemnaceae, Najadaceae, Philydraceae, Ponta-

mogetonaceae, Pontederiaceae, Restionaceae, and Typhaceae.

188 Gardens’ Bulletin, Singapore — XXX (1977)

2. Parasitic plants. These are represented by two families, Orobanchaceae

and Rafflesiaceae.

3. Coastal plants. Represented by three families, Casuarinaceae, Goodenia-

ceae and Portulacaceae. Aizoaceae also has a coastal member in Sesuvium, while

the remaining two genera in Malaya are not very common herbs.

4. Weeds or plants in cultivation. These are represented by 17 families,

Pinaceae, Basellaceae, Bixaceae, Caricaceae, Caryophyllaceae, Cactaceae, Cheno-

podiaceae, Crassulaceae, Cruciferae, Moringaceae, Pedaliaceae, Plantaginaceae,

Plumbaginaceae, Salicaceae, Turneraceae, Bromeliaceae and Iridaceae.

5. Very rare plants. In many cases these families are represented by a

solitary species in Malaya. There are 15 such families. They are, Aceraceae,

Ancistrocladaceae, Clethraceae, Cunoniaceae, Daphniphyllaceae, Droseraceae,

Elaeagnaceae, Epacridaceae, Hydrophyllaceae, Illiciaceae, Magnoliaceae, Mono-

tropaceae, Myricaceae, Cannaceae and Juncaceae.

6. Others. These remaining 23 families are often small and uncommon. They

are Actinidiaceae, Aizoaceae, Campanulaceae, Cornaceae, Dichapetalaceae, Gony-

stylaceae, Juglandaceae, Linaceae, Malpighiaceae, Olacaceae, Opiliaceae, Protea-

ceae, Sabiaceae, Saxifragaceae, Schisandraceae, Simaroubaceae, Styracaceae,

Symplocaceae, Theaceae, Umbelliferae, Smilacaceae and Xyridaceae.

It is unlikely that members of group 1 will ever be found on limestone, but

it is probable that members of group 2 are present on limestone. Members of

group 3 could conceivably be present on coastal limestone barring any physiological

antagonism to limestone. When more limestone areas are disturbed by man, some,

particularly the weed members of group 4 will probably be recorded. Members

group 5 which are very rare plants will, even if they are recorded on limestone,

also be very rare. New records for limestone from families not yet represented will

therefore most likely be from group 6, but there will not be many species, for most

of these families are small. The greatest number of new records will probably be

from families already represented on the limestone.

Phytogeography

The 1216 species recorded from the limestone can be grouped according to

their geographical distribution into a number of classes, viz:

PANTROPIC: Species occurring throughout the tropical regions of the world.

PALEOTROPIC: Species occurring in the tropics of the Old World, from Africa

and India through Malesia and to the Pacific.

INDO-MALESIAN: Species disturbed in mainland Asia as well as in Malesia;

including India to the Himalayas, Burma, S. China, Indochina, Thailand,

Malesia and southwards to Australia and the Pacific Islands.

ASIATIC: Species only in mainland Asia, sometimes only from Thailand.

MALESIAN: Species occurring in Malesia, including Malaysia, Indonesia, —

Philippines, New Guinea and sometimes to Australia and the Pacific.

ENDEMIC: Species occurring only in Malaya. These are subdivided into

(a) Species restricted to limestone in Malaya.

(b) Species not restricted to limestone in Malaya.

Malayan Limestone Flora 189

The result of this analysis is shown in Table 2.

TABLE 2. GEOGRAPHICAL DISTRIBUTION OF THE SPECIES

NUMBER OF SPECIES

Pterido- Gymno- Angio-

phytes sperms sperms Total To

PANTROPIC 6 — 14 20 1.7

PALEOTROPIC 8 --- 27 35 29

INDO-MALESIAN 40 4 315 359 29.6

ASIATIC (Asia mainland) 5 1 228 234 19.0

MALESIAN 32 1 225 257 21.2

ENDEMIC (total) 5 — 266 272 22.5

(a. restricted to limestone 2 — 127 129 10.7)

(b. not restricted to limestone 3 — 139 143 11.8)

WEEDS AND ESCAPES 1 —_ 38 39 2 fd

From the table it can be seen that the pantropic element is negligible; this is

because most species of this group are weeds (plants that occur in cultivated

habitats as undesirables, also in lawns and other open places and often as members

of secondary vegetation) and have been included under that group. The widely

distributed paleotropic element is also small. There is no predominance of either

the Asiatic or Malesian element, both being equally represented except for the

ferns. The weeds and escapes are noticably few in number; this is probably because

there are few areas on the limestone that support a secondary vegetation. However,

this is not to say that the limestone hills have been free from interference, but, that

when the limestone hills are disturbed it usually means that the habitat itself is

bodily removed as for instance in mining and quarrying.

Krasan (1882) and Steenis (1934) suggested that on limestone mountains

plants might be able to descend abnormally low. Although there are no really high

limestone mountains in Malaya this tendency can be seen. Here the mountain

massif has many peaks rising to over 1500 m, with Gunong Tahan rising to 2187 m.

Plants known from 800-2000 m (very rarely from lower elevation, if ever) have

been ae on low elevation (usually jess than 300 m) on the limestone and not

elsewhere.

The species recorded include Antrophyum semicostatum, Leptochilus

decurrens, Vittaria angustifolia, Alyxia angustifolia, Alyxia pumila, Corybas,

mucronatus, Distylium stellare, Liparis compressa, Paphiopedilum lowii, Schefflera

elegans, Sciaphila asterias, Scycopsis dunnii and Toxocarpus curtisii.

5. AFFINITIES OF THE FLORISTIC COMPONENTS TO LIMESTONE

The component species of the Malayan limestone can be arranged in the

following four groups:

I. EXCLUSIVES: These are species which, in Malaya, are restricted to the

limestone and include, (a) Species endemic to Malaya. (b) Species not

endemic to Malaya.

II. PREFERENTS: Species with a preference for the limestone field, appear-

ing in both fields (limestone and non-limestone) but more abundantly in the

limestone field.

Ill. INDIFFERENTS: Species with no affinity for either field, appearing in

both fields without exhibiting any difference in abundance between them.

This group includes most epiphytes and plants of secondary vegetation.

IV. STRANGERS: Species appearing accidentally on the limestone field;

frequently collected from the non-limestone field but only rarely from the

limestone field.

The criterion of fidelity (Brown-Blanquet, 1932) is used in constructing this

grouping, fidelity being indicative of the degree with which a species is restricted

to a particular kind of community (in this case the limestone community).

190 Gardens’ Bulletin, Singapore — XXX (1977)

The fidelity of each species and thus the allocation to the various groups is

based solely on field work. Species under groups I and II are termed the

characteristic species.

Rarely collected species which have been recorded both from limestone and

non-limestone fields are included under group II (indifferents) instead of under

group IV (strangers). This is because the limestone areas are probably the least

botanised of all the vegetation types in Malaya and chances are that rare species

already recorded from limestone are more likely to turn up from it again rather

than from the other vegetation types.

Weeds and plants escaped from cultivation usually fit into groups III (indiffe-

rents) or IV (strangers). However, here they are listed separately.

The true and intrinsic reasons for a plant to be on the limestone field could

also be used to classify the floristic components. Any of the plants found on

limestone could be:

(a) Calciphilous (chalk-loving) in the true physiological sense (Jackson, 1928).

They may be obligate calciphiles, in which case the limestone field is a prerequisite

for growth and survival, or non-obligate. The non-obligates have a physiological

need or affinity for limestone but are able to survive without the limestone field.

A large number of species in my group I (exclusives) are probably obligate

calciphiles while some (those that are non-endemic which are also found away

from the limestone outside Malaya) are probably non-obligate calciphiles. These

non-obligate calciphiles will also include a large number of species from my

group II (preferents).

(b) Indifferents; species that are able to tolerate or are unaffected, physiologically,

by the limestone habitat; as in my group III.

Obligate calcifuges cannot tolerate the limestone field whereas the non-obligate

calcifuges may often appear as strangers on the limestone field when the other

criteria of space, competition and moisture are favourable. The non-obligates

include at least some of my group IV.

However, this classification can only be effected by experimental work.

The species list of the different groups follows and are summed up in Table 3.

PTERIDOPHYTES

Adiantum zollingeri

Antrophyum parvulum

Cheilanthes farinosa

*Doryopteris allenae

Doryopteris ludens

Drynaria bonii

Heterogonium alderwereltii

Heterogonium pinnatum

Lygodium polystachyum

Nephrolepis dicksonioides

*Polystichum lindsaeifolium

Pyrrosia stigmosa

Schizaea inopinata

Tectaria amplifolia

Tectaria devexa

GYMNOSPERMS

Cycas siamensis

ANGIOSPERMS

DICOTYLEDONS

Actephila excelsa

Aeschynanthus longicaulis

I. EXCLUSIVES

*Amaracarpus saxicola

Andrographis tenuiflora

*Aporuellia sumatrensis var.

ridleyi

*Adisia biflora

*Ardisia langkawiensis

*Ardisia meziana

Argostemma diversifolium

*Barleria siamensis var.

glabrescens

Begonia curtisii

*Begonia foxworthyi

*Begonia ignorata

*Begonia kingiana

*Begonia nurii

*Begonia phoeniogramma

Boea acutifolia

*Boea brachycarpa

*Boea caerulescens

*Boea divaricata

*Boea lanata

*Boea minutiflora

*Boea paniculata

*Boea parviflora

*Boea suffruticosa

*Endemic to Malaya.

Boea treubii

*Boea verticillata

Boerhavia chinensis

*Buxus holttumiana

*Buxus malayana

*Buxus rupicola

*Callicarpa angustifolia

*Canarium perlisanum

Canscora pentanthera

*Canthium aciculatum

Capparis diffusa

Celtis philippensis

*Chirita caliginosa

Chirita hamosa

Chirita involucrata

Chirita lacunosa

*Chirita rupestris

*Chirita sericea

*Cleistanthus parvifolius

Cnesmone laevis

*Cnesmone subpeltata

Colona javanica

Cynoctonum mitreola

Dendronide sinuata

Dichiloboea speciosa

Malayan Limestone Flora

Dicotyledons (cont.)

*Dicliptera rosea

Dimocarpus longan ssp.

longan var. longan

Diospyros daemona

*Diospyros holttumii

Diospyros retrofracta

Diospyros transitoria

Diospyros undulata

*Dischidia tomentella

*Drypetes nervosa

*Embelia calcarea

Euphorbia antiquorum

Excoecaria oppositifolia

*Fagraea calcarea

Ficus calcicola

Garuga floribunda

Glossocarya mollis

*Glycosmis calcicola

*Glycosmis calcicola var.

kelantanica

Glyptopetalum zeylanicum

*Gongylosperma lanuginosum

Goniothalamus subevenius

Grewia viminea

Gymnanthera insularum

*Gymnostachyum robinsonii

Heritiera pterospermoides

*Homalium kunstleri

Homalium undulatum

*Hoya occlusa

Hydnocarpus ilicifolia

*Impatiens crytoneura

Impatiens marosepala

Impatiens mirabilis

Impatiens opinata

*Impatiens ridleyi

Impatiens scortechinii

*Impatiens tipusensis

Impatiens vaughanii

*Isonandra perakensis var.

kelantanensis

*Isonandra perakensis var.

perakensis

*Jasminum cordatum

*Jasminum curtisii

*Jasminum sp. A aff.

trinerve

*Jasminum sp. B

*Justicia microcarpa

*Justicia robinsonili

*Justicia rupestris

*Justicia subalternans

Justicia valida

*Kopsia griffithii var.

paucinervia

*Lagerstroemia langkawiensis

*Lasiobema flavum

Leptopus australis

Leucas mollissima

Ligustrum confusum

Lysimachia peduncularis

*Madhuca calcicola

*Mallotus bracteatus

Mallotus brevipetiolatus

Mallotus cuneatus

Maytenus curtisii

*Melodinus perakensis

Microphium pubescens

*Miliusa parviflora

*Millettia pterocarpa

Monophyllaea glabra

*Monophyllaea hirticalyx

Oidentandia rosettifolia

*Ophiorrhiza fruticosa

Ophiorrhiza kunstleri

*Ophiorrhiza longerepens

Ornithoboea flexuosa

Orophea cuneiformis

*Orophea hirsuta

Orophea polycarpa

*Osmanthus scortechinii

*Paraboea bakeri

*Paraboea bettiana

*Paraboea ferruginea

*Paraboea laxa

*Paraboea vulpina

Parishia rosea

*Pavetta pauciflora

Pentacme siamensis

Pentaspadon curtisii

Peperomia dindigulensis

Peperomia portulacoides

*Peperomia sp. A.

*Phanera decumbens

Phyllanthus columnaris

Phyllanthus ridleyanus

*Pilea fruticosa

*Piper collinum

Pisonia aculeata

Piscnia umbellifera

*Pistacia malayana

*Plectranthus kunstleri

Polygala cardiocarpa

Polygala malesiana

Polygala triphylla

*Polytrema cupreum

*Popowia velutina

*Rungia minutiflora

Sageretia thea

Sapium insigne

*Sauropus calcareus

Sauropus macranthus

Sauropus suberosus

*Scheffera subracemosa

Scleropyrum wallichianum

*Scolopia steenisiana

Semecarpus glomerulata

Solanum biflorum

*Sonerila elliptica

Sonerila tenera

*Stenothyrsus ridleyi

Stephania venosa

Sterculia angustifolia

Sterculia lancaviensis

*Strobilanthes pachyphyllus

Tarenna angustifolia

*Tarenna calcarea

Tarenna curtisii

Tetrameles nudiflora

Timonius atropurpureus

*Toxocarpus pauciflorus

Trigonostemon verticillatus

*Tylophora calcicola

Vernonia curtisii

*Vernonia rupicola

Vitex siamica

191

*Vitis scortechinii

Wikstroemia

androsaemifolia

*Zanonia clarkei

*Zizyphus pernettyoides

MONOCOTYLEDONS

*Aglaonema costatum

Amorphophallus carnosus

Amorphophallus

haematospadix

Arisaema fimbriatum

Arthraxon prionodes

Asparagus racemosus

*Boesenbergia curtisii

*Calamus balingensis

Calanthe rubens

Calanthe vestita

*Carex malaccensis

Carex speciosa

*Corymborchis brevistylis

Cymbopogon calcicola

*Dendrobium langkawiense

Dendrocalamus dumosus

*Dendrocalamus elegans

Dichanthium annulatum

*Dichanthium mucronulatum

Dracaena curtisii

Dracaena yuccaefolia

Eulalia quadrinervis

Eulophia keithii

*Fimbristylis calcicola

*Fimbristylis malayana

*Fimbristylis trichophylla

*Globba albiflora var. aurea

Goodyera hispida

Habenaria carnea

*Habenaria kingii

Hapaline brownii

*Isachne langkawiensis

Kaempferia elegans

Kaempferia pulchra

*Liberbaileya gracilis

*Malaxis reniloba

*Malleola undulata

*Maxburretia rupicola

*Oberonia calcicola

Oberonia caudata

*Oberonia transversiloba

*Pandanus calcicola

*Pandanus irregularis

*Pandanus piniformis

Paphiopedilum niveum

Pollia subumbellata

Pomatocalpa naevatum

Pomatocalpa setulense

*Pothos lorispatha

*Pteroceras tanyphyllum

*Raphidophora kunstleri

Sarcanthus termissus

Spathoglottis hardingiana

*Stachyphrynium cylindricum

*Thelasis succosa

Trichoglottis winkleri var.

minor

*Typhonium filiforme

*Typhonium fultum

*Endemic to Malaya.

192

PTERODOPHYTES

Adiantum malesianum

Adiantum soboliferum

Asplenium adiantoides

Asplenium squamulatum

Cyclopeltis crenata

Hemionitis arifolia

Pteridrys syrmatica

*Pteris longipinnula var. b

Pyrrosia penangiana

GYMNOSPERMS

ANGIOSPERMS

DICOTYLEDONS

*Argostemma inaequilaterum

Argostemma pictum

Berrya cordifolia

Chirita viola

Cladogynos orientalis

Cleistanthus gracilis

*Cleistanthus kingii

Colona merguensis

Croton cascarilloides

Cymaria dichotoma

*Debregeasia squamata

Deeringia polysperma

*Dehaasia curtisii

esmodium rugosum

PTERIDOPHYTES

Adiantum stenochlamys

Asplenium macrophyllum

Asplenium pellucidum

Asplenium salignum

Davallia denticulata

Davallia solida

Drynaria quercifolia

Drynaria rigidula

Humata pectinata

Loxogramme avenia

Loxogramme scolopendrina

Nephrolepis falcata

Photinopteris speciosa

Phymatodes nigrescens

Phymatodes papillosum

Phymatodes scolopendria

Pityrogramma calomelanos

Pteridium aquilinum var.

wightianum

Pteridium caudatum var.

yarrabense

Pteris ensiformis

Pteris tripartita

Pteris vittata

Pyrrosia adnascens

Pyrrosia varia

Tectaria variolosum

Thelypteris immersa

Trichomanes bipunctatum

Gardens’ Bulletin, Singapore — XXX (1977)

Il. PREFERENTS

*Drypetes oxyodonta

Epithema saxatile

Fagraea carnosa

Fagraea curtisil

Ficus curtipes

Garcinia minutiflora

Gomphostemma crinitum

Hedyotis coronaria

*Heterostemma piperifolium

Hopea ferrea

*Hoya citrina

Illigera pulchra

*Ixora clerodendron

Jasminum wrayi

*Justicia henicophylla

Lasiobema curtisii

*Lasiobema strychnoideum

*Leea saxatilis

Litsea glutinosa

*Madhuca ridleyi

Mallotus dispar

Mallotus miquelianus

Memecylon laevigatum

*Miliusa amplexicaulis

Monophyllaea horsfieldii

Murraya paniculata

*Orophea maculata

*Paraboea capitata

*Peperomia kotana

*Phyllanthus filicifolius

Procris pedunculata

il. INDIFFERENTS

GYMNOSPERMS

Cycas rumphii

Podocarpus polstachyus

ANGIOSPERMS

DICOTYLEDONS

Actephila ovalis

Adenia nicobarica

Aeschynanthus parvifolia

*Aglaia argentea

Aglaia odoratissima

Aglaia splendens

Agrostistachys gaudichaudii

Allophylus cobbe var. glaber

Allophylus cobbe var.

villosus

Alyxia selangorica

Antidesma japonicum

*Ardisia fulva

*Ardisia kunstleri

Ardisia lancelolata

Ardisia oxyphylla

*Ardisia solanacea var. elata

*Ardisia vaughani

Argyreia maingayi

Argyreia mollis

*Artabotrys grandifolius

Atalantia monophylla

*Atalantia roxburghiana

Rinorea bengalensis

Sauropus villosus

Sumbaviopsis albicans

Terminalia triptera

Trigonostemon viridissimus

Vitis discolor

Wikstroemia indica

Zizyphus oenoplia

MONOCOTYLEDONS

Abdominea minimiflora

Amomum testaceum

Apluda mutica

Appendicula torta

Arachnis flos-aeris

Bulbophyllum fenestratum

*Bulbophyllum flammuliferum

Bulbophyllum lilacinum

Dendrobium salaccense

Dioscorea calcicola

Dracaena congesta

*Dracaena graminifolia

*Globba fasciata

Habenaria reflexa

Pholidota pallida

*Pothos macrocephalus

*Sarcanthus rugulosus

Trichoglottis retusa

Uncifera tenuicaulis

Becheria parviflora

*Begonia debilis

Begonia guttata

*Beilschmiedia lumutensis

Bombax anceps

Bridelia ovata

Bridelia tomentosa

Callicarpa lanata

Canthium didymum

Carallia brachiata

Casearia capitellata

Cassia timoriensis

Cinnamomum iners

Clausena excavata

*Cleidion javanicum

Cleistanthus decurrens

*Cleistanthus glaucus

*Cleistanthus macrophyllus

Clerodendron penduliflorum

Clerodendron serratum

Clidemia hirta

Combretum porterianum

Congea vestita

Connarus sp.

Cordia griffithii

Cordia obliqua

Cratoxylum maingayi

Cyclea laxiflora

*Dalbergia kunstleri

Dalbergia phyllanthoides

Dalbergia scortechinii

*Endemic to Malaya.

Malayan Limestone Flora

Dicotyledons (cont.)

Decaspermum fruticosum

*Dehaasia longipedicellata

Dendrocnide stimulans

Desmos cochinchinensis

Desmos dasymaschalus var.

wallichii

Desmos dunalii

*Diospyros adenophora

Diospyros buxifolia

Diospyros cauliflora

*Diospyros ellipsoidea

Diospyros ferrea

Diospyros frutescens

Diospyros toposiodes

Dischidia benghalensis

Dischidia hirsuta

*Dischidia scortechinii

Ehretia timorensis

*Elatostema curtisii

Elatostema latifolium

*Enicosanthum congregatum

Eriobotrya bengalensis

Erismanthus obliquus

*Ervatamia peduncularis

Erythroxylum cuneatum

*Eugenia pendens

*Eugenia porphyranthera

Euonymus cochinchinensis

Fagraea blumei

Ficus binnendykii

Ficus deltoidea

Ficus elastica

Ficus hispida

Ficus microcarpa

Ficus sagittata

Ficus subulata

Ficus sundaica

Ficus superba var. japonica

Ficus tinctoria ssp. gibbosa

Ficus virens var. glabella

Ficus villosa

Garcinia murdochii |

Garcinia nigrolineata

*Garcinia opaca

Geophila repens

Glochidion rubrum

Glycosmis puberula

Glycosmis rupestris

*Glycomis sapindoides

Glytopetalum quadrangulare

Gomphia serrata

Gomphostemma javanicum

Gomphostemma microcalyx

*Goniothalamus fulvus

Grewia acuminata

*Gymnostachyum decurrens

*Gymnostachyum

diversifolium

Hedvotis tenellifiora

Hedyotis verticillata

Helicteres angustifolia

Helicteres hirsuta

Helixanthera axillaris

Helixanthera pulchra

Hemigraphis ridleyi

Heritiera littoralis

Holarrhena curtisii

Homalanthus populneus ©

Homalium dasyanthum

Hoya coronaria

Hoya latifolia

*Hoya maingayi

Hoya parviliora

Hoya ridleyi

Hydnocarpus wrayl

Iodes cirrhosa

Iodes ovalis

Ipomoea illustris

Ixora nigricans var. ovalis

Ixora pendula

Ixora umbellatta var.

multibracteata

Jacquemontia paniculata

Jasminum adenophyllum

Jasminum insularum

*Justicia pectinella

*Justicia ptychostoma

*Justicia subcymosa

*Justicia uber

Knema globularia

Knema laurina

Kopsia pauciflora

Laportea interrupta

Leea aequata

Leea rubra

Leea sambucina

Leptonychia glabra

Ligustrum robustum

Litsea norohae

Loeseneriella pauciflora

Macaranga tanarius

Macrosolen cochinchinensis

*Maesa pahangiana

Malaisia scandens

Mallotus wrayi

Mammea brevipes

Marsdenia tinctoria

*Medinilla scortechinii

Meiogyne virgata

Melochia umbellata

*Melodinus orientalis

Melothria affinis

*Memecylon dichotomum

Memecylon edule

Memecylon floribundum

*Memecylon kunstleri

Memecylon pauciflorum

*Memecylon wallichii

Microdesmis casearifolia

Micromelum minutum

Milusa longipes

Mimusops elengi

Momordica subangulata

Morinda elliptica

Morinda umbellata

Mucuna biplicata

Mycetia malayana

Myrsine porteriana

Naravelia dasyoneura

Neolitsea zeylanica

Oldenlandia ovatifolia

Oldenlandia pterita

Ophiorrhiza discolor

Ophiorrhiza hispidula

*Endemic to Malaya.

193

*Ophiorrhiza pallidula

Ophiorrhiza remotiflora

Orophea enterocarpa

Orthosiphon aristatus

Oxymitra biglandulosa

Pachycentria constricta

Paederia tomentosa

Paramignya scandens

Paranephelium

macrophyllum

Payena lucida

Petunga hirta

Phanera glauca

Phyllanthus oxyphyllus

Phyllanthus pulcher

Phyllanthus sikkimensis

*Piper mucronatum

*Piper scortechinii

Piper umbellatum

Planchonella obovata

“Plethiandra sessiliflora

Pogonanthera pulverulenta

Poikilospermum suaveolens

*Polyalthia brunneifolia

*Polyalthia lateritia

Polytrema uber

*Prema rubens

Prismatomeris malayana

Pseuderanthemum

crenulatum

Pseuderanthemum

graciliflorum

Pseuduvaria macrophylla

Pseuduvaria setosa

Psychotria angulata

*Psychotria cantleyi

Psychotria montana

Psychotria rhinocerotis

Pterolobium densiflorum

Pterospermum jackianum

Pterospermum pectiniforme

*Quisqualis parvifolia

Radermachera lobbii

Randia densiflora

Rauwolfia reflexa

*Richeriella malayana

Rinorea horneri

Rostellaria procumbens

*Rubus angulosus

Ruellia repens

*Rhus perakensis

Salacia grandiflora

Salacia korthalsiana

Salacia macrophylla

Salomonia ciliata

Saraca declinata

Saraca thaipingensis

Secamone micrantha

Schefflera junghuhniana

*Schefflera musangensis

Schefflera subulata

Schefflera tomentosa

Schefflera venolusa

Scolopia spinosa

Scurrula ferruginea

Scutellaria discolor var.

discolor

194

Dicotyledons (cont.)

Sida javensis

Stauraninera grandifolia

Steiecnocarpus Caulllorus

Sverculia rubiginosa

Suveoius asper

Sctreoius iuicifolius

Sireoius (axoides

*Strobilanthes leucopogon

Sirycnnos axillaris

*jarenna puichra

*Jarenna ridieyl

‘Lermunatia Calamansanai

*|oxocarpus Curtisi

Treina tomentosa

Trigonostemon aurantiacus

Trichosanthes tricuspidata

*Trigonosteinon villosus

*Tristania subauriculata

Trivaivaria macrophylla

Turpinia ovalifolia

Tylophora perakensis

Tylophora tenuis

Vaccinium littoreum

Vatica cinerea

Ventilago gladiata

Ventilago oblongifolia

Villebrunea sylvatica

Viscum orientale

Vitis furcata

Vitis glaberrima

Vitis hastata

*Vitis kunstleri

*Vitis mollissima

Vitis pyrrhodasys

Vitis repens

*Vitis wrayi

Wrightia dubia

Wrightia laevis

MONOCOTYLEDONS

*Adenoncos major

Adenoncos parviflora

Adenoncos sumatrana

Aglaonzma oblongifolium

Agrostophyllum

bicuspidatum

Alocasia denudata

Alocasia lowii

PTERIDOPHYTES

Abacopteris urophylla

Antrophyum callifolium

Antrophyum semicostatum

Arcypteris irregularis

Asplenium phyllitidis

Asplenium unilaterale

Athyrium cordifolium

Athyrium esculentum

Athyrium montanum

Gardens’ Bulletin, Singapore — XXX (1977)

Amorphophallus prainii

Amorphopnalius variabilis

Auyariuin hurmile

Anauvendrum marginatum

*Arenga hookeriana

*Arenga weslermoutil

Afrisaeuia roxburghil

borassogendron suachadonis

Bulvvpiylium apodum

Bulbophyllum concinnum

Bbuloopnylium

Meuibranaceum

Buriuiannia championii

Burmannia lutescens

Calamus concinnus

Calamus siamensis var.

mailaianus

Calanthe ceciliae

Caianthe triplicata

Camarotis apiculata

Carex breviscapa —

Caryota mitis

Catiunbium speciosum

Centhotheca lappacea

Ceratostylis pendula

Chlorophytum orchidastrum

Chrysopogon fulvus

Chrysopogon orientalis

Coelogyne asperata

*Coelogyne pallens

Coelogyne pandurata

Colocasia gigantea

Corybas mucronatus

Corymborchis veratrifolia

Cymbidium finlaysonianum

Dendrobium aloifolium

Dendrobium spurium

Dichanthium caricosum

Dioscorea bulbifera

Dioscorea esculenta

Dioscorea filiformis

Dioscorea glabra

Dioscorea prazeri

Dracaena angustifolia

*Dracaena nutans

Ephemerantha luxurians

Epipremnum giganteum

Eria leptocarpa

IV. STRANGERS

Athyrium pinnatum

*Athyrium prescottianum

Blechnum finlaysonianum

Crypsinus enervis

Ctenopteris alata

Ctenopteris moultoni

Cyclosorus extensus

Cyclosorus interruptus

Cyclosorus megaphyllus

Cyclosorus unitus

*Endemic to Malaya.

Eria pulchella

Fimbristylis fusca

Fiubristyius fuscoides

*Forrestia monosperina

Hanguana malayana

*Hoimalomena deitoidea

*iguanura polyimorpna

Lasia aculeata

Liparis caespitosa

Liparis gibbosa

Malaxis calophylla

Matieola dentitera

Microsaccus ampullaceus

Microsaccus brevifolius

Microsaccus javensis

Neyraudia reynaudiana

*Oberonia flava

Oplismenus compositus

*Pandanus alticola

*Peliosanthes lurida

Phalaenopsis cornu-cervi

Phalaenopsis decumbens

Phreatia secunda

Podochilus lucescens

Podochilus microphyllus

Pogonatherum paniceum

Pollia sumatrana

Polystachya flavescens

Pomatocalpa spicatum

Pothos scandens

Pteroceras ciliatum

Pteroceras hirsutum

Renanthera histrionica

Rhychelytrum repens

Sarcanthus sacculatus

Sarcanthus scortechinii

Sarcanthus subulatus

Schismatoglottis calyptrata

*Schismatoglottis mutata

*Sciaphila asterias

Scindapsus hederaceus

Scindapsus perakensis

Scleria lithosperma

Stemona tuberosa

Thelasis micrantha

Thelasis triptera

*Tupistra grandis

Vandopsis gigantea

Drynaria sparsisora

Humata heterophylla

Lemmaphyllum accedens

Lepisorus longifolius

Leptochilus decurrens

Lindsaya lucida

Lygodium flexuosum

Microlepia speluncae

Microsorium musifolium

Microsorium punctatum

Malayan Limestone Flora

Pteridophytes (cont.)

Nephrolepis biserrata

Nephrolepis hirsutula

Nephrolepis radicans

Oleandra undulata

Pteris mertensioides

*Pteris scabripes

Pyrrosia floccigera

Taenitis blechnoides

Tectaria barberi

Tectaria griffithii

Tectaria macronta

Trichomanes christii

Trichomanes humile

Trichomanes motleyi

Vittaria angustifolia

Vittaria elongata var.

angustifolia

GYMNOSPERMS

Agathis dammara

Gnetum cuspidatum

Gnetum gnemon var.

tenerum

Podocarpus neriifolius

ANGIOSPERMS

DICOTYLEDONS

Acrotrema costatum

Adenosma capitatum

Aeschynanthus radicans

Agelaea borneensis

Alchornea rugosa

Alsomitra pubigera

Alstonia scholaris

*Alyxia angustifolia

*Alyxia pumila

Anaxagorea javanica

Antidesma montanum

Antidesma tomentosum

*Aporosa stellifera

Aralidium pinnatifidum

Ardisia andamanica

Ardisia crenata

Ardisia colorata var.

complanata

Ardisia pendula

*Ardisia playtyclada

Ardisia ridleyi

*Ardisia tahanica

Ardisia villosa

Azadirachta excelsa

Baccaurea lanceolata

Balanophora fungosa

Barleria prionitis

Barringtonia asiatica

*Barringtonia fusiformis

Barringtonia macrostachya

Bauhinia acuminata

*Beilschmiedia pahangensis

Biophytum adiantoides

Blumeodendron kurzii

Brassaiopsis polyacantha

Breynia vitis-idaea

Bridelia stipularis

Buchanania sessilifolia

Caesalpinia crista

Calophyllum curtisii

Cananga odorata

Canarium pilosum

195

Ficus trichocarpa var.

obtusa

Firmiana malayana

Canarium pseudodecumanum Garcinia cowa

Capparis pubiflora

Cardiopteris javanica

Casearia grewiaefolia

Centranthera hispida

Chloranthus elatior

Chukrassia tabularis

Citrus macroptera

Claoxylon longifolium

Cleistanthus hirsutulus

Cleistanthus polyphyllus

Clerodendron paniculatum

Cnesmone javanica

Coffea canephora

*Coffea malayana

Coleus scutellarioides

Colubrina asiatica

Combretum latifolium

Connarus semidecandrus

Cotylelobium malayanum

Croton argyratus

Croton erythrostachys

Croton laevifolius

Cryptocarya griffithiana

Curanga amara

Cynometra malaccensis

*Cyrtandra cupulata

*Cyrtandra lanceolata

*Dacryodes kingii

Dehassia microcarpa

Derris elliptica

Derris thyrsiflora

Desmodium umbellatum

Dillenia indica

Diospyros hermaphroditica

Diospyros malayana

Diospyros pilosanthera

Diospyros rigida

Diospyros wallichii

Diploclisia glaucescens

Dipterocarpus oblongifolius

Dischidia rafflesiana

Distylium stellare

Dryobalanops aromatica

Dryobalanops oblongifolia

Dysoxylum arborescens

Elaeocarpus pedunculatus

Endospermum diadenum

Erythrina variegata

Eugenia chlorantha

Eugenia claviflora

Eugenia spicata

Euonymus javanicus

Exacum tetragonum

Fagraea auriculata

Fagraea ceilanica

Ficus annulata

Ficus botryocarpa

Ficus montana

Ficus oligodon

Ficus parietalis

Ficus racemosa

Ficus scortechinii

Ficus semicordata

Ficus stricta

*Endemic to Malaya.

*Garcinia eugeniaefolia

Garcinia merguensis

*Garcinia montana

Glochidion obscurum

Glochidion perakense

Glycosmis chlorosperma

Gmelina asiatica

Gmelina villosa

Gomphandra quadrifida

var. quadrifida

Gomphostemma curtisii

Goniothalamus scortechinii

Goniothalmus uvarioides

Grewia paniculata

Guettarda speciosa

Gymnopetalum

cochinchinense

Hedyotis congesta

Homonoia riparia

Hopea dryobalanoides

Horsfieldia tomentosa

*Hoya revoluta

Hunteria zeylanica

Hydnocarpus castanea

Hydnocarpus woodii

Hydnophytum formicarum

Hygrophila angustifolia

Hypserpa cuspidata

*Ilex maingayi

Ixora brunonis

Ixora congesta

Ixora grandifolia

Ixora lobbii var.

stenophylla

Ixora nigricans

*Ixora scortechinii

Jasminum bifarium

Justicia vasculosa

Kibara chartacea

Knema cinerea var.

patentinervia

Knema cinerea var. rubens

Knoxia corymbosa

Koiledepas longifolium

*Kopsia macrophylla

Lantana camara var.

aculeata

Lasianthus stipularis

Lepistemon binectariferum

Leucaena leucocephala

Leucas zeylanica

*Lindera concinna

Lithocarpus elegans

Lithocarpus urceolaris

Litsea polyantha

Ludwigia hyssopifolia

Luvunga eleutherandra

Maesa striata

Mallotus eriocarpus

*Mallotus griffithianus

Mallotus oblongifolius

Mallotus peltatus

Mallotus philippensis

Mallotus repandus

196

Dicotyledons (cont.)

Medinilla crassifolia var.

hasseltii

Melaleuca cajuputi

Melanolepis multiglandulosa

Melastoma polyanthum

*Memecylon acuminatum

Memecylon oleaefolium

Mesua ferrea

Millettia hemsleyana

Millettia sericea

*Mitrephora maingayi

Moghania strobilifera

Naravelia laurifolia

Nauclea junghuhnili

Neesia synandra

Neonauclea calycina

Nepenthes sp.

Ophiorrhiza communis

Palaquium obovatum

Palaquium ottolanderi

Parameria polyneura

Parashorea lucida

Pavetta indica

Pavetta naucleiflora

Peltophorum pterocarpum

Phanera integrifolia

Phoebe lanceolata

Piper boehmeriaefolium

Piper caninum

*Piper porphyrophyllum

Piper retrofractum

Pittosporum ferrugineum

Polyalthia cauliflora var.

beccaril

Polyalthia cinnamomea

*Polyalthia hypogaea

Polyalthia motleyana var.

glabrescens

Polyalthia rumphii

Polyalthia stenopetala

Polygonum chinense

Premna pyramidata

Psychotria rostrata

Psychotria sarmentosa

Psychotria viridiflora

Pterygota alata

Pterisanthes coriacea

Rauvolfia perakensis

Reissantia indica

Rhododendron longiflorum

var. longiflorum

Rhynchoglossum obliquum

Rinorea anguifera

Rinorea macrophylla

Sageraea elliptica

Sauropus brevipes

*Schefflera elegans

Shorea guiso

Shorea leprosula

Shorea ovalis

Solanum decemdentatum

Spondias dulcis

Stauranthera umbrosa

Streblus laxiflorus

Gardens’ Bulletin, Singapore — XXX (1977)

Tamarindus indica

*Tarenna appressa

Tetracera scandens

Thespesia populnea

Thunbergia fragans var.

javanica

Tinomiscium petiolare

Tinospora crispa

Trema orientalis

Tristania merguensis

*Turraea brevitlora

Urophyllum corymbosum

Urophyllum glabrum

Utricularia minutissima

Uvaria cordata

Uvaria javana

Viburnum sambucinum

Vitex pubescens

Vitis lanceolaria

Vitis martinelli

Vitis novemfolia

Vitis peduncularis

Wikstroemia polyantha

Xanthophyllum glaucum

*Xerospermum wallichii

Xylocarpus granatum

Xylopia malayana

Zippelia begoniaefolia

MONOCOTYLEDONS

Acampe longifolia

*Achasma macrocheilos

Achasma megalocheilos

Achasma triorgyale

Aerides odoratum

Agrostophyllum hasseltii

Agrostophyllum majus

*Amomum biflorum

*Anadendrum latifolium

Anadendrum montanum

Aneilema nudiflorum

Appendicula anceps

Appendicula cornuta

Appendicula undulata

Areca triandra

Arenga pinnata

Ascochilopsis myosurus

Bulbophyllum pulchellum

Bulbophyllum sessile

*Calamus ornatus var.

horridus

Carex perakensis

Coelogyne rochussenii

*Corymborchis rhytidocarpa

Costus globosus

Costus speciosus

Crinum defixum

*Cryptocoryne affinis

*Cryptocoryne minima

*Cryptocoryne purpurea

Curculigo latifolia

Cymbidium dayanum

Cyperus trialatus

*Endemic to Malaya.

Cyrtosperma lasioides

Dendrobium acerosum

Dendrobium excavatum

Dendrobium farmeri

Dendrobium indivisum

Dendrobium leonis

Dendrobium planibulbe

Dendrobium pumilum

Dendrobium secundum

Dendrobium subulatum

*Dendrobium tetrodon

Dioscorea hispida

Dioscorea polyclades

Dioscorea prainiana

Dioscorea pyrifolia

Dipodium pictum

Donax grandis

*Dracaena porteri

Ephemerantha fimbriata

Epipremnopsis media

Eria citrina

Eria leiophylla

Eria nutans

Eria pannea

Eria vestita

Eurycles sylvestris

Geodorum citrinum

Globba patens

Gymnosiphon aphyllus

Hippeophylum scortechinii

Homalomena griffithii

Homalomena humilis

Homalomena rubra

Iguanura geonomaeformis

Ischaemum indicum

Ischaemum timorense

Leptaspis urceolata

*Licuala modesta

Liparis compressa

Livistona saribus

Malaxis latifolia

Malaxis micrantha

Musa malaccensis

Oberonia anceps

Oberonia dissitiflora

Oberonia spathulata

Onchosperma horridum

*Orchidantha longiflora

Pandanus odoratissimus

Pandanus recurvatus

Panicum sarmentosum

Paphiopedilum lowili

Paspalum conjugatum

Peliosanthes violacea

*Phaeomeria maingayi

Plectocomia griffithii

Poaephyllum pauciflorum

Podochilus tenuis

Pollia sorzogonensis

Pollia thyrsiflora

Polytrias amaura

Pomatocalpa kunstleri

Pomatocalpa latifolium

*Pothos latifolius

rc. ..

Malayan Limestone Flora 197

Monocotyledons (cont.)

Raphidophora beccarii Setaria palmifolia Thelasis carinata

Raphidophora korthalsii Staurochilus fasciatus Thrixspermum album

*Raphidophora maingayi Stenotaphrum helferi Thrixspermum amplexicaulis

Renanthera elongata Tacca leontopetaloides Th 1 :

Sarcanthus machadonis Taeniopyllum culciferum armen pics

Schoenorchis micrantha Taeniophyllum filiforme Trichoglottis musera

Scindapsus scortechinii Taeniophyllum obtusum Tropidia curculigoides

Scleria purpurascens Thecostete alata *Zingiber spectabile

WEEDS AND ESCAPES

PTERIDOPHYTES Erechtites Nagi tania age Solanum torvum

. Eupatorium odoratum Spondias pinnata

seserecoiirms Nae caaian Euphorbia hirta Tridax procumbens

GYMNOSPERMS Flacourtia jangomas Urena lobata

af Hyptis rhomboidea Vernonia cinerea

Ph ag savant

Mikania cordata MONOCOTYLEDONS

Muntingia calabura P tetas

DICOTYLEDONS Murraya koenigii Chrysopogon aciculatus

Abutilon indicum ssp. Passiflora foetida var. Coix lacryma-jobi

indicum hispida Colocasia antiquorum

Acalypha lanceolata Physalis minima Cyperus kyllingia

Ageratum conyzoides Pilea microphylla Digitaria violascens

Aleurites moluccana Piper nigrum Eleusine indica

Bidens pilosa Scoparia dulcis Imperata cylindrica var.

Boehmeria nivea Solanum ferox major

Cyathula prostrata Solanum nigrum Rhoeo spathacea

TABLE 3. AFFINITIES OF THE FLORISTIC COMPONENTS

NUMBER OF SPECIES

ee eet a

EXCLUSIVES sg ema 182 62 257 21.1

PREFERENTS 9 — 53 19 81 6.6

INDIFFERENTS 27 2 285 104 415 34.1

STRANGERS 45 3 257 131 424 34.9

WEEDS AND ESCAPES 1 — 29 9 39 3.2

The species known in Malaya only from limestone (exclusives, which

includes endemic and non-endemics) total 257. This is an increase of 62 over

Henderson’s (1939) figure of 195. However the percentage which is 21.1% of

the present total is less than Henderson’s figure of 26% of his total. This is

because of the even greater increase in the overall total (from 745 to 1216) of

the species recorded from the limestone. The total number of species endemic and

restricted to the limestone remains unchanged with no new additions. Henderson

recorded about 130 species, the present figure is exactly 129 species.

The characteristic species of the limestone which are the exclusives and

preferents number 338. This is 27.7% of the total flora. The indifferents and

strangers are represented by 839 species (69.0% of the total flora), forming the

bulk of the flora. The weeds and escapes are poorly represented by only 39

species (3.2% of the total flora).

198

Gardens’ Bulletin, Singapore — XXX (1977)

SECTION If — The Flora

PTERIDOPHYTES

INTRODUCTORY KEY TO THE FERNS

The numbers on the right hand side refer to the numbers in the main key

with which one should continue.

1. Fronds of two distinct forms; nest and foliage

1. Fronds not as above.

. Coendveset 2. Drynaria spp.

2. Filmy ferns; laminar part of fronds one cell thick .............,ccsssesessencees

« sa» MRRGERTEEY ~ee ber eeed> + <3 +a.05n05 -neethathen toe: on eeeeeeee ene 6. Trichomanes spp.

2. Not filmy ferns; fronds thicker, usually several cells thick.

3. Fronds twining

4. Fronds simple, erect, grass-like

os nhawe vn DEORE OTE MEIC dg tlle pee 11. Lygodium spp.

3. Fronds not twining.

4. Fronds not as above.

5. Fronds simple, entire or lobed.

6. Fronds entire.

ey ee yr 12. Schizaea inopinata

Base of fronds cordate or sagittate, stipitate. ......... 16

Athyrium cordifolium, Hemionitis arifolia, Doryop-

teris spp.

Base of fronds simple, if cordate then sessile.

8. Stipe articulated

8. Stipe if present, not articulated.

UTIs 19. Oleandra undulata

Sori marginal or sub-marginal ..................

MAY osereoey or) re eee ee: 20. Vittaria spp.

Sori not marginal or sub-marginal.

10.

Sori indusiate

heterophylla,

OTERO 23. Humata

Asplenium spp. (some)

Sori non-indusiate.

‘:

11.

Sori? acrostichoid ......4gekea 26.

Sori not acrostichoid.

12.

Sori elongate along veins ......

» ea, Tene 28. Loxo-

gramme spp., Antrophyum spp.

Sori not elongate.

13.

Frond pubescent with

stellate hairs ............ 3.

Pyrosia spp. .

Frond not so pubescent

PET yy ee Eee 39.

Lemmaphyllum_ accedens,

Lepisorus longifolius,

Microsorium spp. Cryp-

sinus enervis.

Malayan Limestone Flora 199

6. Fronds lobed to more than half way to rachis.

14. Sori only at the end of lobes ........................ 43.

Ctenopteris alata

14. Sori differently arranged.

Ne ee Sidr tanned Source enittemaras«.: 45.

Humata pectinata, Doryopteris spp.

i ne 47.

Polypodium papillosum, Phymatodes spp.

5. Fronds compound, pinnate or more divided.

16. Undersurface of fronds chalky-white .........0...00..0... 50.

Cheilanthes farinosa, Pityrogramma calomelanos

16. Not so.

17. Sori acrostichoid or sub-acrostichoid ............ 52.

Photinopteris speciosa, Heterogonium spp.

17. Sori differently arranged.

18. Sori marginal or sub-marginal, elongate ... 55.

Lindsaya lucida, Pteridium spp., Pteris spp.

18. Sori not marginal or sub-marginal, if marginal

or sub-marginal, then not elongate.

19. Sori marginal or sub-marginal.

20. Sori on reflexed marginal flaps

Soaked i dhcp Git plane 65. Adiantum spp.

20. Sori otherwise.

21. Fronds simply pinnate ...... 70.

Ctenopteris moulton, Nephro-

lepis spp. (some)

21. Fronds more divided ...... 73.

19. Sori not marginal or sub-marginal.

aPC ios cna ci dass ot sess one suss oeak i:

Taenitis blechnoides, Blechnum finlay-

sonianum, Athyrium spp., Asplenium

spp. (some)

22. Sori rounded.

23. Sori indusiate.

24. Fronds pinnate or bipinnati-

fid 86.

Polystichum lindsaefolium,

Pteridrys syrmatica, Cyclo-

peltis crenata, Nephrolepis

spp. (some), Thelypteris im-

mersa, Cyclosorus spp.

24. Fronds bipinnate or more

amply divided ............ 95.

Tectaria spp., Méicrolepia

speluncae

23. Sori non-indusiate ............... 101.

Abacopteris urophylla, Acrypteris

irregularis

16.

Gardens’ Bulletin, Singapore — XXX (1977)

FERNS — MAIN KEY

Fronds of two distinct forms, nest and foliage leaves .................esee0es 2

Fronds not differentiated into nest and foliage leaves ....................068+ 2

Nest leaves lobed 2 cm OF More. dlQD 5... u.sscrsc++cocvscnessanneneeennene 2

Nest leaves not or only very shallowly scalloped ......... Drynaria bonii

Foliage leaves pinnate; pinnae stalked .................. Drynaria rigidula

Foliage leaves 1QDE ..... ascscnexsnae+eagsie «sets oa ees peat cs ga needane aaa 4

Rhizome covered by short appressed scales. Sori in two irregular rows

« sid byLsald dw vida He aia PARSE GS VRE TS » oan ela anaes Drynaria_ sparsisora

Rhizome covered by long, semi-erect scales. Sori in two regular rows

oo veces cetubensbet tp premelurne rutys tuleuebiiersehascauenle mee Drynaria quercifolia

Sori enclosed by the hollow base of indusium. Sporangia on an elongate

receptacle. Filmy feist... ...<.:.2.. conasssscshes0ncssesansieunys tans <neeen 6

Sori different, Not ‘Mlmy. Tere”... : .7<;-s2.<cdsaseus sv pees canna peer teen 10

Rhizome.-erect; fronds tubtéd :: scm odwkl.... Trichomanes obscurum

Rhizome creeping; fronds borne at intervals .................ceceeeeeeeeeeees 7

Fronds simple, orbicular to broadly ovate ......... Trichomanes motleyi

Fronds’ pinniate'‘or rfiore dissected 30200...4/0. ATI 8

Fronds with false .Wegus cps fits ian ox sabesintensanp > nieptenenrayhonie cine 9

Fronds. with no. false VeqHie jes. -20sne>5 cdesceydewersnnes Trichomanes humile

Soral-lips of two triangular lobes ............... Trichomanes bipunctatum

Soral-lips not lobed, trumpet-like ...................65 Trichomanes christii

Fronds. twining, Jong chim ooops sicngep sen eia ans ccweweuen pa seane tegen 11

Fronds different: .....0..<éend uateh ee we ab WLM ees ESS Se 12

Sterile leaflets regularly lobed; plant of forest shade .....................04

scecseddetevabnsseeastischegaves Mian tata htaetn tee Tereeama Lygodium polystachyum

Sterile leaflets not lobed or only the basal leaflets lobed; usually in partly

Open places. ..... ... ice sunsnmpeeys+aseae any ae ae Lygodium flexuosum

Fronds simple, erect, narrow and grass-like ............ Schizaea inopinata

Fronds different... d:it-.s..rcsnderestesiowseacady est pany Ulleue ee te hein staan ttt 13

Fronds simple, entire or lobed ............0:..<aglesdsshitas pean 14

Fronds compound, pinnate or more divided ...............ccceceeeeeeeeeeen ones 49

Fronds entire or almost entire, base cordate, sagittate, or simple ......... 15

Fronds lobed, usually to more than half-way to rachis ..................06 43 ©

Frond base cordate or sagittate, stalked ........s0:...csasoessncesnntenspasvenannn 16

Frond base not cordate or sagittate, or if appearing cordate then

gy EEE OO oo eecepecesessosinudanaede» ty aid anne aoieine inna in 19

Fronds: broadly: lanceolate ..............ssesesseeesreees Athrium cordifolium

Malayan Limestone Flora

17.

18.

19.

20.

2h.

22.

235

24.

25.

26.

ay.

28.

29.

30.

31.

32.

33.

Stipes grooved on the adaxial side; sori superficial ... Hemionitis arifolia

Stipes not grooved; sori sub-marginal ..................cccccceseececeeeeeeeees

Hydathodes present as a series of sub-marginal white spots; marginal

eNIENR IMINO fois. os. cs denn- vcs cccs cviee.oceceds Doryopteris ludens

Hydathodes absent; marginal veins mostly jointed ... Doryopteris allenae

Stipes articulated at a point between the rhizome and blade ...............

eae ator eatec anata caclvaecesasrrudthacaces caste vers Oleandra undulata

ge nn RUDD 0 FORER sey Oy Eee Poe ee Ae eae

ee ee INNS = 000 RERIS-MUAO OTN So ss oleic ceva ig os gals inside se seese’scaupaieee

SOF! MOC MiAPial OF SUD-NATPINAL ooo... 5 <5. cece kc cece sect eesesenceceeses

Fronds sessile, 20 by 0.2—0.3 cm ..................05. Vittaria angustifolia.

Fronds stipitate, to 90 by 0.5 cm ...... Vittaria elongata var angustifolia

Stl WIGUSIOES 5s ess oce- Tevvexed: -heue- dopa vedpeeses 6 MEARE. Gal bkns -eep ane. de...

NG Se ESE FEIT OPEL DST PDO ig at SAE

Rhizome long creeping; fronds dimorphic ......... Humata heterophylla

Rhizome short, erect; fronds not dimorphic .....................cceeceeeeees

os OU. | a eee eres Cm Met i), ere Aes Oe

Feomidis: pinnate... dizhin.ods..to. akin acidiie.ne. os Asplenium salignum

Veins of fronds united in a sub-marginal vein ...... Asplenium phyllitidis

Veins of frosd-all free si). 225). becca). cbes 238 Asplenium squamulatum

Sori acrostichoid or sub-acrostichoid ............... Leptochilus decurrens

Sori not acrostichoid or sub-acrostichoid ..................ecececeeeceeeeeceees

Suen ere tS MCE ONC CL ST. SUL Sater cacne oun wclc os indcs-cuae.cecccasese

Sent Mot elomeet- lone: Veins! 20.22. eceicc ree Pe

Sori obique to the midrib, in roughly parallel rows ......................4.

Sori along many veins, forming a network ......................ccceeeeeeeeees

Midrib raised on the lower surface, almost flat above .....................

STE AE, / OLE EEE DENS EE Pe Ee Loxogramme scolopendrina

Midrib raised on the upper surface, almost flat below .....................55

aN eo ius tl ends beh each atdbenttalindees os oe Loxogramme avenia

Parapevees Mi SOT Club-shaped .. +... ... 0. feauccars asp - Wide ory bees omar ones

Paraphyses in sori thread-like ..................... Antrophyum callifolium

Stipes to 10 cm; fronds to 30 by 10 cm...... Antrophyum semicostatum

Stipes not distinct; fronds smaller, to 15 by 2 cm ....................020ee

ergy Gap map lee os Reka) sageny age 7k See ae Antrophyum parvulum

Fronds densely or sparsely covered by stellate hairs, more on the lower

NUNC a hciees - eye < Lge atete Chbice © Beg. nae ees - 45+ cage paeravae ablaseli gees pie eee

eee RS SCT POMPE Cla) DTOAG oo. i. li ccc ducectcccteccccecccetececess

Sterile fronds orbicular to broadly ovate ...... Pyrrosia nummularifolia

201

202

34.

35.

36.

ay;

38.

39.

40.

41.

42.

43.

45.

46.

Gardens’ Bulletin, Singapore — XXX (1977)

Sori in a single row on either side of the midrib, to about 0.5 cm across

spediepeepipesssoreagcsaese regs snennnnceeienpeareaesh eee Pyrrosia angustata

Fronds dimorphic; sterile shorter and sometimes wider ..................-

Fronds not dimorphic; sterile about the same size and shape ............

Sterile fronds 1 — 1.5 cm wide; fronds when dried, the margins involuted

the upper surface becoming concave; scales of rhizome not closely

BPPTESSEU soo... ; ssa ncilelhegoneceeeasepeet saan aen eee Pyrrosia adnascens

Sterile fronds 2 — 3.5 cm wide; fronds sine dried, the margins revoluted,

the upper surface becoming convex; scales of rhizome not closely

Spprested weiss Liv. tohe.. Heard Tia bods. cet kee Pyrrosia varia

Lower surface of frond densely brown-stellate pubescent ..................

a 8's aewad dunainw ss seine getline Pachbanviag .eaiaal he tee cane ntcere: Manian Pyrrosia stigmosa

Fronds about 1.8 cm wide; sori on the apical half or whole of frond; no

clear marginat the. edge... 200/42. 228OT.. Sa Pyrrosia floccigera

Fronds 3-6 cm wide; sori on the apical third of frond; with a clear

margin to 1 cm wide at the edge ..................... Pyrrosia penangiana

Sori in two rows, one on either side of the midrib ........................04.

Sori scattered all over the surface ...............cceceeeeeees peat - 2p ope ends

Fertile part of frond narrowed; frond less than 15 cm long ...............

eT ee ee Te mes ee seeseeeee. Lemmaphyllum accedens

Fertile part of frond not narrowed; frond to over 30 cm long ...............

a veeeas Sande wqdqaigomninee tepiliaey atte aaate net ren Ramee meee ... Lepisorus longifolius

Fronds stipitate’:. i5..52.0c2.5000<s9) tesa. cs appeee aatOle ene ee

Fronds sessile, widening abruptly from the base: .s5..:ipncis ie ee

winie ankceo dadlee og tes eg gree eaenio a en aeececeseeccsseeeeeeeese, Microsorium musifolium

Stipes 1 cm or less; fronds widening very gradually from the base .........

svevasecvedddeuin elle deed DEW adelc AED Le hDGIE ENE iceEmanne Microsorium punctatum —

Stipes 5 cm or more; fronds with broad bases ......... Crypsinus enervis

Sori only at the ends of the lobes of frond; usually 2—5 on each lobe

oy wo sibs «win dialuidcaspinaty 5 #G'anitidinenela peal P Ena Ctenopteris alata

Sori continuous along margins of fronds .................csceeeceeeeceeneneeees

Sori not so; in two rows, one on either side of midrib .....................06.

Hydathodes present as a series of submarginal white spots; marginal

veins mostly free. o.4).0:30000 BAER. AR, RE Doryopteris ludens

Hydathodes not present; marginal veins mostly joined ................+++

a ct nacadbescuscescp scenes epiinscnsnds wees ashes sitet

Malayan Limestone Flora

47.

48.

49.

50.

51.

52.

54.

Do-

56.

57.

58.

=. B

Veins anastomosing. Rhizome stout, to over 1 cm diameter ...............

Veins all free, forked once. Rhizome slender, 0.2 —- 0.3 cm diameter ......

Se epee Htabebrets OES we seta den cng 500 asesseeeeeee Polypodium papillosum

Fronds with 1-4 pairs of lobes; sometimes simple ........................

0 he le Baie ate eee ps ae Teo et fe a Phymatodes_ scolopendria

Fronds with 7-10 pairs of lobes; never simple .......................:.:0ee

ensaeaene teases 2s als ass ss, ha 2k4 css ia snd ves Phymatodes nigrescens

Underside of fronds completely or partially covered by chalky-white

SON Metta ies canna te celduiea Laka SCAM IEUAGLOY. 2352. L0se 220.00. Wk aa.

Underside of fronds not chalky-white ...................ccccceeeeceeeeeeeeeeees

Fronds to 20 cm long, often less; sporangia near the margin of frond

en 5 Ra eee ee Cheilanthes farinosa

Fronds much larger, 30-60 cm long; sporangia scattered all over the

eeeeTertMCe OF LOU oo. 5.0.0 cea> os 4 ye3ca8e5 «* Pityrogramma calomelanos

Sori acrostichoid or sub-acrostichoid, on the apical part of frond or

IRIN: OSM: SCN isi ea ae ria dhs inset Sun somes dignasicanenedss-

Sori not acrostichoid or sub-acrostichoid ..................ccceececeeeeeeee ces

ra OMT A AWG A RR ON. Photinopteris speciosa

ED uss <4 sccawans sx onbue anew hide dass wied ove kes. dUees (NEL ee TOI

Srcemteems ct SAY Ce) IO Heterogonium alderwereltii

oR aa ee eC Se ne Heterogonium pinnatum

Sori marginal or submarginal, continuous in a short or long band ......

Sori not marginal or submarginal, if marginal or submarginal, rounded

I CN SE Sean Seat ace teak twink suas anus hcthckoaensasesesesscese

Sori short banded, uniting the end of several veins ...... Lindsaya lucida

Seed > AUN Swans ik sannenapainas RN Mette adh inns ns hgndy <5

Fronds tripinnate or quadripinnatifid; of long continued growth .........

Fronds pinnate or tripinnatifid or rarely tripartite; not of long continuous

cla rig ce a ect ga le nea pg SRSA UOT ~ SELEY ICA ask AA 0

Ultimate segments widely spaced, connected by a wing decurrent from

the preceeding segment ............ Pteridium caudatum var. yarrabense

Ultimate segments close together, not connected by a wing ...............

Ye, BUTE aeeesescseeeseeesscessseees Pleridium aquilinum var. wightianum

PMUM RRR MNII, OUMEAIN S556) Syd ca cdccaasescscccescscsvescess Pteris vittata

a aN coats oc ock ss cdlacccqcccbdcccsscwandedqedsasseeesse

Fronds dimorphic in mode of branching; sterile, with deeply lobed

pinnae; fertile, with only the basal pinnae having a single basiscopic

IR Fe Od. as svat bans deséadsceadovewe Pteris ensiformis

204

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

7h.

TZ,

74.

Gardens’ Bulletin, Singapore — XXX (1977)

Pinnae, margin entire or serrulate .................eceeeeees Pteris scabripes

Fronds tripartite, basal pinnae almost as large as the terminal one .........

a sio-n ssln'o s,s.» 6's un o's ODAC APEEDS > Mp RE ep eetoae seit tani Oe lk een Pteris tripartita

Fronds not tripartite. .00y.5liici. 0.200. .cetbe deel en calves dallas Danae

Fronds less than 50 cm long; lowest vein in each pinnae lobe 3-4 times

FORCE)... FRAO Rah A ii Re Pteris mertensioides

Sori marginal or:submarginah: ov. . 21). .//Waviiiwa oven. 1a. Gee

Sori not. marginal. or. submargimal ............casccsccaccesanessennsenss SRE

Sori on small ‘reflexed margmal flaps 17.00.52... 2201.22 ee

Sofi 1.otherwise 913. isa. sien gana. cewel- ceties «eee eee a ee

Fronds simply pinnate «....62..2cccesccesecssees « coevrdaesieud taco teaeueenaann

Fronds tripinnate’or more ‘divided '):.//)2..VGs......19018). SAUL, BO

Stipes, rachises and stalks of pinnae winged ...... Adiantum soboliferum

Stipes, rachises and stalks of pinnae not winged .................cceeeeeeeeees

Rachis and pinnae hairy; indusial flaps almost circular .....................

PROT NRE OR MVOSEY ee Ger BY meg nt. hme canto ... Adiantum malesianum

Rachis hairy on the upper surface only; pinnae glabrous; indusial flaps

broader, than loge «iscsi vee eben huis ge easacecnees Adiantum zollingeri

Fronds tripinnate, basal angle of leaflet usually more than a right-angle

Joana is: -yetan aan Sts aa con poet ek ein as Adiantum stenochlamys

Fronds quadripinnate or more divided, basal angle of leaflet usually

less than "9. Tight-Aneye | o.0\s ccwecencs oneav aden beta meaea et Adiantum tenerum

Fronds sunply -qeniiate.:.3.9.5.:4-a3ue anes Te ey Rr

Fronds tripinnatifid..or more :dswided) sic. ..cs. aegis idan se seeemanaeiee ane

Sori indusiate; fronds 30 Ci OF MORE Jecschevaccevtes beige .cdsee cr Pearene ieee

Sori non-indusiate; fronds ;10-20 sont oo) Aaisinsfeesmendcnen Doane eee

snap sun vhele cece s Deglsall seve yiteee Ctenopteris moultoni, Nephrolepis radicans

Apex of ‘pinnae ‘roundéd/7), Sarat na aan Nephrolepis radicans

Apex. of. pinnae. pointed: .ii.:<iaianes oaeeke y= dbanenbbite tonnciieasan eens

Pinnae about 5 cm long, strongly falcate ............... Nephrolepis falcata

Pinnae about 8 cm long, weakly falcate ............ Nephrolepis hirsutula

Fronds tripinnatifid or tripinnate, no false veins; sori longer than broad

sbidal aphes . Jalvs SoU ey ae Wes ecessvererseedecverepess aoe

Fronds quadripinnate or more divided, false veins present; sori about as

Wide as J6mg 4,5. Rhos: eee eee Davallia denticulata

port clompate « . j.i:nincmiv eee ean 1-5 colws'to ees o'vislers wecalniabnelen «heen

Sori ‘rounded!’ /,, 211212, .20j0a0ek Le sbleinn shoetGleodhk. AUST IOILU: AenGnnne!

Sori in a longitudinal band halfway between the margin and costa ......

saEON eee gE Sun bie he ve »3'y 05.3 ve'cbG bobs COMER AMAA POREr ee ae Taenitis blechnoides

Sori different | »........cscsceeredseietedesscueeeaddvedsvoth adelante

Malayan Limestone Flora

76.

fee

78.

79.

80.

81.

82.

83.

84.

85.

86.

87.

88.

89.

90.

Sori elongate along and on either side of the midrib ...........................

We0o Ait. Ot. bnidieiwt. lon endeal cv. t2cu. Blechnum _finlaysonianum

Sori, on one or both sides of vein; transverse section of the upper part

of stipe shows vascular strand with two arms (easily seen in fresh

ak a ead ia en ert ce, SER obs ce Ed « cigs anas benes@dsves vee petees

Sori, all on one side of vein; transverse section of the upper part of stipe

aiowe vascular strand: with! four AMIS: -.20i. 52s. k ii... oo. ce ec ceceeccu ens

Fronds simply pmmnate; pinnae entire or lobed ....................s..cccseseces

Fronds more amply divided. Large plant, stipe and frond to 2 m long

LEC SLI SP Pa OF Se aCe ey eee asceeeeseseeeee. Alhyrium esculentum

a, Oe a ae es Se

RS BIER TOQUE a ick. wiry Degen iy Scenes oh hyans xe Athyrium pinnatum

Lateral veins of pinnae forked once, basiscopic veinlet simple or forked

a SNE a OE AS a oe a OL Athyrium montanum

Lateral veins of pinnae in clinic groups, each with 3-4 pairs of veinlets

(LA Snel eg lenge gamain, Hope be opt Ba ede Athyrium prescottianum

Rhizome slender, creeping; fronds at interval ...... Asplenium unilaterale

Rhizome stout, short creeping; fronds tufted ......................0..cce cee

Lower pinnae gradually reduced .................. Asplenium pellucidum

Lower pinnae not reduced ..................... AD CAINS. SOA...

PaO OY 2 CID sacainw stn ndanpectne nyt Pub nh Rg Asplenium adiantoides

ie OA OY GHD oon i acce cen nsh acne pees Asplenium macrophyllum

Sori indusiate; indusia sometimes deciduous ................ccccccccccceceeceeee

Son non-iIndusiate 2... 250.7 00.00. eo a a bs Re, | he a9 ae Ee ie

Fronds bipinnate or more amply divided ................... 0... cccceceeeeeeee es

EOGES SO 2D CIN WHEE oo 5. dain dvi ges los nae0’ Polystichum lindsaeifolium

an tt A WN HCN AS . CAPS ss cis acs cia de Gdaccadecaceac

A tooth is present in the sinus between lobes ......... Pteridrys syrmatica

a ea cue sdglegewaumené

Sori in three irregular rows on either side of the midrib ..................

i Easy een Baty oy oO Caen APY Reon, OP Ce Cyclopeltis crenata

I MLR scence ndvcavacncessscecucscnsutcccccasteres

Ot e@ftaneed On elmer side, Of the. cOSsta.. .......... 00. <rasdececmaneemaracesiles.

Ot atraticed On,either Side of the costule ..............cccccccccascsecccseccs

Sor ‘on- marginal lobes sii! 30. .tv. eee. Nephrolepis dicksonioides

Sori not on marginal lobes ......................0000 Nephrolepis biserrata

205

101

206

91.

92.

oa:

94.

95.

96.

O7.

98.

99.

100.

101.

Gardens’ Bulletin, Singapore — XXX (1977)

Veins of the adjacent lobes of pinnae fused to form an excurrent vein in

the sinus. If this is not distinct then lobes only two-third to the costae

vapuneeb vd ead actacdionscweubbabeesilen Oh dhWivea Ne O4E POM MOONE cUgth ERI An ian ti inn 92

Veins of the adjacent lobes of pinnae not fused, at the most meeting at

the sinus. Lobes almost to the costae ............... Thelypteris immersa

Sori confined to the lobes of the pinnae, not on the lower veins ......... 93

Sori not confined to the lobes of the pinnae, also on the lower veins ...... 94

Pinnae mostly lobed two-third to the costa. Lowest pair of pinnae not

TOUUICG se ravcauees <0 avienasn oats foo tee eae eee ee Cyclosorus extensus

Pinnae lobed one-quarter to one-third to the costa. Lowest pair of pinnae

often Teducea tay auricles 70 eee Cyclosorus interruptus

Pinnae to 1 cm wide; costae densely pubescent beneath .....................

aa scnaceanainomaye 4 gn'dainy Wha ae Sg pein Say Ueto Mla ea Cyclosorus unitus

Pinnae to 3 cm wide; costae minutely pubescent below .....................

catresentsWactstesth goss ieee teen tt eo tee an ame aan name Cyclosorus megaphyllus

Basiscopic lobe of the lowest pair of pinnae enlarged .....................04. 96

Not so. Fronds deeply dissected: $0.0 2005. es .cs. «ses ape atadan eee een 100

All sori on anastomosing VeINS .................0ceee eee Tectaria amplifolia

All or nearly all sori. on.free ‘yeims i460: dens sndlespimsenes heres ea 97

Fronds glabrous on the upper surface, or with some hairs near the

SUMUSES 52, - o mel'e cae g guidisen + ac te eo atonbecun ly snure bald wate c/a age ta a 98

shee aga sie aieladin ae de Dee otaama i aan ane Miaka a Tectaria_ variolosa

Scales stiff, dark purple-brown. Fertile frond not contracted ...............

gh nu o-nee ¢ouegecdll oe ¢ o/c ohielyOil te tins tain aetnane nanan Tectaria griffithii

Veins anastomosing in the costal and costular areoles only (forming a

single row of areoles on either side of the costae and costules) ............

saia's sian apn! wid cle tall» Aa Sanaa ky Ae aida te assesses. Tectaria devexa

Under-surface of lamina glabrous, glabrecent or rarely pubescent .........

OTETOTTT TPE CA hy Tr FEY Pe ei Microlepia_ speluncae

Under-surface of lamina densely pubescent .................c.sceceeeeeeeee ones

ea le ciated se ne Microlepia speluncae var. villosissima

Pinnae not lobed, at most crenate or toothed only ...............cececeeeeees

5 Pees Wide bon 0 Zsiaiv'c b's snes Sn gale loyal ea wees. Abacopteris urophylla

Pinnae deeply lobed, often pinnate at the base; basiscopic lobe of lowest

Pinnne -Calarged 3.0)... .45..ssiasdessesesserduesesselieene Arcypteris irregularis

Malayan Limestone Flora 207

Adiantaceae

Adiantum malesianum Ghatak, Bull. Bot. Surv. Ind. 5: 73. 1963; Holtt., Fl. Mal.

2: 638. 1966.

A. caudatum L., Mant. (1771) 308; Holtt., lc. 599.

Distributed in N.E. India, S. China, Thailand, Indo-China, Sumatra, Sarawak

and Philippines. Common on limestone in Malaya but also found on other rock

types. A fern of moderately shaded places.

Adiantum soboliferum Wall. apud Hk., Spec. Fil. 2: 13. 1851; Holtt., Fl. Mal.

2: 598. 1966.

Distributed throughout the Old World tropics. In Malaya recorded only from

the north and usually on limestone.

Adiantum stenochalamys Bak., Ann Bot. 5 : 29. 1891; Holtt., Fl. Mal. 2 : 602. 1966.

A. opacum Copel., Phil. J. Sc. 1, Suppl. 255, t. 3. 1906.

In Malaya, nearly always found on rocks near the sea. Found inland only on

rocks along the Tahan river and on limestone along Sungei Betis in Kelantan.

Adiantum tenerum Sw., in Hk. et Bk., Syn. Fil. (1868) 124; v.A.v.R., Mal. Ferns

(1908) 330.

A tropical American species now widely cultivated in tropical, subtropical or

even temperate countries. Recorded on limestone as escapes from cultivation; on

shady hill slopes.

Adiantum zollingeri Mett. ex Kuhn, Ann. Mus. Bot. Lugd. Bat. 4: 280. 1869;

Holtt., Fl. Mal. 2: 638. 1966.

A. caudatum L. var. subglabrum Holtt., l.c. 600.

Distributed in Ceylon, S. India, Thailand and Indochina. In Malaya recorded

only from limestone in Kedah and Perlis.

Antrophyum callifolium Bl., Enum. Pl. Jav. (1828) 111; Holtt., Fl. Mal. 2: 605.

1966.

A common forest fern, epiphytic or on rocks. Recorded from limestone in

South Kelantan and Central Pahang.

Antrophyum parvulum BI]. Enum. PI. Jav. (1828) 110; Holtt., Fl. Mal. 2 : 605. 1966.

Distributed in Java. In Malaya this species is widely distributed and common

on limestone, on rocks; with only one record as an epiphyte (Henderson 1939).

It has been recorded from limestone in the south in Johore and apparently absent

from the vast tract of forest separating the Johore from the northern limestone.

It has been collected away from limestone only twice, both on Penang Hill.

Antrophyum semicostatum Bl., Enum. Pl. Jav. (1828) 110; Holtt., Fl. Mal. 2: 605.

1966.

An epiphytic fern of mountain forest. Recorded from low elevation on lime-

stone probably as epiphytes in Kelantan. Resembles A. callifolium but distinguished

by the club-shaped paraphyses.

Cheilanthes farinosa (Forsk.) KLf., Enum. Fil. (1824) 202; Holtt., Fl. Mal. 2 : 592.

1966.

Pteris farinosa Frosk., Fl. Aegypt. Arab. (1775) 187.

Aleuritopteris farinosa Fée, Gen. Fil. (1850-52) 153.

208 Gardens’ Bulletin, Singapore — XXX (1977)

Distributed in most tropical and temperate regions with two records from

Malaya, both from limestone. One was from Bukit Baling in Kedah and the other

from Bukit Chintamani in Pahang the Chintamani specimen was growing in rock

crevices from vertical cliffs.

Doryopteris ludens (Wall.) J. Sm., Hist. Fil. (1875) 289; Holtt., Fl. Mal. 2: 594.

1966.

Pteris ludens Wall. apud Hk., Spec. Fil. 2: 210. 1858.

Distributed from N. India and S. China southwards through Malesia. Restrict-

ed to limestone in Malaya and all collections except two are from Kedah and

Perlis; the exceptions are from Gua Batu, Selangor and Batu Kurau, Perak.

Doryopteris allenae Tryon, Contr. Gray Herb. 91:91, 97. 1962; Holtt., Fl. Mal.

2: 638. 1966.

This species is similar to D. ludens in appearance. It differs however in having

the rhizome short creeping rather than long creeping, the marginal veins in the

sterile lamina mostly jointed rather than free and in not having any hydathodes

on the upper surface of the fronds.

Endemic to limestone in Malaya, recorded from Perak and Selangor. Usually

in shaded localities from rock crevices.

Hemionitis arifolia (Burm.) Moore, Ind. Fil. (1859) 114; Holtt., Fl. Mal. 2 : 596.

1966.

Asplenium arifolium Burm., Fl. Ind. (1768) 213.

Distributed in Ceylon, India, Burma, Indochina, Malesia and the Philippines.

In Malaya only from the north, nearly always from limestone.

Pityrogramma calomelanos (L.) Link, Handb. Gew. 3 : 20. 1833; Holtt., Fl. Mal.

2: 593. 1966.

Acrostichum calomelanos Linn., Sp. Pl. (1753) 1072.

Pantropic, originating in tropical America. Common in Malaya, from the low-

lands to 1,300 m. Frequently one of the early colonisers of open ground. Recorded

once from the base of limestone.

Taenitis blechnoides (Willd.) Sw., Syn. Fil. (1806) 24, 220; Bedd., Handb. 410, t.

242; Holtt., Fl. Mal. 2: 586. 1966.

Pteris blechnoides Willd., Phytogr. (1794) 13.

Vittaria angustifolia Bl., Ennum. Pl. Jav. (1828) 199; Holtt., Fl. Mal. 2: 610. 1966.

Distributed throughout Malesia. In Malaya, Sumatra, Borneo and Java this

is a mountain epiphyte at 600-1800 m. On limestone in Malaya this has been

collected as a low epiphyte from 20-300 m; recorded from Kedah, Kelantan and

Perlis; uncommon.

Vittaria elongata Sw., Syn. Fil. (1806) 302; Holtt., Fl. Mal. 2: 614, 1966. var.

angustifolia Holtt., in l.c.

This variety is endemic to Malaya and found only in the North-west and on

Pulau Tioman in the South-east. It is both an epiphyte and rock plant and

recorded from only one limestone locality as an epiphyte. The typical variety is a

common lowland and mountain epiphyte with a paleotropical distribution.

Malayan Limestone Flora 209

Dennstaedtiaceae

Arcypteris irregularis (Pr.) Holtt., Reinw. 1: 193. 1951; Fl. Mal. 2: 538. 1966.

Polypodium irregulare Pr., Rel. Haenk. 1 : 25. 1825.

A common forest species in Malaya. Recorded once from the small limestone

outcrop in deep forest in Johore.

Asplenium adiantoides (L.) C. Chr., Ind. Fil. 1905; Holtt., Fl. Mal. 2 : 431. 1966.

Trichomanes adiantoides L., Sp. Pl. (1753) 1098.

Distributed in Burma, Thailand and southwards to Malesia, Australia and

Polynesia. Also in Madagascar.

Resembles A. macrophyllum Sw. but generally smaller. Intermediates occur

but the extreme forms are very different. More field work and cultivation of this

species will help to clarify the growth form of this species. Presently they are best

kept apart.

This is a fern of rocky places and nearly always from limestone. It is very

common on scrubby summits of some hills around Gua Musang in Kelantan.

Asplenium macrophyllum Sw., Schrad. Journ. 1800/2: 52. 1801; Holtt., Fl. Mal.

2: 431. 1966.

Found in most parts of Malaya on rocks and as epiphytes. It is common on

coastal areas in the east coast and on limestone in Selangor.

Asplenium pellucidum Lam., Encyl. 2 : 305. 1786; Holtt., Fl. Mal. 2 : 428. 1966.

Common in Malaya from the lowlands to the hills, on rocks or as epiphytes:

often on limestone.

Asplenium phyllitidis Don, Prodr. Fl. Nep. (1825) 7; Holtt., Fl. Mal. 2 : 420. 1966.

Common in Malaya, usually epiphytic. Recorded from limestone in Kelantan.

Asplenium salignum Bl., Enum. Pl. Jav. (1828) 175; Holtt., Fl. Mal. 2 : 421. 1966.

A. filiceps Copel., Philip. J. Sc. Sc : 285. 1910.

Common in Malaya, usually as an epiphyte, on the lowands, hills and moun-

tains. Fairly common on limestone under partial shade as a low epiphyte or on

rocks.

Asplenium squamulatum BI., Enum. Pl. Jav. (1828) 174; Holtt., Fl. Mal. 2 : 426.

1966.

Distributed throughout Malesia. Recorded in Malaya from Pahang, Kelantan

and Selangor, usually on limestone.

Asplenium unilaterale Lam., Encys. 2 : 305. 1786; Holtt., Fl. Mal. 2 : 438. 1966.

Widely distributed in Malaya, on rocks in moist shady places. Uncommon on

limestone.

Athyrium cordifolium (Bl.) Copel., Philip. J. Sc. 3c: 300. 1908; Holtt., Fl. Mal.

2: 548. 1966.

Diplazium cordifolium Bl., Enum. Pl. Jav. (1828) 190.

Common in lowland and mountain forest in Malaya. Recorded only once from

limestone.

Athyrium esculentum (Retz.) Copel., Philip. J. Sc. 3c : 295. 1908; Holtt., Fl, Mal.

2: 562. 1966.

Hemionitis esculenta Retz., Obs. Bot. (1791) 38.

Common in wet places in the lowlands of Malaya. Recorded once from lime-

stone at the base of hill. 3

210 Gardens’ Bulletin, Singapore — XXX (1977)

Athyrium montanum (v.A.v.R.) Holtt., Fl. Mal. 2: 555. 1966.

Diplazium montanum v.A.V.R., Bull. Jard. Bot. Ser. If, 28: 19. 1918.

Common in Malaya in lowland and hill forest. Recorded from limestone in

Johore only.

Athyrium pinnatum (Blanco) Copel., Philip. J. Sc. 3c : 297. 1908; Holtt., Fl. Mal.

2: 560. 1966.

Allantodia pinnata Blanco, Fl. Filip. Ed. 2 (1845) 571.

Widespread in Malaya with one record from limestone.

Athyrium prescottianum (Wall.) Holtt., Fl. Mal. 2: 557. 1966.

Asplenium prescottianum Wall., Cat. (1829) 235 (nom. nud.)

Endemic, an uncommon species. Recorded once as an epiphyte from limestone.

Blechnum finlaysonianum Hk. et Grev., Ic. Fil. (1831) t. 225; Holtt., Fl. Mal.

2: 445. 1966.

A common species in lowland and hill forest, once recorded from limestone.

Cyclopeltis crenata (Fée) C. Chr., Ind. Fil. Suppl. 3 : 64. 1934; Holtt., Fl. Mal.

233527, 1966,

Hemicardion crenatum Fée, Gen. Fil. (1852) 283, t. 22A.

Distributed in Burma, S. China and western Malesia. This is a plant of rocky

places, usually but not always from limestone, in Malaya. Widely collected from

limestone, in part shade.

Davallia denticulata (Burm.) Mett., in Kuhn., Fil. Deck. (1867) 27; Holtt., Fl. Mal.

22309. 1966.

A common rock plant and epiphyte; recorded a number of times from lime-

stone.

Davallia solida (Frost.) Sw., Schrad. Journ. 1800/2: 87. 1801; Holtt., Fl. Mal.

2: 360. 1966.

Trichomanes solidum Forst., Prodr. (1786) 86.

A common coastal epiphyte and rock plant in Malaya. Uncommon inland.

Recorded only once on limestone; this was from a large local population on the

summit of Batu Tapah, growing in part shade on rocks and as a low epiphyte. It

is probably not uncommon on limestone, on such dry, rocky, scrubby summits.

Heterogonium alderwereltii Holtt., Sarawak Mus. J. 5 : 163. 1949, Fl. Mal. 2 : 522.

1966.

Pleocnemia membranifolia p.p. quoad Bedd., Handb. Suppl. (1892) 48.

Plant like that of H. pinnatum. The typical specimen however has larger

sterile fronds with 4~7 pairs of pinnae. The sori is not acrostichoid but is either

distinct or elongate along veins. The indusia is distinct though less prominent in

mature sori.

Distributed in Sumatra. Restricted to limestone in Malaya (except for a

specimen from Patani, Kedah); not as common as H. pinnatum.

For notes see under H. pinnatum.

Heterogonium pinnatum (Copel.) Holtt., Sarawak. Mus. J. 5: 163. 1949, Fl. Mal. |

2: 524. 1966.

Stenosemia pinnata Copel., Phil. J. Sc. 1, Suppl. 2 : 48. 1892.

Pleocnemia membranifolia p.p. quoad Bedd. Handb. Suppl. (1892) 48.

___ Distributed in Sumatra, Borneo and the Philippines. Restricted to limestone

in Malaya, and the commonest fern on limestone.

Malayan Limestone Flora 211

Typically this fern is distinguished from H. alderwereltii Holtt. mainly by the

absence of indusia and the sterle fronds having only one instead of four to seven

pairs of free lateral pinnae. H. alderwereltii has distinct sori or extended sori along

the veins while typical H. pinnatum has acrostichoid sori, and if atypical, the sori

extend along veins but still without indusia.

However, there is great variation in the morphology of these two species, thus

delimited. There are H. pinnatum plants with up to 7 pairs of free pinnae (in the

sterile fronds) and which have, in the fertile fronds, an acrostichoid condition in

which the sori are non-indusiate. The sterile lamina with up to 7 pairs of free

pinnae would seem to belong to H. alderwereltii while the acrostichoid non-

indusiate sori would seem to belong to H. pinnatum. Apart from the extreme forms

of H. pinnatum in which the sterile fronds have normally one pair of free pinnae,

the form and size of the sterile fronds are insufficient characters to separate these

two species.

There are identical plants in which the sori are distinct or elongate sub-

marginally. In some instances they are indusiate and in others they are not. Once

these two species are kept distinct, one can say that the non-indusiate plants would

be abnormal H. pinnatum (the normal plants have acrostichoid sori). However,

this argument would be equally valid if one says that the non-indusiate condition

is found on abnormal H. alderwereltii which have lost their indusia. In both

indusiate and non-indusiate forms there often exists a condition in which sori are

found on fronds with the size and form of a sterile frond. These sori are either

very distinct on vein endings or often form submarginal bands.

This range of variation can be found in a single population such as that on the

base of Bukit Takun, Selangor, growing on limestone and organic debris. Only

the extreme forms would seem to keep these two species apart and I suspect (as

Beddome thought) they are variations of one species. This is as much as I can

conclude from field observations; experimental culture from spores is required.

Besides these variations there are indusiate as well as non-indusiate forms

which are free veined, a character now found in H. sagenoides (Mett.) Holtt. Also,

the form in which the fertile fronds have a broad lamina and distinct sori

encroaches on one of the characters of the remaining species of Heterogonium

found in Malaya, H. saxicolum (Bl.) Holtt. Other characters, however, seem to

keep H. pinnatum and H. alderwereltii distinct from these latter two species. The

former two are also different in being exclusively limestone plants in Malaya with

an exception of one specimen of H. alderwereltii collected from Patani, Kedah.

In the light of recent collections, the taxonomic status of this complex of

Heterogonium needs looking into, especially the limestone-inhabiting groups.

For convenience, the two species are retained here, often distinguished only

by the presence or absence of an indusium. H. pinnatum is the more commonly

collected species, appearing on most limestone outcrops except in the extreme

north and Kelantan. Both are found in shady, rocky places and most luxuriently

in situations with humus accumulation.

It is interesting to note that H. pinnatum has been collected from the very small

and isolated Johore limestone, forming a link with the more southern Sarawak

populations.

Humata heterophylla (Sm.) Desv., Prodr. (1825) 323; Holtt., Fl. Mal. 2 : 366. 1966.

Davillia heterophylla Sm., Mem. Ac. Turin. 5: 415. 1793.

A widely distributed species in Malaya usually in exposed places, on rocks

or as epiphytes. Recorded on limestone from Kelantan.

212 Gardens’ Bulletin, Singapore — XXX (1977)

Humata pectinata (Sm.) Desv., Prodr. (1827) 323; Alston, Phil. J. Sc. 50: 175

1933; Holtt., Fl. Mal. 2 : 369. 1966.

Davallia pectinata Sm., Mem. Ac. Turin. 5 : 415. 1793.

Distributed from Sumatra to New Guinea, usually on coastal rocks and trees.

Inland collections in Malaya are restricted to limestone, growing as low epiphytes

or on rocks in shade or part shade. Abundant on the rocky, scrubby summit of

Batu Tapah.

Lindsaya lucida Bl., Enum. Pl. Jav. (1828) 216; Holtt., Gard. Bull. S.S. 9: 131.

1937, Fl. Mal. 2: 328. 1966.

L. lobbiana Hk. Sp. Fil. 1: 205. 1848; C. Chr., Gard. Bull. S.S, 4: 396,

1929; Holtt., Gard. Bull. S.S. 5:61. 1930.

In Malaya, a streamside plant of low country, widespread. Recorded once

from the base of a limestone hill growing on limestone boulders by a stream.

Microlepia speluncae (L.) Moore, var. villossima C. Chr., Gard. Bull. S.S. 4 : 399.

1929; Holtt., Fl. Mal. 2: 314. 1966.

Nephrolepis biserrata (Sw.) Schott, Gen. Fil. (1834) t. 3; Holtt., Fl. Mal. 2 : 380.

1966.

Aspidium biserratum Sw., Schrad. Jour. 1800/2 : 32. 1801.

A very common fern of open or partly shaded places. Recorded once from

limestone (on Gua Musang) as a secondary element about a year and a half after

fire destroyed the original vegetation.

Nephrolepis dicksonioides Chr., Verh. Nat. Ges. Basel 11 : 241. 1895; Holtt., Fl.

Mal. 2 : 376. 1966.

Distributed in Celebes, Borneo and New Guinea. In Malaya it is restricted to

limestone, on exposed or partly exposed situations. Common on many hills and

often forming dense thickets.

Nephrolepis falcata (Cav.) C.Chr., Dansk. Bot. Ark. 9: 15. 1937; Holtt., Fl. Mal.

2:381. 1966.

Tectaria are Cav., Descr. (1802) 250.

N. barbata Copel., in Holtt., Gard. Bull. S.S. 9 : 132. 1937.

A common fern on rocky places and as epiphytes. Sometimes on limestone.

Nephrolepis hirsutula (Forst.) Pr., Tent. Pterid. (1836) 79; Holtt., Fl. Mal. 2 : 382.

1966.

Polypodium hirsutulum Forst., Prodr. (1786) 81.

Common in Malaya in open places. Recorded once from the disturbed base

of limestone in secondary scrub vegetation.

Nephrolepis radicans (Burm.) Kuhn, Ann. Lugd. Bat. 4: 285. 1869; Holtt., FI.

Mal. 2 : 381. 1966.

Polypodium radicans Burm., FI. Ind. (1768) 233, t. 66.

Oleandra undulata (Willd.) Ching, Lingnan Sc. Jour. 12: 565. 1933; Holtt., FI.

Mal. 2 : 384. 1966.

Polypodium undulatum Willd., Sp. Pl. 5 : 155. 1810.

A mountain fern, common in Malaya above 1000 m., often on acid peat and

humus. An unlikely species to be found on limestone. However, one specimen

(Kadim K491) positively of this, is labelled as from Gunong Tempurong, in forest

Malayan Limestone Flora 213

at 250-550 m. The main range of mountains which rises abruptly to over 650 m

just over a mile to the east of the Gunong Tempurong massive could provide the

source for the spread of this species. The plant was probably growing on an

accumulation of humus over the limestone.

Polystichum lindsaeifolium Ridl., J. Mal. Br. R. As. Soc. 4:61 1926; C. Chr.,

Gard. Bull. S.S. 4 : 393. 1929; Holtt., Fl. Mal. 2: 489. 1966.

Endemic to limestone in Selangor, Pahang, Kelantan and Perak, uncommon.

Pteridium aquilinum (L.) Kuhn, var. wightianum (Ag.) Tryon, Rhodora 43 : 1-70.

1941; Holtt., Fl. Mal. 2 : 634. 1966.

Pteris aquilina L., Sp. Pl. (1753) 1075.

Pteridium aquilinum (L.) Kuhn, in Holltt., lc. 389.

Recorded from limestone as a secondary element.

Pteridium caudatum (L.) Maxon var. yarrabense Domin, in Tryon, Rhodora

43 : 63. 1941; Holtt., Fl. Mal. 2: 634. 1966.

Pteris esculenta Forst., Pl. Escul. (1786) 74.

Pteridium esculentum (Forst.) Nakai, Bot. Mag. Tokyo 39: 109. 1925; Holtt.,

lc. 390.

Recorded on limestone as an element of secondary vegetation.

Pteridrys syrmatica (Willd.) C. Chr. et Ching, Bull. Fan Mem. Inst. Bot. 5: 131.

1934; Holtt., Fl. Mal. 2 : 530. 1966.

Aspidium syrmaticum Willd., Sp. Pl. 5 : 277. 1810.

Distributed in Ceylon, Thailand, Indochina and Malesia to the Philippines.

In Malaya, this fern is found mainly in the north and nearly always from limestone.

It is a rock fern of shady places.

Pteris ensiformis Burm., Fl. Ind. (1768) 230. Holtt., Fl., 2 : 399. 1966.

Sometimes on limestone.

Pteris longipinnula Wall., Cat. (1829) 108; Holtt., Fl. Mal. 2: 404. 1966. var.

b, in Holtt., l.c. 405.

This variety is endemic to Malaya and chiefly from limestone, though not

exclusively so. The typical variety is distributed from India to China and south-

wards throughout Malesia. It differs from the typical variety by the narrower

pinnae and by the presence of the branched basal pair of pinnae.

Pteris mertensioides Willd., Sp. Pl. 5: 394. 1810; Holtt., Fl. 2: 404. 1966.

Pteris scabripes Wall. apud Hook., Sp. Fil. 2: 165. 1858; Holtt., Fl. Mal. 2: 399.

1966.

Endemic, usually on rocks by streams, also in lowland forest and rarely on

limestone, not a common species.

Pteris tripartita Sw., Sshrad. Jour. 1800/2 : 67. 1801; Holtt., Fl. Mal. 2: 408 1966.

P. marginata Bory, Voy. 2 : 192. 1804.

Pteris vittata L., Sp. Pl. (1753) 1074; Hoiltt., Fl. Mal. 2 : 396. 1966.

Not uncommon on disturbed localities on limestone, by quarry edges and on

disturbed summits.

214 Gardens’ Bulletin, Singapore — XXX (1977)

Tectaria amplifolia (v.A.v.R) C. Chr., Ind. Fil. Suppl. 3: 176. 1934; Holtt., FA.

Mal. 2: 515. 1966.

Aspidium amplifolium v.A.v.R., Bull. Jard. Bot. Buit. Ser. II, 11:2. 1913.

Distributed in Sumatra. Common on and restricted to limestone in Malaya.

Apparently unknown from the extreme North-west. A plant of shady places.

Tectaria barberi (Hk.) Copel., Philip. J. Sc. 2c : 414. 1907; Holtt., Fl, Mal. 2 : 508.

1966.

Polypodium barberi Hk., Sp. Fil. 5 : 100. 1864.

Tectaria devexa (Kze) Copel., Philip. J. Sc. 2c: 415. 1907; Holtt., Fl. Mal. 2 : 505.

1966.

Aspidium devexum Kze, Bot. Zeit. (1848) 259.

A. membranaceum Hk., Sp. Fil. 5: 105. 1864.

Distributed in Ceylon, Thailand and S. China. Restricted to limestone in

Malaya, widely distributed and fairly common.

Tectaria griffithii (Bak.) C.Chr., Ind. Fil. (1867) 300; Holtt., Fl. Mal. 2 : 636 1966.

Nephrodium griffithit Bak., Syn. Fil. (1867) 300.

Tectaria multicaudata Ching, Sinensia 2: 20. 1931; Holtt., F. Mal. 2: 507.

1966.

T. malayense Christ, Philip. J. Sc. 2c : 187. 1907.

Tectaria macrodonta (Fee) C. Chr., Ind. Fil. Suppl. 3 : 181. 1934; Holtt., Fl. Mal.

2: 505. 1966.

Sagenia macrodonta Fée, Gen. Fil. (1852) 313 t. 24.

A number of specimens collected from north Malaya are doubtfully identified

as this species. One of these is from limestone. They are smaller than the typical

form of this species which are represented in India. They resemble T. variolosa

but the fronds are hairy on both surfaces and hardly dimorphic, and the sori are

not all on free veins.

Tectaria variolosa (Wall.) C. Chr., Contr. U.S. Nat. Herb. 26: 289. 1931; Holtt.,

Fl. Mal. 2: 506. 1966.

Aspidium variolosum Wall., Hook., Spec. Fil. 4: 51. 1862.

Grammitidaceae

Ctenopteris alata (Bl.) Holtt., Fl. Mal. 2 : 232. 1966.

Davallia alata Bl., Enum. Pl. Jav. (1828) 230.

Prosaptia alata Christ, Ann. Buit. II, 5 : 127. 1905.

Ctenopteris moultoni (Copel.) C. Chr. et Tard. Not. Syst. 8: 181. 1939; Holtt., Fi.

Mal. 2 : 229. 1966.

Polypodium moultoni Copel., Philip. J. Sc. 10c : 149. 1915.

Hymenophyllaceae

Trichomanes bipunctatum Poir., in Lamk., Encyl. 8: 69. 1808; Holtt., Fl. Mal.

Zz: 99. 1966.

Crepidomanes bipunctatum Copel., Phil. J. Sc. 67: 59. 1938.

The only common species of this family (except in the very north) on limestone.

Malayan Limestone Flora 215

Trichomanes christii Copel., Phil. J. Sc. 1. Suppl. 251. 1906, ibid., 51 : 185. 1933;

Holtt., Fl. Mal. 2: 100. 1966.

Crepidomanes christii Copel., Phil. J. Sc. 67 : 60. 1938.

Trichomanes humile Forst., Prodr., (1786) 84. Copel., Phil. J. Sc. 51: 164. 1933;

Holtt., Fl. Mal. 2: 98. 1966.

Crepidopteris humilis Copel., Phil. J. Sc. 67 : 58. 1938.

Trichomanes motleyi Bosch, Ned. Kruidk. Arch. 5: 145. 1861; Copel., Phil. J. Sc.

51:201. 1933; Holtt., Fl. Mal. 2:92. 1966.

Polypodiaceae

Crypsinus enervis (Cav.) Copel., Gen. Fil. (1947) 207; Holtt., Fl. Mal. 2: 199.

1966.

Polypodium enervis Cav., Descr. (1802) 245.

Drynaria bonii Christ., Not. Syst. 1 : 186. 1910; Tardieu-Blot et Christ., Fl. Gen.

Indochine 7 : 515. 1941.

Rhizome creeping. Nest leaves small, ovate, base cordate-sagittate, margin

entire scalloped or shallowly lobed, 5-10 by 3.5—6.5 cm. Foliage leaves with stipes

8-20 cm long, narrowly winged; lamina deeply lobed, 20-45 by 12-20 cm. Lobes

3-7 pairs, 8-20 by 2-4 cm, base decurrent along the rachis. Sori non-indusiate and

small, scattered on the lower surface of frond.

Distributed in S. China, Indochina and Thailand. A new record (Chin 1764)

for Malaya. Common as low epiphytes and on rocks in Pulau Langgun, Langkawi

on the northern side of the island.

Characterised by the small unlobed nest leaves and the dissection of the

foliage leaves. The old rhizome which is fleshy and flattened grows up to 3 by

1.5 cm.

Drynaria quercifolia (L.) J. Sm., J. Bot. 3: 398. 1841; Holtt., Fl. Mal. 2: 182.

1966.

Polypodium quercifolium Linn., Sp. Pl. (1753) 1087.

Drynaria rigidula (Sw.) Bedd., Ferns Brit. Ind. (1869) t. 314; Holtt., Fl. Mal.

2: 183. 1966. :

Polypodium rigidulum Sw., Schrad. J. 1800/2 : 26. 1801.

A common epiphyte in the northern half of Malaya; it is not found in the

south. It is not uncommon on limestone in Langkawi and has been recorded as

far south as Bukit Takun in Selangor. This forms the southernmost record for this

Poe in Malaya. On limestone it has been recorded as epiphytes as well as on

rocks.

eta sparsisora (Desv.) Moore, Ind. Fil. (1862) 348; Holtt., Fl. Mal. 2: 183.

Polypodium sparsisorum Desv., Berl. Mag. 5 : 315. 1811.

Lemmaphyllum accedens (Bl.) Donk. Reinw. 2 : 409. 1954; Holtt., Fl. Mal. 2: 152.

1966.

Polypodium accedens Bl., Enum. Pl. Jav. (1828) 121.

Weatherbya accedens Copel., Gen. Fil. (1947) 191.

Lepisorus longifolius (Bl.) Holtt., Fl. Mal. 2: 151. 1966.

Grammitis longifolia Bl., Enum. Pl. Jav. (1828) 119.

Paragramma longifolia Moore, Copel., Gen. Fil. (1947) 190.

216 Gardens’ Bulletin, Singapore — XXX (1977)

Leptochilus decurrens Bl., Enum. Pl. Jav. (1828) 206; Holtt., FI. va 2: 164 1966.

Acrostichum variabile Hook, Sp. Fil. 5 : 277. 1864.

Perper avenia (BIl.) Presl., Tent. Pterid. (1836) 215; Holtt., Fl. Mal. 2 : 167.

Grammitis avenia Bl., Enum. Pl. Jav. (1828) 117.

Loxogramme blumeana Presl., Tent. Pterid. (1836) 215.

Not common on limestone, but recently found to be abundant on the rocky,

one-layered scrub forest on the summit of Gua Batu Boh, near Gua Musang,

Kelantan; on rocks and as low epiphytes.

Loxogramme scolopendrina (Bory) Presl., Tent. Pterid. (1836) 215; Holtt., Fl. Mal.

2: 168. 1966.

Grammitis scolopendrina Bory, Dup. Voy. (1829) 257.

Fairly common on limestone and widely distributed.

Microsorium musifolium (Bl.) Ching, Bull. Fan Mem. Inst. 4: 295. 1933; Holtt.,

Fl. Mal. 2: 176. 1966.

Polypodium musifolium Bl., Enum. Pl. Jav. (1828) 134.

Microsorium punctatum (L.) Copel., Univ. Cal. Publ. Bot. 16: 111. 1929; Holtt.,

Fl. Mal. 2: 179. 1966.

Acrostichum punctatum Linn., Sp. Pl. Ed. 2. (1763) 1524.

Photinopteris speciosa (BL.) Pres]., Epim. Bot. (1849) 264; Holtt., Fl: Mal. 2 : 187.

1966. |

Lomaria speciosa Bl\., Enum. Pl. Jav. (1828) 202.

Photinopteris rigida Bedd., Fl. Brit. Ind. (1867) t. 211.

Phymatodes nigrescens (Bl.) J. Sm., Ferns Br. & For. (1866) 94; Holtt., Fl. Mal.

2: 193. 1966.

Polypodium nigrescens P\., Enum. Pl. Jav. (1828) 126.

Phymatodes scolopendria (Burm.) Ching, Contr. Inst. Bot. Nat. Acad. Peip. 2 : 63.

1933; Holtt., Fl. Mal. 2: 191. 1966.

Polypodium scolopendria Burm., Fl. Ind. (1769) 232.

P. phymatodes Linn., Mant. (1771) 306.

Recorded a number of times from limestone, ahatiddte on Batu Tapah.

Kelantan, on the scrubby summit growing over boulders and as low epiphytes.

Polypodium papillosum Bl., Enum. Pl. Jav. (1828) 131; Holtt., Fl. Mal. 2: 203.

1966.

Pyrrosia adnascens (Sw.) Ching, Bull. Chin. Bot. Soc. 1 : 45. 1935; Holtt., Fl. Mal.

2: 144. 1966. 7

Polypodium adnascens Sw., Syn. Fil. (1806) 25.

Common in the lowlands of Malaya, epiphytic pres on Tocks; also common on

limestone and widely collected.

Pyrrosia floccigera (Bl.) Ciuag, Bull. Chin. Bot. Soc. 1: 71. 1935; Holtt., Fl. Mal.

2: 147. 1966.

Niphobolus flocciger Bl., Enum. Pl. Jav. (1828) 107.

Malayan Limestone Flora 217

Fairly common in Malaya, usually from 300-1300 m, in partly shaded areas,

epiphytic. Recorded once from limestone at less than 100 m elevation (UNESCO

1962, 216, from Gua Batu Boh, Kelantan).

Pyrrosia penangiana (Hook.) Holtt., Fl. Mal. 2: 146. 1966.

Niphobolus penangianus Hook., Ic. Pl. (1840) t. 203.

Distributed in Sumatra. Recorded from the northern half of Malaya, usually

on limestone; on rocks or as epiphytes.

Pyrrosia stigmosa (Sw.) Ching, Bull. Chin. Bot. Soc. 1:67. 1935; Holtt., Fl. Mal.

2: 148. 1966.

Polypodium stigmosum Sw., Schrad. Jour. 1800/2: 21. 1801.

Distributed in North India and Indochina and from Sumatra to New Guinea.

Restricted to limestone in Malaya, on rocks and epiphytic; it has been reported

as an epiphyte on Rain trees near Gua Batu, Selangor.

Pyrrosia varia (Kaulf.) Farwell, Am. Midl. Nat. 12 : 302. 1931; Holtt., Fl. Mal.

2: 146. 1966.

Niphobolus varius Kaulf., Enum. Fil. (1824) 125.

A widely distributed species, also common on limestone.

Schizaeaceae

Lygodium flexuosum (L.) Sw., Schrad. Jour. 1800/2: 106. 1801; Holtt.. Fl. Mal.

Zt. 1966.

Ophioglossum flexuosum Linn., Sp. Pl. (1753) 1063.

Lygodium polystachyum Wall. ex Moore, Gard. Chron. (1859) 671; Holtt., Fl. Mal.

2:56. 1966.

Distributed in Burma and Thailand. Uncommon and apart from specimens

from Penang Hill (granite), this species is restricted to limestone in Malaya.

Recorded from Perak, Kelantan, Langkawi and Pahang, usually in shady forest.

Schizaea inopinata Selling, Sevensk Bot. Tidskr. 40 : 274. 1946; Holtt., Fl. Mal.

2252. 1966.

Distributed in Sumatra and Philippines. In Malaya, this species is restricted

to limestone in Kelantan, Pahang and Selangor, not uncommon. It resembles the

common S. digitata (L.) Sw. and is frequently mistaken for it. S. inopinata is how-

ever characterized by having the sporangia in two instead of four rows.

Thelypteridaceae

Abacopteris urophylla (Wall.) Ching, Bull. Fan Mem. Inst. Bot. 8: 251. 1938;

Holtt., Fl. Mal. 2: 296 1966.

Polypodium urophyllum Wall., in Hook., Sp. Fil. 5: 9. 1863.

Cyclosorus extensus (Bl.) Ching, Bull. Fan Mem. Inst. Bot. 8 : 182. 7938: Holtt.,

Fl. Mal. 2: 264. 1966.

Aspidium extensum Bl., Enum. Pl. Jav. (1828) 156.

Cyclosorus interruptus (Willd.) Ching, Bull. Fan Mem. Inst. Bot. 8 : 184. 1938;

Holtt., Fl. Mal. 2 : 262. 1966.

Pteris interrupta Willd., Phytogr. 1 : 13. 1794.

218 Gardens’ Bulletin, Singapore — XXX (1977)

Cyclosorus megaphyllus (Mett.) Ching, Bull. Fan Mem. Inst. Bot. 8: 225. 1938;

Holtt., Fl. Mal. 2: 268. 1966.

Aspidium megaphyllum Mett., Ann. Mus. Lugd. Bat. 1 : 233. 1864.

Cyclosorus unitus (L.) Ching, Bull. Fan Mem. Inst. Bot. 8: 192. 1938; Holtt., FI.

Mal. 2: 260. 1966.

Polypodium unitum L., Syst. Nat. ed. 10, 2 : 1326. 1759.

Thelypteris immersa (Bl.) Ching, Bull. Fan Mem. Inst. Bot. 6 : 306. 1936; Holtt.,

FL 2; 245.. 1906.

Aspidium immersum Bl., Enum. (1828) 156.

GYMNOSPERMS

Araucariaceae

Agathis dammara (Lambert) L. G. Rich., Comm. Bot. Conif. Cycad. (1826) 83:

Keng, Tree Fl. Mal. 1:41. 1972.

A. loranthifolia Salisb. in Ridl., Flora 5 : 278. 1925.

A. alba (Lam.) Jeff. in Burk., Dict. I: 62. 1935. Corner, Ways. Trees I: 715.

1940.

Usually found on the hills; once recorded (Loh FRI 17201) for limestone

at 600-700 m at Gua Peringat, Pahang. This was growing in deep soil in a gully.

Cycadaceae

Leaves with petioles about 100 cm long, leaflets 0.7-0.9 cm wide. Male cone

about 20 cm) long >) 407-24.:.c0d dodge aes lal aren Cycas siamensis

Leaves with petioles about 200 cm long, leaflets 1.3-2.0 cm wide. Male cone

about 35 cm lome yi peccdskpcectabn anee ee tk eee aaa on Saeed Cycas rumphii

Cycas rumphii Mig., Bull. Sc. Phys. Neerl. 839, 45; Hk. f., F.B.I. 5: 657; Ridl..

Fl. 5 : 284. 1925.

Cycas siamensis Mig., Bot Zeit. (1863) 334; Hk. f., F.B.I. 5 : 657; Ridl. Fl. 5 : 285.

1925.

Distributed in Burma and Thailand; in Malaya, restricted to limestone, and

common on Langkawi islands.

Gnetaceae

A shrub or small tree, to 3 m tall; flowers not embedded in thick hair masses.

ee et ee eee Pe oe ee Gnetum gnemon var. tenerum

A liana, more than 10 m long; flowers embedded in thick hair masses. ...............

a vos Gnd cs'ovwavan dulro0 0 bis apmpondin anc pl a ad i eepelccadio i a ne Gnetum cuspidatum

Gnetum cuspidatum Bl., Rumphia 4:5. 1848; Henders., J. Mal. Br. As. Soc. —

17 : 87. 1939; Markgraf, Fl. Mal. I, 4: 343. 1951.

G. penangense Ridl., Fl. 5 : 276. 1925.

Gnetum gnemon Linn., Mant. 1: 125. 1767; Markgraf, Bull. Jard. Bot. Btzg II,

(10) : 436. 1930; Rid]. Fl. 5:273. 1925; Markgraf, Fl. Mal. I, 4: 340. 1951.

var. tenerum Markgraf, Fl. Mal. I, 4: 341. 1951.

Malayan Limestone Flora 219

Podocarpaceae

i A en Se Se ee ee ee Podocar pus polystachyus

Male cones solitary or sometimes 2-3 together. Leaves usually larger, 5.5—20

i I Ee on eee A ae OA Ae Podocar pus _neriifolius

Podocar pus polystachyus R. Br. [in Mirb., Mem. Mus. Hist. nat. Paris 13:47. 1825

nom. nud.] ex Endl., syn. Conif. (1847) 215. Ridl., Fl. 5: 282. 1925; Keng,

Ttee Fl, Mal, 1: 53: 1972.

Usually coastal except on limestone hills where it is very widely distributed

and common.

Podocar pus neritifolius D. Don, in Lambert, Desc. Gen. Pinus 1 : 21. 1824; Ridl., FI.

5 : 281. 1825; Keng, Tree Fl. Mal. 1 : 51. 1972.

Usually on hill and mountain forest, rarely below 300 m. Recorded on lime-

stone from Gua Batu Selangor. (Ng FRI. 1634).

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A List of Holttum’s New Taxa and Name-Changes in Ferns to

July 1975

J. A. CRABBE

British Museum (Natural History)

London

Little did I think, when I embarked on this abstracting, that I would be

delving among some 420 titles by R. E. Holttum which range over botany,

horticulture, philosophy, etc. (see Flora Malesiana Bulletin 28: 1975), and that

IT would find around 700 items for this present list.

I have not included anything published after his 80th birthday on 20 July

1975, nor some few names superfluously replicated, nor some invalidly published

(usually without latin diagnoses and usually rectified later). The genus Polystic-

hopsis (17: 149, 152) is, to me, obscure and I suspect there may be other items in

this work and also in a similar treatment (The classification of ferns, in Biol.

Rev. 24: 267-296. 1949).

It is with the greatest pleasure that I present this list, acknowledging with

gratitude much helpful discussion with colleagues, and trusting it will serve a useful

purpose.

For brevity, I have numbered the bibliography, then quoted those numbers

followed by page numbers, New species give the type and its location, a semicolon

separating holotype from isotypes. Where Holttum designates no type but lists

specimens he has seen, I have quoted the first in that list, not thereby implying

nid other status. Basionym authors followed by their genus are enclosed in round

rackets.

Abacopteris menisciicarpa (Bl. — Aspidium) 24:290

A, peltochlamys (C. Chr. — Dryopteris) 24:295

A, rubicunda (v.A.v.R. — Phegopteris) 24:292

A. salicifolia (Wall. ex Hook. — Meniscium) 24:288

Alsophila parvifolia 2:19. — Mentawi Is., Kloss 14579 (SING)

Amauropelta bergiana (Schlechtend. — Polypodium) 57:133

. bergiana var, calva 57:133.— Cameroon Mt, Richards 4308 (K)

. bergiana var. tristanensis 57:134. — Tristan da Cunha, Carmichael (K)

. heteroptera (Desv. — Nephrodium) 57:132

. membranifera (C, Chr. — Dryopteris) 57:132

. oppositiformis (C. Chr. — Dryopteris) 57:135

. Strigosa (Willd. — Aspidium) 57:134

. tomentosa (Thouars — Polypodium) 57: 134

AMPHINEURON gen. nov. 50:45. figs 19, 19a. — type species: A. opulentum

A. immersum (Blume — Aspidium) 62:203

A. opulentum (Kaulf.— Aspidium) 50:45

A. terminans (Hook: — Nephrodium) 54:82

>Prr>rrprpyp

221

222 Gardens’ Bulletin, Singapore — XXX (1977)

Araiostegia membranulosa (Wall. ex Hook. — Davallia) 52:230

Arcypteris brongniartii (Bory — Polypodium) 21:195

A. gigantea (Cesati— Nephrodium) 21:195

A. irregularis (Presl — Polypodium) 21:193. ff. 1-3

A. macrodonta (Fée — Dictyopteris) 21:194

Arthropteris wollastonii (Ridl.— Polypodium) 39:229

Aspidium maingayi (Bak.— Gymnogramine) 5: 207. ff. 1-9

Asplenium kinabaluense 7:281, t. 62 — Borneo, Clemens 29830 (SING)

A. malayo-alpinum 7:280. t. 61 — Borneo, Holttum 25485 (SING)

A. nidus var. phyllitidis (Don pro spec.) — 2:20

Athyrium amplissimum (Bak. — Asplenium) 24:569

A. angustipinna 23:8 — Malaya, Holttum SFN 39461 (SING)

. angustisquamatum 16:273.— Malaya, Holttum SFN 37359 (SING)

. curtisii 22:545. — Malaya, Molesworth-Allen 2338 (SING)

. insigne (Holtt.— Diplazium) 24:562

. kunstleri (Holtt. — Diplazium) 24:564

latisquamatum (Holtt.— Diplazium) 24:563

malaccense (Presl.— Diplazium) 24:552

montanum (v.A.v.R. — Diplazium) 24:555

. prescottianum (Wall. ex Hook. — Asplenium) 24:557

. procumbens (Holtt. — Diplazium) 24:572

. Tiparium (Holtt.— Diplazium) 24:554

. Simplicivenium (Holtt.— Diplazium) 24:573

. subintegrum (Holtt.— Diplazium) 24:557

. velutinum (Holtt. — Diplazium 24:567

Belvisia longissima 41:328.— New Guinea, Pullen 5356 (K)

Bolbitis nitens 27:453. — Malay Peninsula, Molesworth-Allen 3384 (K)

B. singaporensis 16:271.— Malaya, Hullett s.n., March 1882, 2 sheets (SING)

Cerosora sumatrana 27:450.— Sumatra, Meijer 3959 (L)

CHINGIA gen. nov. 50:31. — type species: C. ferox

. acutidens 60:17. — Celebes, Alston 15708 (BM)

. atrospinosa (C. Chr. — Dryopteris) 60:19

. Clavipilosa 60:23. N Borneo, Mt Kinabalu, Holttum 44 (SING; K)

christii (Copel. — Cyclosorus) 60:25

ferox (Bl. — Aspidium) 50:31

horridipes (v.A.v.R. — Dryopteris) 60:26

imponens (Cesati— Polypodium) 60:21

malodora (Copel. — Dryopteris) 60:19

. longissima (Brack. — Goniopteris) 60:17

. mMuricata (Brause — Dryopteris) 60:20

. paucipaleata 60:26. — Philippine Is., Luzon, Price 781 (PNH; K)

. perrigida (v.A.v.R. — Phegopteris) 60:27

. Supraspinigera (Rosenst. — Dryopteris) 60:27

. pricei 60:22. — Philippine Is., Luzon, Price 512 (PNH; K) 3

. pseudoferox 60:24. — Malay Peninsula, Perak, Matthew s.n., 6 Feb 1908 (K)

. Sambasensis 60:24. W Borneo, Brooks s.n, Sept 1908 (BM; K, L)

. urens 60:28. — Philippine Is., Luzon, Mt Makiling, Price 2367 (PNH; K)

Christella sect. PELAZONEURON 57:144 — type species: Polypodium patens

C. afzelii (C. Chr. — Dryopteris) 57:146

C. altissima 57:141.— Natal, Buchanan 103b (K)

> >>> > DP DL DD SD

anaaAANAAARAAAAAANANA

Fern names of R. E. Holttum 223

(Christella cont.)

anaaaaaaaaaaanaanaan ananaanann

. arida (D,. Don — Aspidium) 62:206

. callensii (Alston — Cyclosorus) 57:146

. assamica (Bedd.-— Nephrodium multilineatum var.) 62:210

. chaseana (Schelpe — Thelypteris) 57:148

. Clarkei (Bedd. — Pleocnemia) 62:203

crinipes (Hook. — Nephrodium) 62:208

cylindrothrix (Rosenst.— Dryopteris) 62:208

dentata (Forsk. — Polypodium) 55:306, [transfer previously made by Brownsey

& Jermy in Brit. Fern Gaz. 10:338. 1973]

distans (Hook. — Nephrodium) 57:140

. evoluta (Clarke & Baker — Nephrodium amboinense var.) 62:208

friesii (Brause — Dryopteris) 57:145

. gueinziana (Mett.— Aspidium) 57:147

. guineensis (Christ — Dryopteris) 57:145

hilsenbergii (Pres! — Nephrodium) 57:142

jerdonii (Ching — Cyclosorus) 62:206

lebeufii (“‘leboefii’?) (Baker — Polypodium) 62:206

meeboldii (Rosenst.— Dryopteris) 62:208

microbasis (Baker — Nephrodium) 57:146

modesta 57:144. — Madagascar, Decary 16,485 (P)

. molliuscula (Kuhn — Aspidium) 62:205

. multifrons (C. Chr. — Dryopteris) 57:145

namburensis (Bedd. — Polypodium) 62:206

. papilio (Hope — Nephrodium) 62:208

. papyracea (Bedd.— Nephrodium) 62:206

. prolixa (Willd. — Aspidium) 54:83

. zeylanica (Fée — Nephrodium) 62:208

Cibotium arachnoideum (C. Chr. — Cibotium cumingii var.) 32:166

Colysis acuminata (Bak. — Gymnogramme) 24:162

CORYPHOPTERIS gen. nov. 50:33. — type species C. viscosa

didymochlaenoides (Clarke — Nephrodium gracilescens var.) 62:203

hirsutipes (Clarke — Nephrodium gracilescens var.) 62:203

viscosa (Bak. — Nephrodium) 50:33

Crypsinus laciniatus (Presl — Phymatodes) 24:198

stenophyllus (Bl. — Polypodium) 24:199

subfasciatus (Holtt.— Phymatodes) 24:202

Ctenitis manilensis (Presl. — Lastrea) 47:36

setosa (Presl — Lastrea) 47:20

Ctenopteris alata (Bl. — Davallia) 24:232

ANAAANANAAN

. brevivenosa (v.A.v.R. — Polypodium) 24:228

. contigua (Forst. — Trichomanes) 24:230

khasyana (Hook. — Polypodium) 24:233

. leysii (Bak. — Polypodium) 24:232

. minuta (Bl. — Polypodium) 24:228

. moultonii (Copel. — Polypodium) 24:229

. obliquata (Bl. — Polypodium) 24:229

. Sparsipilosa 16:268.— Malaya, Holttum SFN 23522 (SING)

. Subminuta (v.A.v.R. — Polypodium) 24:228

Cyathea subgen. Cyathea sect. GYMNOSPHAERA (BI. — pro gen.) 32:115

C. subgen. SPHAEROPTERIS (Bernh. — pro gen.) 32:124

subgen. Sphaeropteris sect. SCHIZOCAENA (J. Sm. — pro gen.) 32:141

224 Gardens’ Bulletin, Singapore — XXX (1977)

(Cyathea cont.)

fepeteerne 2 MM NANNNAAANANAANAANAANAAN ANNHAAAN AANAARA AN, Gee

subgen. Sphaeropteris sect, Schizocaena subsect. SARCOPHOLIS 32: 153. fig. 30.

Type species: Cyathea rosenstockii —

subgen, Sphaeropteris sect. Sphaeropteris subsect. FOURNIERA (Bommer —

pro gen.) 32:138. fig. 27

. acanthophora 30:51.—N. Borneo, Clemens 34012 (BO; US)

aeneifolia var. macrophylla 32:134.—-NE New Guinea, Hoogland & Schodde

7209 (L)

. aeneifolia var, melanacantha (Copel. — Cyathea) 32:134

. agathetii 30:51. — Indonesian Borneo, Kostermans 12870 (BO; A, K, L)

. alleniae 30:52. — Pahang, Molesworth Allen 4127 (K; US)

angustipinna 30:52. — Sarawak, Richards 1675 (K; SING, US)

archboldii var, horrida 32:99. NE New Guinea, Hoogland & Pullen 5506 (K)

. atrospinosa 30:53. —- NE New Guinea, Hoogland & Pullen 6090 (K)

atrox var. inermis 32: 130. figs. 24, 25. — NE New Guinea, Hoogland & Schodde

7457 (L)

. australis subsp. norfolkensis 33:244. — Norfolk Id., Metcalf s.n. (NSW, P5817)

. carrii 30:53. — Papua, Carr 13526 (K; BM, L)

. catillifera 30:53. Papua, Brass 4549 (BRI)

. cicatricosa 33:274. — New Caledonia, Green 1778 (K)

. coactilis 31:533.— Papua, Schodde 1887 (CANB)

cucullifera 30:54. —- NE New Guinea, Hoogland & Pullen 5497 (K; BO, BRI,

L, SY, Us)

decurrens subsp. epaleata 33:249. — Tahiti, Grant 4233 (K)

deminuens nom, nov. 32:145.— Alsophila parvifolia

dicksonioides 31:529.—- New Guinea, Hoogland & Schodde 7171 (L)

discophora 30:54. N. Borneo, Clemens 31698 (B; BO, US)

eriophora 30:55. — Papua, Carr 14439 (K; BM, L)

excavata 8:306, t.31. — Malaya, Holttum SFN 23538 (SING)

gigantea (Wall. — Alsophila) 8:318

glaberrima 30:55. — Papua, Brass 27092 (L; US)

hooglandii 30:56. -NE New Guinea, Hoogland & Schodde 7203 (L)

inaequalis 30:56. — Papua, Brass 23547 (L; A)

insulana 30:56. — Papua, Brass 24725 (A; L)

insularum 30:57. — Papua, Brass 27419 (L; K, US)

jacobsii 57:499.— S. Sumatra, Jacobs 8113 (L; BO, K)

kanehirae nom. nov. 32:113.— Alsophila arfakensis

latebrosa var, indusiata 8:305.— Malaya, Holttum SFN 23539 (SING)

loerzingii 30:58. — Sumatra, Lérzing 14904 (L; BO)

lunulata subsp. vitiensis (Carr. — Alsophila vitiensis) 33:264

. macrophylla var. quadripinnata 32:141.— NE New Guinea, Womersley NGF

13959 (K)

. mesosora 30:57. —-_NE New Guinea, Robbins 1032 (L)

. microchlamys 30:58. — Philippine Is., Ramos 30475 (US; P)

. nigrolineata 30:58. NE New Guinea, Hoogland & Pullen 5495 (K; BO, BRI,

L, SYD, US)

. higropaleata 30:59. NE New Guinea, Pullen 666 (L)

. nilgirensis 35:468. — India, Gamble 17799 (K)

nothofagorum 41:327.— NE New Guinea, Pullen 5358 (K)

. Odsora 30:59.—N. Borneo, Clemens 51188 (K; A, L, MICH, UC) >

. pallidipaleata 30:60.— Celebes, Eyma 776 (BO)

. parvipinna 30:60.— Papua, Brass 25837 (L; K, US)

. pseudomuelleri 30:61. — W. New Guinea, Brass 9430 (A; BO, L, MICH, UC)

- pycnoneura 31:533.— New Guinea, Pullen 562 (CANB: BM, L)

. robusta (C. Moore — Alsophila) 33:265

Fern names of R. E. Holttum 225

(Cyathea cont.)

C. rubella 30:61. — Papua, Hoogland 4487 (K; A, BM, BRI, L)

. Sarasinorum nom, nov. 30:61.— Alsophila contaminans var. longepaleata

. semiamplectens 30:62.—- NE New Guinea, Womersley 11500 (BRI; L)

. setifera 30:62. N. Moluccas, Main & Aden 1306 (BO; K, L)

solomonensis 33:252.— Solomon Is., Whitmore & Womersley BSIP 1035

(LAE)

subtripinnata 31:534. — Papua, Schodde 1763 (CANB)

. stelligera 33:250.— New Caledonia, McKee 9863 (NSW)

. stipitipinnula 30:62.— N. Borneo, Clemens 33156 (K; A, BO, L, UC)

. womersleyi 30:63. -NE New Guinea, Womersley & Millar 8470 (K; A, BO.

L, SYD)

vandeusenii 31:529.—- New Guinea, Brass 29989 (L; K)

. verrucosa 30:63. — Sumatra, Matthew s.n. (K)

AA AA|ANM AAAN

yclosorus deminuens 34:135.— New Guinea, Brass 32106 (K)

dicranogramma (v.A.v.R.— Dryopteris) 24:267

ecallosus 16:269.— Malaya, Holttum SFN 31294 (SING)

heterocarpus var, glaucostipes (Bedd.— Nephrodium glaucostipes 24:27]

. jacobsii 31:530. — Sarawak, Jacobs 5086 (L: K, SAR, G, US, B, S-PA)

. Menisciicarpus (Blume — Aspidium) 44:39

. oligodictyus (Bak. — Acrostichum) 31:530

. polycarpus (Bl.— Aspidium) 24:283

Davallia dimorpha 9:122.— Malaya, Holttum SFN 31298 (SING)

D. trichomanoides var. lorrainii (Hance — Davallia lorrainii) 24:361

Dicksonia lanigera 30:64. W. New Guinea, Pulle 1010 (L; BM)

D. mollis 30:64. — Philippine Is., Elmer 9874 (K; L, MICH, US)

Dicranopteris subgen. ACROPTERYGIUM (Diels, pro sect. sub Gleichenia

subgen. Mertensia 26:261

. Clemensiae 26:275 —N. Borneo, Clemens 28745 (BM; K, L, US)

. linearis var. alternans (Mett. — Gleichenia dichotoma var, 26:278

. linearis var, altissima 26:276.— Malay Peninsula, Corner SF 31447 (SING:

KL)

linearis var. demota 26:275 — N. Borneo, Clemens 29535 (BM; K, L, US)

linearis var. dichotoma (Thunb. — Polypodium dichotomum) 26:277

linearis var. ferruginea (Blume — Gleichenia ferruginea) 26:278

linearis var. glabra 44:40— Bismarck Archipelago, Noona Dan Exped.

2608 (C)

linearis var. inaequalis (Rosenst. — Gleichenia) 26:278

linearis var. latiloba 26:277. — Philippine Is., Merril] 975 (US)

linearis var. montana 26:276.— Malay Peninsula, Molesworth Allen 2720

(SING)

linearis var. rigida (Blume — Gleichenia hermannii var.) 26:277

linearis var. subpectinata (Christ — Gleichenia subpectinata 26:277

linearis var. subspeciosa 26:278.— N. Borneo, Topping 1516 (SING; US)

speciosa (Pres]l — Hicriopteris) 26:273

aAAAANNA

O 9959 9S9 9995 995

iplazium angustipinna (Holtt.— Athyrium) 45:637

armatum (Copel. — Lastrea) 34:135

curtisii (Holtt.— Athyrium) 45:637

dielsii (Brause — Alsophila) 34: 136

. insigne 9:123.— Malaya, Holttum SFN 21635 (SING)

. kunstleri 15:88.f.2.— Malaya; Holttum SFN 31194 (SING)

. latilobum 43:308. — Thailand, Hansen, Seidenfaden & Smitinand 11155 (C; K)

. latisquamatum 9:124.— Malaya, Holttum SFN 31311 (SING)

whelvieivleic)

226 Gardens’ Bulletin, Singapore — XXX (1977)

(Diplazium cont.)

. procumbens 15:95.f.4. — Malaya, Holttum SFN 36503 (SING)

. Tiparium 15:97.f.5.— Malaya, Hume 8186

. simplicivenium 15:100.f.6.— Malaya, Holttum SFN 36507 (SING)

. Speciosum var. major (Bedd. — D, sorzogonense var.) 15:103

. stellatopilosum (Brause — Dryopteris) 34: 136

. Stipitipinnula 41:328 New Guinea, Hoogland 9090 (K)

subintegrum 9:125.— Malaya, Holttum 31350 (SING)

. Velutinum 9:126.— Malaya, Holttum SFN 31221 (SING)

. woodwardioides (Presl — Brachysorus) 47:30

wluluieieieioioie)

Diplora translucens 44:41. — Bismarck Archipelago, Noona Dan Exped. 1824 (C)

Dryopteris brunnea var. glabrata (Clarke — Polypodium distans var.) 9:127

D. haniffii 16: 272. f. 1.— Malaya, Holttum SFN 31286 (SING)

D. urophylla var. nitida 7:249, 251 (text-f.) — Borneo, Holttum 25592 (SING)

Elaphoglossum angustifrons 40:319. NE New Guinea, Womersley 11272 (LAE;

. apiculatum 40:320.— W. New Guinea, Brass 13624 (G)

. apoense 40:320. — Philippine Is., Edafio PNH 710 (MICH)

. arachnoideum 40:320.— Papua, Brass 24541 (A)

. brevifolium 16:270.— Malaya, Holttum SFN 20750 (SING)

calanasanicum 64:196. — Philippine Is., Luzon, Price 2933 (K; PNH)

favigerum 40:321.— W. New Guinea, Brass 13440 (G; MICH)

heterostipes 40:321.—C. Indonesian Borneo, Endert 4424 (BO)

idenburgensis 40:321. —- W. New Guinea, Brass 12283 (G; MICH)

indrapurae 40:321.— Sumatra, Alston 14275 (BM)

latemarginatum 40:322.— Papua, Brass 24896 (A)

malayense 40:322.— Malay Peninsula, Henderson 17765 (K; SING)

melanochlamys 40:322.—-W. New Guinea, Eyma 5408 (BO)

negrosensis 40:323.— Philippine Is., Elmer 9885 (MICH; BO)

nesioticum 40:323.— Papua, Brass 24882 (A)

. nigripes 40:323. — W. New Guinea, Pulle 493 (BM; BO, L, U)

ophioglossoides (Goldm. — Acrostichum) 64:197

. pallescens 40:323.— Papua, Brass 22879 (A)

peninsulare 16:270. — Siam, Eryl Smith 2021 (SING)

planicosta 40:324.—- NE New Guinea, Jermy 4216 (BM)

. pullenii 40:324.-_NE New Guinea, Pullen 5034 partim (L)

. resiniferum 40:324.— Papua, Wakefield 1466 (BM)

. spongophyllum Bell ex. 40:325.— N. Borneo, Clemens 31869 (BO)

. Stenolepis Bell ex. 40:325.— N. Borneo, Clemens 28019 (US)

. Ssumatranum 40:325.— Sumatra, Matthew s.n. (K)

. thamnopteris 40:326.— Papua, Brass 22990 (A)

. vepriferum 40:326.—-NE New Guinea, Clemens 7417 (B; G)

ot oe subgen, DIPLOPTERYGIUM — (Diels, pro sect. sub subgen. Merten-

sia :261

. alstonii 26:267.— Sumatra, Alston 14981 (BM)

. angustiloba 26:263.—N. Guinea, Brass 4960 (BM; K)

. brevipinnula 26:264.— Sarawak, Bell 2042 (BM)

clemensiae (Copel. — Hicriopteris) 28:14

clemensiae var, membranacea (Holtt.— Gleichenia papuana var.) 28:14

. deflexa 26:264.— Addendum, New Guinea, Papua, Brass 27171 (L)

hirta var. amboinensis (v.A.v.R.— Gleichenia amboinensis) 26:269

hirta var. amoena (v.A.v.R.— Gleichenia amoena) 26:269

. hirta var. candida (Rosénst.—Gleichenia candida) 26:269

cleFehefelutestutcielctencleltlolutonstolesol relate:

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Fern names of R. E. Holttum 227

(Gleichenia cont.)

. hirta var. lanuginosa (v.A.v.R.—G. ornamentalis var.) 28:27

. hirta var. ornamentalis (Rosenst. — Gleichenia ornamentalis) 26:269

. hirta var, virescens (Hieron. — Gleichenia dolosa var.) 26:270

. hooglandii 41:327. - NE New Guinea, Hoogland & Schodde 7692 (CANB)

loheri var. major 26:272.— Celebes, Bunnemeijer 11965 (BO; K, L)

matthewii 26:265.— Sumatra, Matthew s.n., 31 Jan. 1912 (K)

microphylla var. vulcanica (Bl. — Gleichenia vulcanica) 24:66

paleacea (Copel. — Hicriopteris) 26:265

papuana 26:266.——N. Guinea, Brass 22597 (BM)

papuana var. membranacea 26:266.— N. Guinea, Brass 24763 (BM)

peltophora var. schizolepis C. Chr. ex. 26:262.— Sarawak, Mjoberg 9 (P)

pulchra (Copel. — Sticherus) 26:271

. Sumatrana 26:264.— Sumatra, Alston [‘1484” sphalm.] 14841 (BM)

truncata var. bracteata (v.A.v.R.—G. laevigata var.) 26:271

truncata var. celebica 28:22. — (Central) Celebes, 2600-3000 m. alt. [nothing

else quoted]

truncata var. involuta 26:272.— Sumatra, Winkler (Rosenst., Fil. Sumatr.

exsicc. 182) (L; BM, P)

truncata var. plumiformis (Presl — Mertensia) 26:272

. venosa (Copel. — Sticherus) 29:20

. volubilis var. peninsularis 26:265.— Malay Peninsula, Bintang FMS. Mus. s.n.,

June 1917 (K)

Grammitis crispatula 16:268.— Malaya, Burkill SFN 12705 (SING)

Haplodictyum bulusanicum 56:61.— Philippine Is., Luzon, Elmer 16585 (K; BO,

FL-G, L)

Heterogonium alderwereltii nom. nov. 18:163.f.1.— Pleocnemia stenosemioides

. giganteum (Blume — Aspidium) 18:166

. gurupahense (C. Chr. — Dryopteris sagenoides var.) 25:272

. pinnatum (Copel. — Stenosemia) 18:163

sagenoides (Mett. — Aspidium) 18:161.f.2 A&B.

. Saxicolum (Blume — Aspidium) 18:165. f.2 D.

. subaequale (Rosenst.— Aspidium) 28:274

subglabrum 25:273. — Sumatra, Lérzing 5520 (BO)

. subsageniaceum (Christ — Aspidium) 25:273

. teysmannianum (Bak. — Acrostichum) Posthumus ex. 19:29

. wigmanil (Racib. — Aspidium) 25:27]

Histiopteris caudata (Copel. — Lepidocaulon) 42:459

H. pilosa 42:460.— Sumatra, Matthew s.n. (K)

H. squamulata 42:459.— New Guinea, Millar & Holttum 18640 (K)

Hymenophyllum johorense 3:408 cum fig. — Malaya, Holttum 10755 (SING)

Lastreopsis parishii var. chupengensis (Ridl. — Lastrea chupengensis) 24:498

Lepisorus longifolius (Bl.— Grammitis) 24:151

Leptochilus simplicifolius 3:409.— Malaya, Henderson 17977 (SING)

Lindsaya cultrata var. parvula 4:61 cum fig. — Malaya, Nur 7297 (SING)

L. cultrata var. plumula (Ridley. — Lindsaya) 24:329

L. integra 4:67 cum fig. — Malaya, Holttum 20934 (SING)

L. kinabaluensis 7:237.— Borneo, Clemens 27744 (simply pinnate), Holttum

25433 (bipinnate) (SING)

L. malayensis 4:69 cum fig. — Malaya, Haniff 4032 (SING)

AAA A AgAgAAAAANDDADAD

sofaofesfeofasfastesf=sfecfe=

228 Gardens’ Bulletin, Singapore — XXX (1977)

(Lindsaya cont.)

L. scandens var. terrestris 24:327. — (sine typus)

L. tenera var. gigantea (Hook. — L. flabellata var.) 4:65.

Lomagramma brassii 38:224.— Papua, Brass 24947 (A)

L. copelandii 11:201. tt. 9 & 10.— Philippine Is., Ramos & Edafio PBS 28826

(PNH; BO)

. cordipinna 11:202. tt. — Samoa, Safford s.n., March 1888 (US)

cultrata (Baker — Acrostichum) 11:203

grosseserrata 11:203. tt. 10 & 11.— Siam, Winit 1019 (SING)

leucolepis 38:224.— W. New Guinea, Brass 12950 (L; BM, BO, MICH, GH)

. matthewii (Ching — Campium) 11:206

. merrillii 11:208.t. 14. — Philippine Is., Merrill 8282 (PNH)

. tahitensis 37:223. — Tahiti, Vesco s.n. (P; B, K)

omariopsis amydrophlebia (Slosson— Stenochlaena) 14:617

boivinii 13:53. — Madagascar, Boivin 1947/2 (P)

congoénsis 14:622. ff. 8 & 9.— Portuguese Congo, Gossweiler 8219 (BM)

. crassifolia 13:57. Madagascar, Humblot 529 (P; K)

intermedia (Copeland — Stenochlaena) 6:270

jamaicensis (Underw. — Stenochlaena) 14:614

kingii (Copeland — Stenochlaena) 6:273

kingii var. petiolulata Ros. ex. 6:273.—- New Guinea, Bamler 50 (UC)

. kunzeana (Underw. — Stenochlaena) 14:617

lanceolata 14:619. — Brazil, Sellow s.n. (B)

. latiuscula (Maxon — Stenochlaena) 14:617

lineata (Presl — Olfersia) 47:9

longicaudata (Bonap. — Stenochlaena) 13:58

maxonii (Underw. — Stenochlaena) 14:617

mexicana 14:617.— Mexico, Hahn 56 (P)

muriculata 14:624. f.10.— Belgian Congo, Jespersen 26 (BR)

nigrescens 14:627. ff. 13 & 14.— Fernando Po, Mann s.n. (K)

nigro-paleata 14:618. ff.4&5.— Peru, Klug 2082 (US; BM, K, NY)

. raciborskii (C. Chr.) 6:272.— Stenochlaena Raciborskii

. rossii 14:625. ff. 11 & 12. — Nigeria, Ross 22 (BM)

. setchellii (Maxon) 6:276. — Stenochlaena Setchellii

. speciosa 14:620. — Brazil, Sellow 370 (B)

. Subtrifoliata (Copeland — Stenochlaena) 6:274

. underwoodii 14:614. ff, 1 & 2.— Jamaica, Jenman s.n. (NY)

Macrothelypteris ogasawarensis (Nakai — Dryopteris) 48:27

M. polypodioides (Hook. — Alsophila) 48:29

M. torresiana var. calvata (Bak. — Nephrodium setigerum var.) 48:28

Se al al we we ae

Seepage wieilelgic laluigl ls) |<

Mesoneuron brackenridgei (Mett.— Aspidium) 44:44

M. wantotensis (Copel. — Dryopteris) 34:134

MESOPHLEBION nom. gen. nov. 50:29. — Mesoneuron

. subgen, PLESIONEURON 50:30. — type species: M. tuberculatum

. arenicola 63:228.— Sumatra, Meijer 4478 (L)

. auriculiferum (v.A.v.R.— Dryopteris) 63:231

. beccarianum (Cesati— Nephrodium) 63:230

. Carolii 63:227. — Sarawak, Hose 712 (K; BM)

. chlamydophorum (Rosenst, ex. C. Chr.— Dryopteris) 63:231

. crassifolium (Bl.) 50:30.— Aspidium crassifolium

. dulitense 63:229.— Sarawak, Hose s.n., 1894 (K)

. echinatum (Mett.— Aspidium) 63:226

—<SS<SS52588

Fern names of R. E. Holttum 229

(Mesophlebion cont.)

. endertii (C. Chr. — Dryopteris) 63:228

. hallieri (Christ — Aspidium) 63:227

. motleyanum (Hook. ex. Hook. & Bak.— Nephrodium) 62:209

. Oligodictyon (Baker — Acrostichum) 63:227

pallescens (Brause — Dryopteris) 63:230

. persquamiferum (v.A.v.R.— Dryopteris) 63:226

. rufescens 63:231.— Papua, Brass 27957 (K; BO, L, LAE)

teuscheri (v.A.v.R. — Dryopteris) 63:228

. trichopodum (C. Chr. — Dryopteris) 63:226

. tuberculatum (Cesati— Nephrodium) 50:30

. vinosicarpum (v.A.v.R.— Dryopteris) 63:229

Metathelypteris dayii (Bedd. — Nephrodium) 62:205

M. fragilis (Baker — Polypodium) 57:127

Microsorium congregatifolium (v.A.v.R.— Pleopeltis) 24:178

M. sarawakense (Bak.— Polypodium) 24:175

NANNOTHELYPTERIS gen. nov. 50:38.— type species: N. aoristisora

N. aoristisora (Harr. — Polypodium) 50:38

N. inaequilobata 56:67. — Philippine Is., Luzon, Cuming 272 (K; B, E, G, L, US)

N. nervosa (Fée — Phegopteris) 56:66

N. philippina (Presl — Physematium) 56:66

Oreopteris elwesii (Bak. ex Hook. & Bak. — Nephrodium) 62:204

Orthiopteris kingii (Bedd. — Dicksonia) 24:306

Parathelypteris salazica 57:136.— Réunion, Cadet 1981-bis (P)

PARATHYRIUM gen. nov. 27:449. — species typica: P. boryanum [= Dryoathy-

rium Ching]

P. boryanum (Willd. — Aspidium) 27:449

. forsythii-majoris (C. Chr. — Dryopteris) 27:449

. macdonellii (Bedd. — Asplenium) 27:449

. parvisorum (C. Chr.— Dryopteris) 27:449

. pterorachis (Christ — Athyrium) 27:449

. unifurcatum (Bak. — Nephrodium) 27:449

. viridifrons (Makino — Athyrium) 27:449

Phymatodes crustacea (Copel. —- Lecanopteris) 24:190

P. subfasciata 16:267.— Malaya, Holttum SFN 21559 (SING)

Pleocnemia andaiensis (Baker — Polypodium) 61:350

. brongniartii (Bory — Polypodium) 61:350

. chrysotricha (Baker — Nephrodium) 20: 187

. conjugata var. elatior 20:179. — Sumatra, Matthew 512 (BO)

. dimidiolobata 20: 184. ff. 16, 17. — Solomon Is., Brass 2648 (BO)

. elegans (Copel. — Tectaria) 61:348

. hemiteliiformis (Racib. — Pleocnemia leuzeana var.) 20:179. f. 11.

intermedia 61:349. — Sarawak, Anderson S 32555 (K)

. irregularis (Presl— Polypodium) 61:347

macrodonta (Fée — Dictyopteris) 61:348

. megaphylla 61:357. - New Guinea, Jermy 8149 B (cult. K)

olivacea (Copel. — Tectaria) 20:181. ff. 8, 10

. pleiotricha 20: 182. ff. 12, 13. North Borneo, Elmer 21421 (SING; BO)

. presliana 20: 183. ff. 14, 15. — Philippine Is., Edafio BS 78717 (SING; BO)

. seranensis 20:187.— Moluccas, Rutten 1850 (BO; SING)

. submembranacea (Hayata — Aspidium) 61:352

SSSSS555888

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230 Gardens’ Bulletin, Singapore — XXX (1977)

(Pleocnemia cont.)

tripinnata 20: 185, ff. 18, 19. — New Guinea, Lauterbach 560 (SING; BO)

winitii 20: 181. — Thailand, Winit 36 (SING)

PLESIONEURON (Holtt.— Mesophlebion subgen. Plesioneuron) 63:232

P. altum (Brause — Dryopteris) 63:241

ys yo oO

POI UVUIUUUUUUUU UU UU UU UU UU VUDUUUVUUUUUUUUUUT

. archboldiae (Copel. — Lastrea) 63:242

. attenuatum (Brack. — Lastrea) 63:245

. belense (Copel. — Dryopteris) 63:239

. bipinnatum (Copel. — Dryopteris) 63:242

. costulisorum (Copel. — Lastrea) 63:246

crassum (Copel. — Dryopteris) 63:243

ctenolobum 63:247.— NE New Guinea, Jermy 3604 (BM)

. doctersii 63:246.— W. New Guinea, Docters van Leeuwen 10357 (BO; L)

dryas 63:247.— Papua, Brass 23447 (BM; L, LAE)

. dryopteroideum (Brause — Alsophila) 63:237

falcatipinnulum (Copel. — Dryopteris) 63:244

. fuchsii 63:236.— N. Borneo, Fuchs 21477 (L; K)

. fulgens (Brause — Dryopteris) 63:238

. hopeanum (Bak. — Nephrodium) 63:244

hopeanum var. acutilobum 63:245.— Fiji, Milne 190 (K)

. Irayense (Copel. — Cyclosorus) 63:239

kundipense 63:247. — NE New Guinea, Millar & Holttum NGF 18593 (LAE;

BRI, K)

. marattioides (Alston — Dryopteris) 63:238

. myriosorum (Copel. — Dryopteris) 63:244

notabile (Brause — Dryopteris) 63:237

. ophiura (Copel. — Dryopteris) 63:248

. phanerophlebium (Bak. — Nephrodium) 63:243

. platylobum 63:246.— NE New Guinea, Jermy 3610 (BM)

ponapeanum (Hosokawa — Phegopteris) 63:241

prenticei (Carr. — Lastrea) 63:244

. pullei 63:236.— W. New Guinea, Pulle 905 (L; BM)

quadriquetrum (v.A.v.R. — Dryopteris) 63:242

. rigidilobum 63:248.— NE New Guinea, Walker 774i (BM)

. royenii 63:240 — W. New Guinea, van Royen 5469 (L)

. savaiense (Bak. — Nephrodium) 63:240

septempedale (Alston — Dryopteris) 63:238

subglabrum 63:243.— Solomon Is., Braithwaite RSS 4512 (K)

subterminale 63:241. -NE New Guinea, Brass 32049 (K; L, LAE)

. tahitense 63:241. — Tahiti, Savatier 752 (K)

. translucens 63:240.— Amboina, Brooks 17760 (BO; BM)

. tuberculatum (Cesati— Nephrodium) 63:246

wantotense (Copel. — Dryopteris) 63:245

. wariense (Copel. — Dryopteris) 63:239

. woodlarkense (Copel. — Cyathea) 63:236

. angusticaudata 55:308.— Papua, Brass 24984 (BM; LAE)

. auctipinna 55:310.—N. Celebes, P. & F. Sarasin 119 (BAS)

. basicurtata 55:309. — Sumatra, Robinson & Kloss 148 (BM)

. blastophora (Alston — Cyclosorus) 57:156

. brooksii (Copel. — Dryopteris brooksii) 55:308

. bryanii (C. Chr. — Dryopteris) 55:317

. caudata (Holtt.— Pseudocyclosorus) 55:321

eumatopteris afra (Chr. — Dryopteris) 55:306

Fern names of R. E. Holttum 231

(Pneumatopteris cont.)

. cheesmaniae 55:307.—-W. New Guinea, Cheesman 1269 (BM)

. christelloides 55:311.— N. Borneo, Mt Kinabalu, Clemens 27541 (K)

. comorensis 55:318.— Comoro ls., Hildebrandt 1782 (K; B, BM, FI, L)

. costata (Brack. — Goniopteris) 55:305

. deficiens 55:321. — Flores, Posthumus 3183 (BO; K)

dicranogramma (v.A.v.R.— Dryopteris) 55:30]

ecallosa (Holtt. —- Cyclosorus) 55:310

egenolfioides 55:306.— W. New Guinea, Brass 11497 (L; BM, BO)

. excisa (Holtt.— Pseudocyclosorus) 55:32]

. glaberrima (Richard — Aspidium) 55:318

. glabra (Copel. — Cyclosorus) 55:301

glandulifera (Brack. — Goniopteris) 55:302

hudsoniana (Brack. — Nephrodium) 55:317

. humbertii 55:313. — Madagascar, Humbert 31679 (P)

imbricata 55:322.— Amboina, Brooks 17775 (BM; BO, L)

. incisa 55:317. — Ternate, Alston 16623 (BM)

. inclusa (Copel. — Dryopteris) 55:306

. jermyi 55:310.— NE New Guinea, Jermy 3739 (BM)

kerintjiensis 55:312.— Sumatra, Alston 14149 (BM)

keysseriana (Rosenst.— Dryopteris) 55:320

laevis (Mett. — Aspidium) 55:308

. laticuneata 55:312. - NE New Guinea, Jermy 8044 (BM)

. latisquamata 55:305.— NE New Guinea, Millar & Holttum NGF 15777 (K:

LAE)

. ligulata (Presl — Lastrea) 55:320

. longipes (Bl. — Aspidium) 55:306

. lucida (Bak. — Nephrodium) 55:307

. Macroptera (Copel. — Dryopteris) 55:316

. magnifica (Copel. — Dryopteris) 55:319

. mesocarpa (Copel. — Dryopteris) 55:303

. michaelis 55:313.— Philippine Is., Price 317 (K)

. Microauriculata 55:311.— N. Borneo, Mt Kinabalu, Clemens 27137 (BM)

. microloncha (Chr. — Dryopteris) 55:308

micropaleata 55:319.— N. Borneo, Mt Kinabalu, Holttum 58 (K; SING)

. nephrolepioides (C. Chr. — Dryopteris) 55:320

nitidula (Pres] — Nephrodium) 55:318

novae-caledoniae 55:313. — New Caledonia, Lécard s.n., Oct. 1876 (P)

obliqua 55:309. — Celebes, P. & F. Sarasin 127 (BAS)

obstructa (Copel. — Dryopteris) 55:302

. oppositifolia (Hook. — Polypodium) 55:304

. oxyura (Copel. — Dryopteris) 55:315

petrophila (Copel. — Dryopteris) 55:322

. regis (Copel. — Dryopteris) 55:322

. remotipinna (Bonap. — Dryopteris) 55:311

. rodigasiana (T. Moore — Nephrodium) 55:319

rotumaensis (St. John — Cyclosorus) 55:318

papuana 55:311.— Papua, Brass 22778 (LAE; BM, L)

. parksii (Ballard — Dryopteris) 55:302

. patentipinna 55:307.— NE New Guinea, Walker 8477 (BM)

. pennigera (Forst.— Polypodium) 55:305

. pergamacea 55:315.— New Ireland, Jermy 7877 (cult. K, 123/70, no. 1035)

. prismatica (Desv.— Nephrodium) 55:316

OOOO UOMO UU UU UU UUM VUVUUVUUUUMUUUUUUUUU UU UY

232 Gardens’ Bulletin, Singapore — XXX (1977)

(Pneumatopteris cont.)

a-Ma-Ba-Ma-Na-Ba-Be-Ba- Bene Be- eee

a-Na-Ra-Me-Me- ha"

. sandwicensis (Brack. — Stegnogramma) 55:303

. sibelana 55:301.— Moluccas, Batjan, Alston 16942 (BM)

. sogerensis (Gepp. — Dryopteris) 55:315

. stokesii (E. Brown — Dryopteris) 55:302

. subappendiculata (Copel. — Dryopteris) 55:303

. subpennigera (C. Chr. — Dryopteris) 55:304

. sumbawensis (C. Chr. — Dryopteris) 55:323

. Superba (Brause — Dryopteris) 55:303

. remotipinna (Bonap. — Dryopteris) 57: 159

. tobaica 55:316. — NE Sumatra, Surbeck 14 (L; BO)

. transversaria (Brack. — Nephrodium) 55:319

. truncata (Poir. — Polypodium) 55:314

truncata var, loyalii 55:314. — Darjeeling, Loyal sn. Aug. 1956 (Herb. Punjab

Univ. no. 3172)

unita (Kunze — Gymnogramma) 55:304

. usambarensis 55:312. — Tanzania, Faden et al. 70/293 (EA)

. vaupelii (C. Chr. — Dryopteris) 55:317

. venulosa (O, Kuntze — Dryopteris) 55:315

. versteeghii 55:321. — W. New Guinea, Versteegh BW 10259 (L)

. walkeri 55:323. — NE New Guinea. Walker 9980 (BM)

PODOSORUS gen. nov. 42:455.— Type-species: P. angustatus Holtt.

angustatus 42:455. — Philippine Is., Gutierrez PNH 78332 (K; L)

Polypodium papilliferum 2:22.— Mentawi Is., Kloss 14800 (SING)

POLYSTICHOPSIS (J. Sm. pro subsect. sub Lastrea) — 17:149, 152 [= Arachnio-

SO oO *O Fo

des BI.]

. aristata (Forst. — Polypodium) 24:468

. chinensis (Rosenst. — Polystichum amabile var.) 24:486

. hanifii (Holtt.— Dryopteris) 24:485

. hasseltii (Bl. — Polypodium) 24:487

. puncticulata (v.A.v.R. — Polystichum) 24:487

Pronephrium sect. DIMORPHOPTERIS (Tagawa & K. Iwats. — Dimorphopteris

ee eS OO eo OU

pro gen.) 50:36. ff. 10, 10a

. sect, GRYPOTHRIX 50:36. fig. 8. — type species: Meniscium cuspidatum BI.

. acanthocarpum (Copel. — Dryopteris) 53: 107

. amboinense (Willd. — Aspidium) 53:120

. aquatiloides (Copel. — Dryopteris) 53:108

. articulatum (Houlst. & Moore — Nephrodium) 53:116

. asperum (Presl — Polypodium) 53:112

. brauseanum nom, nov. 53:107.— Dryopteris canescens var. novoguineensis

. beccarianum (Cesati— Meniscium) 53:107

. buwaldae 53:115.— Aru Is., Buwalda 4979 (L)

. celebicum (Bak. — Acrostichum) 53:119

. clemensiae (Copel. — Dryopteris 53:118

. crenulatum 53:123.— Vietnam, Eberhardt s.n. 1896 (P)

. cuspidatum (Bl. — Meniscium) 53:123

. debile (Bak. — Nephrodium) 53:118

. diminutum (Copel. — Dryopteris) 53:115

. elmerorum (Copel. — Dryopteris) 53:118

Fern names of R. E. Holttum 233

(Pronephrium cont.)

DS WUD UUD UD UND UN DUD DU UU UU UD

. euryphyllum (Rosenst. — Dryopteris) 53:112

. exsculptum (Bak. — Acrostichum) 53:117

. firmulum (Bak. — Polypodium) 53:116

. gardneri 49:81.— Ceylon, Gardner 1135 (K), s.n. (BM)

glandulosum (Bl. — Aspidium) 53:118

granulosum (Presl — Polypodium 53:119

gymnopteridifrons (Hayata — Dryopteris) 53:112

hosei (Bak. — Meniscium) 53:120

. insulare (K. Iwats. — Abacopteris) 53:123

kjellbergii 53:118.— Celebes. Kjellberg 2638 (BO)

lakhimpurense (Rosenst. — Dryopteris) 53:110

longipetiolatum (K. Iwats.— Abacopteris) 53:123

megacuspe (Bak. — Polypodium) 53:122

melanophlebium (Copel. — Dryopteris) 53:108

menisciicarpon (Bl. — Aspidium) 53:111

. merrillii (Chr. — Dryopteris) 53:117

micropinnatum 53:108.— NE New Guinea, Darbyshire & Hoogland 8014

(CANB; BM, L, LAE)

millarae 53:115.— NE New Guinea, Womersley & Millar NGF 8500 (LAE)

minahassae 53:119.—N. Celebes, Alston 16570 (BM)

moniliforme (Tagawa & K. [wats.— Dimorphopteris) 50:36

. nitidum (Holtt. — Dryopteris urophylla var.) 53:109

nothofageti 53:118.— NE New Guinea, Walker 8788 (BM)

nudatum (Roxb. — Polypodium 53:111

palauense (Hosok. — Meniscium) 53:121

palopense 53:116.— Celebes, Kjellberg 1999 (BO)

parishii (Bedd. — Meniscium) 53:12

penangianum (Hook. — Polypodium) 53:110

pentaphyllum (Rosenst. — Dryopteris) 53:108

peramalense 53:115.— W. New Guinea, Pulle 415 (BM: L)

repandum (Fée — Goniopteris) 53:109

rhombeum (Chr. — Dryopteris diversifolia var. acrostichoides subvar. 53:120

rubicundum (v.A.v.R. — Phegopteris) 53:123

rubidum (J. Sm. ex Hook. — Polypodium) 53:111

rubrinerve (Mett. — Phegopteris) 53:110

. Salicifolium (Wall. ex Hook. — Meniscium) 53:123

samarense (Copel. — Cyclosorus) 53:119

simillimum (C. Chr. — Dryopteris) 53:116

. simplex (Hook. — Meniscium) 53:122

. solsonicum 64:197.— Philippine Is., Luzon, Price 2903 (K; PNH)

. tibangense (C. Chr. — Dryopteris) 53:118

. thwaitesii (Hook. — Meniscium) 53:122

. triphyllum (Sw. — Meniscium) 53:122

. womersleyi 53: 108. NE New Guinea, Floyd & Womersley NGF 6308 (LAE:

BM)

. xiphioides (Chr. — Dryopteris) 53:119

Pseudocyclosorus camerounensis 57:137.— Cameroun, Meurillon CNAC 726

vUuWUUU

(P; K

. caudatus 34: 133. fig. 7.— New Guinea, Holttum, cult. Kew 545/63, no. 12 (K)

. excisus 34: 133. — New Guinea, Millar & Holttum 18623 (K)

. johannae 57:138.— Comoro Is., Johanna Id., Bewsher 24 (K)

. ochthodes (Kunze — Aspidium) 62:204

. petrophila (Copel. — Dryopteris) 34:133

. pulcher (Bory ex Willd. — Aspidium) 57: 138

234 Gardens’ Bulletin, Singapore — XXX (1977)

Pseudophegopteris aubertii (Desv.— Polypodium) 48:18

P. cruciata (Willd. — Aspidium) 48:21

. cyclocarpa 34:131. fig. 10.— New Guinea, Holttum, cult. Kew 578/63, no. 57

(K)

P. dianae (Hook. — Polypodium) 48:21]

P. henriquesii (Bak. — Polypodium) 48:15

P. hirtirachis (C, Chr. — Dryopteris) 48:22

P. keraudreniana (Gaud.— Polypodium) 48:19

P. kinabaluensis 48:16.— Borneo, Fuchs 21475 (KSEPL; K, L)

P. persimilis (Bak. — Polypodium) 48:20

P. procera (Mann — Phegopteris) 48:18

P. pyrrhorhachis var. glabrata (Clarke — Polypodium distans var.) 48:24

P. rectangularis (Zoll. — Polypodium) 48:19

P. sumatrana 48:22. — Sumatra, Holttum 26211 (K; SING, BO)

P. tenggerensis 58:500.— Java, Buysmann 40 (U)

Pteris simplex 31:531.— E. New Guinea, Cruttwell 985 (K)

P. werneri (Rosenst. — Hemipteris) 42:457

Pyrrosia costata (Presl — Apalophlebia costata 1851) 47:27

[transfer previously made in 1967, with Niphobolus costatus [Presl] Pres] ex Bedd.

(1866) as basionym, by Tagawa & Iwatsuki in Act. Phytotax. Geobot.

22: 100]

P. penangiana (Hook. — Niphobolus) 24: 146

P. stictica (Kze — Niphobolus) 47:31

Rheopteris asplenioides 31:531. -SW New Guinea, Kalkman BW 8528 (L)

Rumohra puncticulata (v.A.v.R. — Polystichum) 9:137

Schizaea paucijuga 16:267. — Malaya, Haniff SFN 1098 (SING)

Schizoloma javanense (Bl. — Lindsaya) 24:349

S. tenerum (Dryander — Lindsaya) 24:348

Selliguea lima (v.A.v.R. — Pleopeltis) 24:157

Sphaerostephanos angustifolius (Pres]l— Nephrodium) 65:66

. appendiculatus (Blume — Gymnogramma) 65:62

. arbuscula (Willd. — Aspidium) 57: 164

. dichrotrichoides (v.A.v.R. — Dryopteris) 65:65

. diversilobus (Presl — Nephrodium) 65:57

. elatus (Bojer — Aspidium) 57: 167

. elatus var. thomensis 57: 167. — Sao Tomé, Moller s.n. 1885 (P)

. ellipticus (Rosenst. — Dryopteris) 65:66

. elmerorum (Copel. — Dryopteris) 65:55

. erectus (Copel. — Cyclosorus) 65:64

. fenixii 65:55. — Batanes Is., Fenix BS 3780 (P; US)

. heterocarpus (Blume — Aspidium) 62:209

. heterocarpus var. bordenii (Christ — Dryopteris) 65:67

. hirsutus (Kunze ex Mett.— Aspidium) 65:63

. lastreoides (Pres] — Pronephrium) 65:54

. latebrosus (Kunze ex Mett.— Aspidium) 62:209

. lobatus (Copel. — Cyclosorus) 65:64

. loherianus (Christ — Aspidium) 65:61

. penniger (Hook. — Nephrodium) 62:209

. magnus (Copel. — Cyclosorus) 65:61

. mindorensis 65:55. — Philippine Is., Mindoro, Price 690 (K; PNH)

. perglanduliferus (v.A.v.R. — Dryopteris) 65:59

NANNNNNANNUNUAUNNURNUNUURAURNRNURNRUNUURNN

Fern names of R. E. Holttum 235

(Sphaerostephanos cont.)

. polisianus 65:65. — Philippine Is., Luzon, Price 431 (K; PNH)

. polycarpus (Blume — Aspidium) 65:56

. porphyricola (Copel. — Dryopteris) 65:59

. productus (Kaulf.— Aspidium) 65:59

. punctatus 65:60.— Philippine Is., Luzon, Elmer 16642 (US; BO, G, K, L,

SYD)

. Sakayensis (Zeiller — Nephrodium) 62:209

. santomasii 65:62. — Philippine Is., Luzon, Price 1034 (K; PNH)

. sessilipinna (Copel. — Dryopteris) 65:53

. simplicifolius (J. Sm. ex Hook. — Aspidium) 65:61

. smithianus (Pres! — Nephrodium) 65:65

. spenceri (Christ — Dryopteris) 65:58

. stenodontus (Copel. — Cyclosorus) 65:56

. subtruncatus (Bory — Polypodium) 48:80

. tephrophyllus (Copel. — Dryopteris) 65:58

. todayensis (Christ — Dryopteris) 65:58

. unitus (L. — Polypodium) 57:165

. williamsii (Copel. — Dryopteris) 65:53

Stenochlaena cumingii 51:122.— Philippines Is., Cuming 226 (K; BM)

Syngramma minima 1:56.— Malaya, Holttum 17498 (SING)

Taenitis cordata (Gaud. — Schizoloma) 29: 112

T. dimorpha 16:274. — Malaya, Holttum SFN 17399 (SING)

T. diversifolia 46:93. — Solomon Is., RSS 4360 (K)

flabellivenia (Bak. — Polypodium) 31:533

. hosei (Bak. — Hemionitis) 46:92

lanceolata (Diels — Syngramma) 46:91

marginata 31:532.— Sumatra, Ernst 701 (Z; K)

pinnata (J. Sm.— Syngramma) 46:90

pinnata var, brachysora (Bak. —- Gymnogramme) 46:91

pinnata var. polypodioides (Bak.— Gymnogramme) 46:91. fig. 4

trilobata nom. nov. 46:93. — Lindsaya trilobata Bak. non Colenso (= L. hosei

C. Chr. non Taenitis hosei (Bak.) Holtt.)

vittarioides 42:460. — Sumatra, Korthals (L; K)

Tectaria grandidentata (Cesati — Polypodium dilatatum var.) 24:514

T. gymnosora 43:308. — Thailand, Hansen, Seidenfaden & Smitinand 11225 (C; K)

T. ingens (Atkinson ex Clarke — Nephrodium) 24:503

T. isomorpha 44:49. — Bismarck Archipelago, Noona Dan Exped. 2369 (C)

TERATOPHYLLUM Mett. emend. 12:355.— Type species: Lomaria aculeata

NANNNNNNNANHNNM ANNNNMN

T. aculeatum var. montanum 6:289, text. ff. 34, 41; tt. 1 & 4.— Malay Peninsula,

Kunstler 8309 (SING)

T. arthropteroides (Christ — Stenochlaena) 6:303. t. 12

T. brightiae (F. von Muell. — Acrostichum) 12:358. t. 28.

T. clemensiae 7:262, text-ff, 1-9. — Borneo, Clemens 31614 (SING)

T. gracile (Blume — Lomaria) 6:291. text-ff. 4, 42 t. 5.

T. koordersii 6:301. text-ff. 48 & 49. t. 11.—Celebes, Koorders 17065B (adult

fronds), 17064B and 17123B (bathyphylls) (BO)

T. leptocarpum (Fée — Lomariopsis) 10:143

T. ludens (Fée — Lomariopsis) 6:298. text-ff, 5, 23-28, 31, 35, 46, 47; t. 10

T. luzonicum 6:297. t. 9. — Philippine Is., Fenix, Bur. Sci 28272 (BO)

T. rotundifoliatum (R. Bonap. — Stenochlaena) 6:294. text-ff. 36, 37, 44. tt. 7 & 8.

236 Gardens’ Bulletin, Singapore — XXX (1977)

(Teratophyllum cont.)

T. wilkesianum (Brack. — Polybotrya) 12:359. tt. 29 & 30

T. williamsii (Underwood — Stenochlaena) 6:292, text-f. 43. t. 6

Thelypteris herbacea 16:268.— Malaya; Holttum SFN 20571 (SING)

. khamptorum 49:82.— Burma, Toppin 4217 (K)

novae-hiberniae 44:50. — Bismarck Archipelago, Noona Dan Exped. 1848 (C)

. paleata (Copel. — Dryopteris) 24:249

. motleyana (Hook. — Nephrodium) 24:247

. unidentata (Bedd. — Lastrea) 24:25]

TRIGONOSPORA gen. nov, 50:29. ff. 4, 4a. — type species: T. ciliata

T. calcarata (Blume — Aspidium) 59:506

T. ciliata (Benth. — Aspidium) 50:29

T. koordersii (Christ — Aspidium) 59:506

T. sericea (Scott ex Bedd. — Lastrea) 62:204

Vaginularia_acrocarpa 44:51.— Bismarck Archipelago, Noona Dan Exped.

2452 AMS)

Vittaria angustissima 16:274.— Malaya, Carr s.n., March 1929 (SING)

V. malayensis 3:409 cum fig. —- Malaya, Burkill & Holttum 8705 (SING)

Xiphopteris hieronymusii (C. Chr. — Polypodium) 24:218

X. sparsipilosa (Holtt.— Ctenopteris) 24:219

Bibliography

(dates of publication given here may differ from those given on title-pages)

A new fern from the Malay Peninsula. Gdns’ Bull. S.S. 4: 56-57. (1927).

Spolia Mentawiensia: Pteridophyta. J. Mal. Br. R. Asiatic Soc. 6: 14-23. (1928).

New species of ferns from the Malay Peninsula. Gdns’ Bull. S.S. 4: 408-410. (1929).

The genus Lindsaya in the Malay Peninsula, with descriptions of two new species.

Gdns’ Bull. S.S. 5: 58-71. (1930).

5 Aspidium maingayi (Baker) Holttum comb. nov. Gdns’ Bull. S.S. 5: 207-211. (1931).

6 On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan Region, Gdns’ Bull.

S.S. 5: 245-313. (1932).

7 The ferns of Mt. Kinabalu (C. Christensen & R.E.H.). Gdns’ Bull. S.S. 7: 191-324.

(1934).

8 The tree-ferns of the Malay Peninsula. Gdns’ Bull. S.S. 8: 293-320. (1935).

9 Notes on Malayan ferns, with descriptions of 5 new species. Gdns’ Bull. S.S. 9: 119-138.

(1937).

10 Further notes on Stenochlaena, Lomariopsis and Teratophyllum. Gdns’ Bull. S.S. 9:

139-144. (1937).

11 The genus Lomagramma. Gdns’ Bull. S.S. 9: 190-221. (1937).

12 A re-definition of the genus Teratophyllum. Gdns’ Bull. S.S. 9: 355-362. (1938).

13. The genus Lomariopsis in Madagascar and the Mascarene Islands. Notulae Systematicae,

Paris 8: 48-62. (1939).

14. New species of Lomariopsis. Kew Bull. 1939: 613-628. (1940).

15 The fern genus Diplazium in the Malay Peninsula. Gdns’ Bull. S.S. 11: 74-108. (1940).

16 2 uma of vascular plants from the Malay Peninsula. Gdns’ Bull. Sing. 11: 267-298.

17 A revised classification of Leptosporangiate ferns. J. Linn Soc. Bot. 53: 123-158. (1947).

18 The fern-genus Heterogonium Presl. Sarawak Museum J. 5: 156-166. (1949).

19 Further notes on the fern-genus Heterogonium Presl. Reinwardtia 1: 27-31. (1950).

20 The fern-genus Pleocnemia Presl. Reinwardtia 1: 171-189. (1951).

21 The fern-genus Arcypteris Underwood (Dictyopteris Pres] sensu Fée). Reinwardtia 1:

191-196. (1951).

22 A new Malayan Athyrium, Kew Bull. 8: 545. (1954).

hk WwWne

Fern names of R. E. Holttum 237

A new fern from Malaya (Athyrium angustipinna). Gdns’ Bull. Sing. 14: 8. (1953).

A revised Flora of Malaya vol. 2. Ferns of Malaya. Singapore: Government Printing

Office. (1955).

Some additional species of Heterogonium. Reinwardtia 3: 269-274. (1955).

Florae Malesianae Praecursores, XVI: On _ the taxonomic subdivision of the

Gleicheniaceae, with descriptions of new Malaysian species and varieties. Reinwardtia 4:

257-280. (1957).

Notes on Malaysian ferns, with descriptions of a new genus and new species. Kew Bull.

13: 447-455. (1959).

Flora Malesiana, Ser. II, Pteridophyta, vol. 1, pt. 1: i-xxiii, 1-61. (Introductory Note;

List of genera of Malayan Pteridophyta; The Morphology of Ferns; General Keys

1 & 2; Keys to Genera; Bibliography; Gleicheniaceae, Schizaeaseae). (1959).

Two problem-species: Schizoloma cordatum Gaud. and Syngramma pinnata J. Sm.

Amer. Fern J. 50: 109-113. (1960).

New species of tree-ferns (Cyathea Sm. and Dicksonia L’Herit). Kew Bull. 16: 51-64.

(1962).

New species of Malaysian ferns. Blumea 11: 529-534. (1963).

Cyatheaceae. Flora Malesiana, Ser. II, 1, pt. 2: 65-176. (1963).

The tree-ferns of the genus Cyathea in Australasia and the Pacific. Blumea 12: 241-274.

(1964),

Cytological observations on ferns from New Guinea, with descriptions of new species

(R.E.H. & S. K. Roy). Blumea 13: 129-139. (1965).

Tree-ferns of the genus Cyathea Sm. in Asia (excluding Malaysia). Kew Bull. 19: 463-487.

(1965).

Studies in the Flora of Thailand 33: Filicinae excl. of the Ophioglossaceae (second list).

Dansk Bot. Ark. 23: 227-244. (1965).

The genera Lomariopsis, Teratophyllum and Lomagramma in the islands of the Pacific

and Australia. Blumea 14: 215-223, (1966).

Two new species of Lomagramma from New Guinea. Blumea 14: 224. (1966).

The genus Arthropteris J. Sm. in Malesia. Blumea 14: 225-229. (1966).

Florae Malesianae Precursores XLIV: The fern-genus Elaphoglossum in Malesia, with

descriptions of new species. Blumea 14: 317-326, (1967).

Four new species of ferns from New Guinea. Blumea 14: 327-329. (1967).

New ferns from Malesia. Kew Bull. 20: 455-460. (1967).

Studies in the Flora of Thailand 38: Some plants new or rare to Thailand with descrip-

tions of new species: Filicinae. Dansk Bot. Ark. 23: 308-309. (1966).

Botanical Report of the Danish Noona Dan Expedition, Filicinae. Dansk Bot. Ark. 25:

35-52. (1967).

A revised Flora of Malaya vol. 2. Ferns of Malaya, ed. 2. Singapore: Government

Printing Office. (1968).

A re-definition of the fern-genus Taenitis Willd. Blumea 16: 87-95. (1968).

A commentary on some type specimens of ferns in the Herbarium of K. B. Presl.

Novitates Botanicae Inst. Bot. Univ. Carol. Prag. 1968: 3-57. (1969).

Studies in the family Thelypteridaceae 1. The genera Phegopteris, Pseudophegopteris

and Macrothelypteris. Blumea 17: 5-32. (1969).

New species of Thelypteridaceae from India, Ceylon and Burma (R.E.H, & P. Chandra).

Kew Bull. 26: 79-82, (1971).

Studies in the family Thelypteridaceae 3. A new system of genera in the Old World.

Blumea 19: 17-52. (1971).

The genus Stenochlaena J. Sm., with description of a new species. American Fern J. 61:

119-123. (1971).

Morphology and anatomy of the genera Davallia, Araiostegia and Davallodes, with a

discussion on their affinities. (T. & U. Sen & R.E.H.). Kew Bull. 27: 217-243. (1972).

Studies in the family Thelypteridaceae 4. The genus Pronephrium Pres]. Blumea 20:

105-126 (1972).

The identity of three type specimens in the Willdenow Herbarium. Amer, Fern J. 63:

81-84 (1973).

238 Gardens’ Bulletin, Singapore — XXX (1977)

55 Studies in the family Thelypteridaceae 5. The genus Pneumatopteris Nakai. Blumea 21:

293-325. (1974).

56 Studies in the family Thelypteridaceae 6. [The genera] MHaplodictyum and

Nannothelypteris, Kalikasan 2: 58-68. (1974).

57 Thelypteridaceae of Africa and adjacent islands. $. African J. Bot. 40: 123-168. (1974).

58 Additions to the Fern Flora of Java. Reinwardtia 8: 499-501. (1974).

59 The genus Trigonospora (Thelypteridaceae) in Malaysia. Reinwardtia 8: 503-507, (1974).

60 Studies in the family Thelypteridaceae 7. The genus Chingia. Kalikasan 3 : 13-28. (1974).

61 The fern-genus Pleocnemia. Kew Bull. 29: 341-357. (1974).

62 [new thelypterid nomenclatural combinations] in B. K. Nayar, A revised nomenclature

of the thelypteridoid ferns. pp 202-212 in B. K. Nayar & S. Kaur, Companion to

R. H. Beddome’s Handbook to the ferns of British India, Ceylon and the Malay Peninsula.

Koenigstein & New Delhi (1974).

63 Studies in the family Thelypteridaceae 8. The genera Mesophlebion and Plesioneuron.

Blumea 22: 223-250 (1975).

64 New Philippine ferns. Kalikasan 3: 196-197 (1975).

65 Studies in the family Thelypteridaceae 9. The genus Sphaerostephanos in the Philippines.

Kalikasan 4: 47-68 (1975).

Philippine Dryopteris

M. G. PRICE

Department of Botany

University of Michigan

Ann Arbor, Michigan, U. S. A.

Summary

A key to the thirteen recognized Philippine species of Dryopteris is presented, with

descriptions of two new species, D. uropinna and D. permagna, and four species not hitherto

reported as Philippine: D. chrysocoma, D. pulvinulifera, D. polita, and D. formosana.

Ctenitis mearnsii is reduced to Nothoperanema hendersonii.

The genus Dryopteris Adanson (nom. cons.) is here circumscribed in the

strict sense adopted by most recent authors, with its nearest relatives in Asia being

along one lineage Arachniodes and on another Nothoperanema. Dryopteris consists

of about two hundred species, mostly north subtropical and temperate, with the

centre of diversity the Himalayan-East Asian region. Distinguishing characters of

the genus are:

Rhizome short-erect, infrequently short-creeping, stipes with 5 or more

vascular bundles at base, axes grooved above, junctions between grooves of different

orders of axes open, lamina pinnate to quadripinnate, architecture usually anadro-

mous at proximal base, then catadromous upwards and outwards, or infrequently

anadromous throughout. Hairs not acicular, nor multicellular and erect, margins

not bearing long sharp cartilaginous teeth, sinuses without cartilaginous membranes

Or protuberances, veins free, not reaching margin. Sori along veins, round,

receptacles strongly raised, indusium round-reniform with a short sinus, somewhat

stalked (rarely much reduced), paraphyses usually present, spores monolete,

variously winged, bullate, or tuberculate.

Arachniodes is always rather strongly anadromous throughout, with dimidiate

pinnules (basiscopic base excised), and usually with sharp cartilaginous teeth. It

is, however, extremely difficult to formulate a verbal boundary; among Philippine

plants Arachniodes hasseltii has blunt or rounded segments and Dryopteris pulvinuli-

fera and D. subarborea are anadromous.

Nothoperanema has often been immersed in Dryopteris or Ctenitis, It agrees

with Dryopteris in soral characters but is vegetatively similar to Peranema and

Diacalpe, and thence to Acrophorus and Stenolepia. The patent stipe paleae, erect

Diacalpe, and thence to Acrophorous and Stenolepia. The patent stipe paleae, erect

conical multiseptate hairs on lamina, and fine dissection clearly mark the genus. It

might more properly be reduced to Peranema (as has Diacalpe) than Dryopteris.

Since Nothoperanema has not been reported by name from the Philippines, I

provide below a brief synonymy and description of the one Philippine species:

Nothoperanema hendersonii (Bedd.) Ching, Acta Phytotax. Sinica 11 (1966) 28.

Lastrea hendersonii Bedd., Ferns Brit. India Suppl. (1876) 17, t. 377. ——

Dryopteris hendersonii (Bedd.) C. Chr., Ind. Fil. (1905) 270; Tagawa, Acta

Phytotax. Geobot. 6 (1937) 155. —— Ctenitis mearnsii Copel., Philip. J. Sci. 81

(1952) 24; Fern Fl. Philip. 2 (1960) 289.

239

240 Gardens’ Bulletin, Singapore — XXX (1977)

Rhizome ascending, paleae brown, to 18 X 3 mm, concolorous, with minute

marginal projections. Stipes bright brown, to 70 cm long and 8 mm thick at base,

paleae linear, patent, brown, subentire, Lamina subquadripinnate, to 70 cm long,

ovate-deltoid, anadromous in sequence of pinnules at base, upwards catadromous,

rachis with reduced linear patent paleae, these grading to stout conical erect

multiseptate hairs forming a tuft inside rachis groove at junctions, also along

costae, costules, and veins both sides of lamina; texture thin papyraceous. Basal

pinnae to 30 cm long, stalk 3 cm, apex acuminate; basal basiscopic pinnule to

9 cm, secondary pinnules very deeply lobed. Middle pinnae ascending, 12-17 cm

long, acuminate, stalk 5-10 mm, basal pinnules c. 3 X 1.7 cm, secondary pinnules

adnate, blunt, shallowly lobed or crenate-serrate, apex toothed. Sori inframedial,

indusium dark brown, minutely farinose, 0.7 mm across.

LUZON. Mt Pulog, Curran et al. FB 16277 (MICH), Jacobs 7301 (A, CAHUP,

PNH); Mt Burnay, Iwatsuki et al. P-774; Benguet, Pauai, McGregor BS 8507

(MICH); Haight’s Place, Mearns BS 4216 (MICH), 4218 (MICH); Mt Singakalsa,

Celestino & Castro PNH 3971 (MICH); Mt Santo Tomas, Price 1868; Bulasan,

Copeland s.n. 7 May 1913 (MICH).

Ecology: Terrestrial in mountain forests, 1900-2700 m.

Distribution: N. India, Nepal, S. China, Japan, Thailand, Java.

Most of the Philippine species of Dryopteris are found in the Benguet highlands

of Luzon; none were collected by botanists until the 1890s. It is likely that

additional species remain to be discovered in the mountains of northern Luzon.

Aside from the delimitation of Dryopteris from Arachniodes, the greatest

difficulty still remaining is the proper treatment of the D. subarborea complex

(species 11-13 below). There are at least three Philippine taxa but their relationship

with plants from elsewhere in Malesia and Polynesia remains to be ascertained. I

am not sure that I have applied the names D. subarborea and D. purpurascens

correctly.

Instead of a full synonymy, I have provided all Philippine references, original

and basionym citations, and reference to important descriptions. Philippine

specimens I have seen are cited; duplicates of my own collections are being

distributed, Ecological information is based on Philippine conditions.

KEY TO THE PHILIPPINE SPECIES OF DRYOPTERIS

1. Indusium c. 2 mm across, persistently helmet-shaped; rhizome paleae orange-

colored

2. Fronds dimorphic (the fertile lamina contracted), deltoid-ovate, not

glandular 2102221, 234 Se Se 1. D. cochleata

2. Fronds uniform, lanceolate, bearing small capitate glandular hairs on axes,

lamina, ATIC? TOSI eh ae Sec 2. D. chrysocoma

1. Indusium exceptionally to 1.5 mm across, shriveling; paleae not orange

3. Lamina lanceolate or narrower, pinnate or bipinnatifid; basal pinnae similar

to middle pinnae

4. Pinnae toothed or shallowly lobed ........................005 3. D. hirtipes

4. Pinnae lobed nearly to COSEA, ses.) aniee a) eee 4. D. wallichiana

3. Lamina ovate or deltoid-ovate, at least bipinnate; basal pinnae more

dissected than middle pinnae, considerably longer and/or broader

Philippine Dryopteris 241

5. Lamina to c, 50 cm long

6. Lamina anadromous throughout; rhizome short-creeping, paleae

golden-brown, acicular ........................0.0008 5. D. pulvinulifera

6. Lamina catadromous above base; rhizome short-erect, paleae brown

or black

7. Stalks of basal pinnae over 15 mm long; indusium minute

len AMR Seer eamledalle: seehpeaee lip eeetet Beas 6. D. polita

7. Stalks of basal pinnae 10 mm or less long; indusium evident

8. Paleae of rhizome and stipe pale brown, very thin, lanceo-

late or ovate; rachis groove strongly papillate within ......

ee a oe ae 7. D. sparsa

8. Paleae at least partially black, not thin, linear or narrowly

lanceolate; rachis groove not papillate

9. Basal basiscopic pinnule of basal pinnae about equal

to succeeding basiscopic pinnules

9. Basal basiscopic pinnule much enlarged

10. Pinnules mostly somewhat falcate (costules ascend-

a tale sea I ahaa 9. D. varia

10. Pinnules never falcate (costules not arcuate) ......

bd hd COS ERS teers, |. 5: 10. D. formosana

5. Lamina over 100 cm long

11. Rhizome short-creeping; lamina glabrous

eres eee e eee eee eee eee er eeeeeeee

11. Rhizome short-erect; lamina with appressed multi-cellular hairs

beneath

12. Sori containing conspicuous paraphyses with large yellow

EE i oan ans 12. D. purpurascens

12. Paraphyses inconspicuous, hyaline, shorter than sporangia,

heads not or only very slightly expanded ......................5.

baboons: DYE | ate b at rcect eh dy Kocsis iiteeiaice Lo. DD.’ permagna

1. Dryopteris cochleata (Don) C. Chr., Ind. Fil. (1905) 258; Ching, Bull. Fan

Mem. Inst. Biol. Bot. 8 (1938) 434; Copel., Fern Fl. Philip. 2 (1960) 282. ——

Nephrodium cochleatum Don, Prod, Fl. Nepal. (1825) 6. D. heleopteroides

Christ, Philip. J. Sci. 2C (1907) 212; Merrill & Merritt, Philip. J. Sci. 5C

(1910) 317.

LUZON. Benguet, Bacani FB 15962 (MICH), Copeland 1837a (MICH), Hernaez

CAHUP 13898, Price 1146, 1823, 1827, Ramos BS 5813 (NY, US, MICH),

Williams 1502 (NY, US); Bontoc, Vanoverbergh 3343 (F); Ilocos Norte, Iwatsuki

et al. P-113, P-151, P-221, Price 3341.

Ecology: Rhizome short-creeping, terrestrial or on limestone, 200-1800 m, in fairly

open seasonally dry grassland, under pine forest, or at borders of secondary growth,

in sites with good drainage, cool nights, and no wind.

Notes: Copeland described D. cochleata as glabrous but the lamina beneath has

appressed multicellular hairs identical to apices of paleae.

Distribution: §, China (not Taiwan), India, Nepal, Burma, Thailand, N. Vietnam,

Java, Bali.

242 Gardens’ Bulletin, Singapore — XXX (1977)

2. Dryopteris chrysocoma (Christ) C. Chr. Ind. Fil. (1905) 257; Ching, Bull. Fan

Mem. Inst. Biol. Bot. 8 (1938) 436. —— Aspidium filix-mas var. chrysocoma

Christ, Bull. Herb. Boiss. 6 (1898) 966.

Rhizome short-creeping, paleae c. 1.5 cm long, narrow, orange-brown, con-

colorous, cells very long and slender, margins with minute regular projections.

Stipe 20 cm X 3 mm, stramineous, bearing broadly ovate paleae 5 X 2.5 mm and

reduced narrow paleae, Lamina lanceolate, 35 xX 15 cm, bipinnate throughout,

thick herbaceous, catadromous except at base, both surfaces and axes bearing erect

capitate orange glandular hairs, rachis beneath also with narrow orange-brown

paleae. Pinnae linear-lanceolate, basal slightly reduced, 17 pairs, to 9 X 2.7 cm,

abruptly narrowed to the acute apex. Pinnules c. 12 X 6 mm, sessile but not

adnate (base narrow), apices broad, rounded, with numerous small teeth. Indusia

2.5 mm across, persistent, convex, orange-brown, bearing numerous capitate orange

glandular hairs.

LUZON. Benguet, Haight’s Place, July 1907, Mearns BS 4219 (MICH).

Ecology: Terrestrial, c. 2100 m; only one Philippine collection.

Notes: Very small plants may be fertile.

Distribution: §. China, N. India & Pakistan, Burma. Reported from Taiwan by

Tagawa, Acta Phytotax. Geobot. 8 (1939) 22, but not mentioned in the recent

Flora of Taiwan.

3. Dryopteris hirtipes (Bl.) O. Ktze., Rev. Gen. Pl. 2 (1891) 813; Christ, Philip.

J. Sci. 2C (1907) 212; Merrill & Merritt, Philip. J. Sci. SC (1910) 317; Copel.

Leafl. Philip. Bot. 3 (1910) 806; Fern Fl. Philip. 2 (1960) 281; Holttum, Ferns of

Malaya, ed. 2 (1966) 635; Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 5 (1973)

6. Aspidium hirtipes Bl., Enum. Pl. Jav. (1828) 148; Christ, Bull. Herb.

Boiss. 6 (1898) 194. —— Nephrodium hirtipes (Bl.) Hook.; Copel., Polyp. Philip.

(1905) 21.

LUZON. Bontoc, Vanoverbergh 3146 (F); Mt Data, Copeland 1887 (MICH),

Price 389, 395, 402; Mt Pulog, Celestino et al. PNH 3026 (MICH), Curran et al.

FB 16278 (MICH); Mt Polis, Price 422, 436: Mt Singakalsa, Celestino & Castro

PNH 3981 (MICH); Mt Santo Tomas, Price 1006, Topping 1187 (NY); Pauai,

J. K. Santos BS 31667 (US); Benguet, Elmer 6529 (NY, US), Price 1672, Topping

248 (GH, US), 302 (GH, US), 303 (MICH, US), 3/1 (NY, US); Mt Ragut,

Loher 850 (US).

MINDANAO. Davao, Mt Calelan, Elmer 10799 (F, GH, MICH, NY, US); Mt

Apo, Copeland s.n. 27 Oct 1904 (MICH); Cotobato, Mt Matutum, Copeland s.n.

1 May 1917 (MICH).

Ecology: Terrestrial in mountain forest, 1350-2650 m.

Notes: Dimensions more variable than hitherto reported, in Philippine material

stipes 12-63 cm, lamina 14-73 cm long, pinnae 2.3-24 x 0.8-3.5 cm. Topping 311

at NY is aberrant in having irregularly anastomosing veins. Reports of this species.

from Mt Arayat (Christ 1898, Copeland 1905, 1960) are based on erroneous

localization of Loher 850, Feb. 1894, by Christ (1898); the specimen in US bears

the same date and elevation (1950 m) as Christ’s published report but with the

name Mt Ragut. Arayat is only about 1000 m high.

Distribution: Sri Lanka and India to S. China & Japan, Malesia, Samoa.

Philippine Dryopteris 243

4. Dryopteris wallichiana (Spreng.) Hyl., Bot. Notis. 1953 (1953) 352; Alston,

Amer, Fern J. 47 (1957) 92; Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 5 (1973)

8. —— Aspidium wallichianum Spreng., Syst. Veget. 4 (1827) 104. —— D. filix-

mas var. parallelogramma (Kunze) Christ, Philip. J. Sci. 2C (1907) 212; Merrill

& Merritt, Philip. J. Sci. SC (1910) 317. —— D. paleacea (Sw.) C. Chr. nom.

illegit.; Copel., Fern Fl. Philip. 2 (1960) 281.

LUZON. Mt Pulog, Curran et al. FB 16247 (MICH), Jacobs 7260 (A, CAHUP,

PNH); Mt Data, Copeland 1875 (MICH).

Ecology: Terrestrial in wet mountain forest, 2100-2800 m.

Notes: Alston (1957) distinguished the tropical American D. parallelogramma

from the Asian D. wallichiana by only the color of the rhizome scales, which I find

to be unreliable. Christensen, Cont, U. S. Nat. Herb. 26 (1931) 280, stated: “‘...a

close comparison of very complete specimens ... has shown me that it is impossible

to find one stable character by which they may be distinguished.” The genus

Dichasium (A. Br.) Fée was established for plants of this species.

Distribution: Sri Lanka, India, Nepal, Burma, S. China & Taiwan, Japan, Java,

N. Borneo, New Guinea, Hawaii, Mexico, Cuba, Jamaica, Hispaniola, Guatemala,

Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia,

Argentina, Rhodesia, Madagascar.

5. Dryopteris pulvinulifera (Bedd.)O. Ktze. Rev. Gen. Pl. 2 (1891) 813; Ching,

Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 466; Sledge, Bull. Brit. Mus. (Nat.

Hist.) Bot. 5 (1973) 10. —— Lastrea pulvinulifera Bedd., Ferns Brit. Ind. (1870)

t. 333.

Rhizome short-creeping, paleae very narrow, bright golden-brown, glossy,

c. 1 cm long, entire. Stipe bases swollen, fleshy, blackish, densely paleate; stipes

to 34 cm long. Lamina deltoid-ovate, acuminate, to 49 cm long, tripinnate +

pinnatifid at base, anadromous throughout, both surfaces with multiseptate dark

appressed hairs and unicellular glossy hyaline appressed glandular hairs, rachis

and costae with lanceolate blackish subclathrate paleae grading distally to bullate

based. Pinnae to 18 free pairs, basal pinnae to 22 cm long, basal basiscopic

pinnule to 7 cm long; larger pinnae gradually acuminate, pinnules acute, teeth of

segments acute (not aristate nor cartilaginous). Sori subterminal on veins, about

medial, indusia c. 1 mm across, reddish-brown, with multicellular hairs at base of

sinus and minute fragile appressed glossy glandular hairs elsewhere and at margins.

LUZON. Mt Data, Copeland 1861 (MICH); Mt Santo Tomas, Price 1616, 1841;

Mt Burnay, Jwatsuki et al. P-827, P-848, Price 3363.

Ecology: Terrestrial, rarely petrophytic among mosses, 1800-2200 m, on slopes

and cliffs in mossy forest.

Notes: Extremely small fronds may be fully fertile; the smallest, such I have seen,

is 4.5 cm long including stipe.

Distribution: Sri Lanka, N. E. India, Nepal.

6. Dryopteris polita Rosest., Fedde Repert. 13 (1914) 218; Holttum, Ferns of

Malaya (1955) 492; DeVol & Kuo, Fl. Taiwan 1 (1975) 378.

Rhizome short-erect, paleae glossy, light brown, subentire, concolorous, to

13 X 2 mm, Stipe to 52 cm by 3 mm, stramineous. Lamina lanceolate-deltoid,

18-45 cm long, catadromous above base, bipinnate, texture firm herbaceous,

beneath with multiseptate appressed reddish-brown hairs, axes with sparse narrow

244 Gardens’ Bulletin, Singapore — XXX (1977)

fibrils. Pinnae narrowly lanceolate, acuminate, evenly tapering, broadest on

basiscopic side; basal pinnae to 17 cm long, stalks 1.6-3.6 cm long, largest basal

basiscopic pinnule to 6.8 X 1.8 cm, lobed 4/5 to costule basally, most pinnules

4-9 mm broad with rounded apex. Sori medial, indusia minute, pale brown, com-

pletely concealed by mature sporangia, sporangial stalks dark reddish-brown

basally, paraphyses none.

MINDANAO. Bukidnon, Mt Dumalucpihan, June-July 1920, Ramos & Edano

BS 39031 (US).

Ecology: Terrestrial.

Notes: Rosenstock described this species from Sumatra as exindusiate and an

isotype in NY is indeed without indusia. However, specimens from Thailand,

S. Japan, the Philippines, and Papua clearly have very small indusia. In Fern FI.

Philip. 2 (1960) 283, Copeland cited the one Philippine specimen as an unnamed

relative of D. sparsa.

Distribution: Thailand and Sumatra to S. Japan and Papua New Guinea.

7. Dryopteris sparsa (Don) O. Ktze., Rev. Gen. Pl. 2 (1891) 813; Christ, Philip.

J. Sci. 2C (1907) 213; Merrill, Philip. J. Sci. 3C (1908) 390; Copel., Leafi. Philip.

Bot. 2 (1908) 389; Fern Fl. Philip. 2 (1960) 283; Holttum, Ferns of Malaya (1955)

492, t. 292; Hatusima, Mem. Fac. Agr. Kagoshima Univ. 5 (1966) 22, ——-

Nephrodium sparsum Don, Prod. Fl. Nepal. (1825) 6; Copel., Polyp. Philip. (1905)

24. —— Aspidium sparsum (Don) Spreng.; Christ, Bull. Herb. Boiss. 6 (1898) 194.

BATAN ISLAND. Mt Iraya, Hatusima & Sato 29055 (PNH).

LUZON. Banawe, Banlugan PNH 72907; Mt Ugo, Ramos BS 5836 (MICH, US);

Mt Santo Tomas, Price 1042, 1840; Baguio, Price 1578, Topping 222 (GH, US),

282 (MICH, US); Twin Peaks, Bartsch 211 (US); Benguet, Merrill 7779 (MICH,

US); Mt Arayat, Loher 1190 (US); Mt Banahaw, Price 362; Mt San Cristobal,

Price 804, 813, 839; Mt Makiling, Price 211, 366, 499, 2398.

NEGROS. Mt Cuernos de Negros, Elmer 9949 (F), 10100 (MICH), Herre 110]

(NY), Price 2612.

PALAWAN. Foxworthy BS 672 (MICH).

MINDANAO. Lanao, Clemens s.n. July 1907 (F); Zamboanga, San Ramon,

Copeland 1727 (MICH, US); Bukidnon, Mt Camates, Ramos & Edario BS 38607

(US); Davao, Mt Apo, Williams 2526 (NY).

Ecology: Terrestrial in mountain forests, 800-2100 m, descending to 400 m in

everwet ravines at base of Mt Banahaw.

Notes: Occasional specimens have globular sessile yellow glands on the lower

laminar surface, a character possibly controlled by a single allele. Examples

are: Herre 1101, Price 813, 1578, 2398. Price 1042 from high elevation on Mt

Santo Tomas has maroon stipe and rachis. Similar plants have been named D.

sparsa var, nitidula (Bedd.) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 472.

Distribution: Sri Lanka & India to S. Japan and Papua New Guinea.

8. Dryopteris uropinna Price, sp. nov. ——- Aspidium erythrosorum auct,

non Eaton: Christ, Bull. Herb. Boiss. 6 (1898) 193. —— Nephrodium erythro-

sorum auct. non (Eaton) Hook.: Copel., Polyp. Philip. (1905) 24. —— D.

erythrosora auct. non (Eaton) 0. Ktze.: Christ, Philip. J. Sci. 2C (1907) 211;

Philippine Dryopteris 245

Copel., Fern Fl. Philip. 2 (1960) 283. —— D. subtriangularis auct. non (Hope

G Chr. : Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 485; Tard. & C. Chr.,

FI. Gén. L’Indo-Chine 7 (1941) 315, fig. 35. —— D. labordei auct. non (Christ)

C. Chr.: DeVol & Kuo, Fl. Taiw. 1 (1975) 377, p.p.

Fig. 1

ce, holotype. A. Frond. B. Stipe scale. C. Pinnule from upper

Jani Lacy.

Fig. 1. Dryopteris urop r

i s ice Glimm

opinna Pri

pinna. (see arrow). Drawn by Ms.

246 Gardens’ Bulletin, Singapore — XXX (1977)

Species e turma D. erythrosorae, ad D. indusiatam arcte accedens, differt

magnitudine minore, ambitu subtriangulari, rhachidi glabrescenti, pinnulis late

rotundati-truncatis, apicibus pinnarum frondisque abrupte contractis et caudatis.

Ho.otypus: Luzon, Ilocos Norte, Mt Burnay, Price 3364 (MICH).

Rhizome short-erect, paleae c. 10 X 1 mm, linear, blackish with narrow brown

margins, entire. Stipes brown, 14-34 cm long, glabrescent above base, Lamina

deltoid to deltoid-ovate, 17-36 cm long, caudate-acuminate, bipinnate + deeply

pinnatifid at base, thin papyraceous, dull brown or green, costae stramineous below,

catadromous above base, above with a few narrow fibrils at axis junctions, below

with small light brown bullate based acuminate paleae along costae and at costule

bases, much-reduced septate hair-like paleae along veins. Pinnae to 16 cm long,

broadest basiscopically, abruptly narrowed and caudate, on stalks to 4 mm long,

basal pair longest; basal basiscopic pinnule slightly longer or shorter than

succeeding, 1.5—4.1 cm long, shallowly to deeply lobed. Pinnules blunt, broadest at

base but not auriculate, when undivided to 20 X 7 mm, with well-spaced small

sharp teeth. Sori medial or slightly inframedial, indusium orbicular, centrally

attached, sinus nearly obscured by overlapping edges, glabrous, shriveling but not

deciduous, to 1.5 mm across, paraphyses short, non-capitate, on sporangial stalks,

spores dark brown, rather irregularly tuberculate.

LUZON. Lepanto, Bagnen, Copeland 1929 (MICH); Ilocos Norte, Mt Burnay,

Iwatsuki et al, P-849, Price 3364. Christ (1898, 1907) also cited a Loher collection

from Mt Santo Tomas, Feb. 1894, 2250 m.

CHINA. Kwangsi, Ching 6263 (NY, US); Yunnan, H. T. Tsai 60474 (GH).

INDIA. Assam, Khasi Hills, 4000 ft., Oct. 1890, G. Mann (NY, US).

THAILAND. Udawn, Tagawa et al. T-645, T-1297 (US).

VIETNAM. Cochin-China, Cavalerie s.n. ex Herb. Jeanpert (US).

TAIWAN. Taito, Tagawa 2670 (MICH); Agyoku, Taihoku, Suzuki 24 (MICH).

OKINAWA. Mt Yonahadake, Hatusima 18371 (US).

RYUKYUS. Ishigaki, Koidzumi, s.n. July 1923 (US).

JAPAN. Kyushu, Yakushima, Iwatsuki 3159 (US).

Ecology: Terrestrial, 1800-2250 m in sheltered well-lighted spots in moist montane

forest.

Notes: Hope cited two specimens of Gustav Mann with his Nephrodium

subtriangulare, neither of which was seen by Ching (1938) who mistakenly named

a Clarke specimen as type. Hope’s description differs from Ching’s plant (D.

uropinna) by rhizome scales castaneous, “‘pinnae all except lowest pair very

slightly narrowed at base, lowest inferior pinnules on lowest pinnae very much

shorter than next three or four.” In Philippine literature this species has been known

as D. erythrosora which however has a creeping rhizome, brown paleae, and much

reduced basal basiscopic pinnules of basal pinnae, among other differences. D. |

indusiata differs by attaining larger dimensions, lanceolate outline, more strongly

paleate stipe and rachis, segments narrower apically, frond apex and pinna apices

not suddenly contracted and caudate. D. labordei has prominently enlarged basal

basiscopic pinnules of basal pinnae, brown (not black) paleae, and gradually

narrowed apices. The specimens above cited from Thailand and Japan differ in

minor respects from the Philippine type. Some Japanese specimens appear to

Philippine Dryopteris 247

intergrade with D. indusiata; they may be hybrids. Another similar species is

represented by Topping 1794 (US) from Mt Kinabalu; it differs by bullate scales

absent, segments hardly toothed, basal basiscopic pinnule not dissected.

Distribution: Himalayan region to S. Japan and Luzon.

9. Dryopteris varia (L.) O. Ktze., Rev. Gen. Pl. 2 (1891) 814; Ching, Bull. Fan

Mem. Inst. Biol. Bot. 8 (1938) 490; Copel., Fern Fl. Philip. 2 (1960) 284.

Polypodium varium L., Sp. Pl. 2 (1753) 1090. —— Aspidium varium (L.) Sw.;

Christ, Bull. Herb. Boiss. 6 (1898) 192; ibid. II, 6 (1906) 996. Polystichum

varium (L.) Presl; Copel., Polyp. Philip. (1905) 18.

LUZON. Bontoc, Madulid et al. PNH 113264, Vanoverbergh 624 (MICH); Ilocos

Norte, Iwatsuki et al. P-202, P-225, P-886, Price 3342, 3398; Baguio, Copeland

1959 (MICH), Elmer 6489 (MICH, NY, US), Hernaez CAHUP 13899, 13900,

13901, Price 386, 1677, Ramos BS 5795 (MICH, US), Topping 177 (US),

Williams 1513 (NY, US); Mt Banahaw, Price 882.

Ecology: Terrestrial in seasonally dry forest, 700-1500 m.

Notes: The Banahaw collection is a new southern range extension.

Distribution: N.E. India, China, Korea, Japan.

10. Dryopteris formosana (Christ) C. Chr., Ind. Fil. (1905) 266; DeVol & Kuo,

Fl. Taiwan 1 (1975) 376. —— Aspidium formosanum Christ, Bull. Herb. Boiss.

II, 4 (1904) 615.

Rhizome short-erect, paleae fuscous, concolorous, subentire, to 20 x 2.5 mm.

Stipes stramineous to brown, to 49 cm long, 6 mm thick at base, paleae blackish,

entire, Lamina deltoid or deltoid-ovate, to 42 cm long, fully tripinnate at base,

firm papyraceous, greyish-green, anadromous at proximal base, elsewhere cata-

dromous. Rachis with reduced narrow blackish paleae, grooves with conical hairs

at junctions, pinna-rachises, costae, and costules with small dark acuminate bullate-

based paleae beneath, veins beneath with appressed multiseptate hairs. Basal pinnae

to 19 cm long, stalk 9 mm, basal basiscopic pinnule notably enlarged, to 13 cm,

secondary pinnules to 3.5 X 1.1 cm, tertiary pinnules to 8 X 3.5 mm, adnate,

minutely aristate; basal acroscopic pinnule to 3.2 xX 1.7 cm. Middle pinnae c.

13 cm long, acuminate, pinnules to 3.4 cm long, secondary pinnules adnate or

confluent, aristate-denticulate. Sori about medial, indusium bright brown, to 1.3 mm

across, glabrescent.

LUZON. Benguet Prov., Mt Santo Tomas. Price 1117; Ilocos Norte Prov., Solsona,

Price 2929, Mt Burnay, Iwatsuki et al. P-851, Price 3384.

Ecology: Terrestrial in mountain forest, 1400-2100 m.

Distribution: Taiwan, Japan.

11. Dryopteris subarborea (Bak.) C. Chr., Ind. Fil. (1905) 295; Holttum, Ferns

of Malaya (1955) 491, t. 291; Copel., Fern Fl. Philip. 2 (1960) 284, p.p.

Polypodium subarboreum Bak., J. Linn. Soc. Bot. 24 (1887) 259. —— D. purpuras-

cens auct, non (BI.) Christ: Christ, Philip. J. Sci. 2C (1907) 213.

Rhizome short-creeping, c. 2.5 cm thick, paleae to 20 X 2 mm, brown, con-

colorous, with marginal projections from one or two adjoining cells bearing a small

globose pale deciduous gland. Stipe stramineous, sparsely paleate, to at least 55 cm

Jong, 12 mm thick at base, with c. 12 vascular bundles forming a single ring

interrupted by the adaxial groove. Lamina to at least 110 cm long, ovate, to fully

quadripinnate basally, anadromous throughout or nearly so, thin papyraceous,

248 Gardens’ Bulletin, Singapore — XXX (1977)

rachis with a few scattered appressed remnants of paleae, rachis groove, pinna-

rachis grooves, and basal portion of pinnule grooves with minute erect capitate

glandular hairs within, lamina otherwise glabrous. Basal pinnae to 60 cm long,

stalk 1.5 cm to basal pinnule, apex acuminate, pinnules to 17 cm long, acuminate,

secondary pinnules to 4.5 X 1.6 cm, blunt or acute, the proximal pinnate or lobed

with rounded undulate-toothed segments, grading distally to shallowly crenate-

dentate. Middle pinnae c. 32 cm long, acuminate, stalk 1 cm, basal pinnules

7.5 X 3 cm, secondary pinnules to 1.6 X 0.9 cm. Sori medial to slightly supramedial,

indusium brown, glabrous, smaller than sorus, 0.5 mm across, paraphyses with

hyaline or yellow bead-shaped heads.

LUZON. Quezon, Real. Llavac, Price 1494, 1499; Laguna, Mt San Cristobal,

Copeland PPE 91 (GH, MICH); Mt Makiling, Elmer 18067 (F, GH, NY),

Price 2988.

MINDORO. Mt Halcon, Merrill 6101 (MICH)?

CATANDUANES. Ramos BS 30192 (US), 30223 (US), 30462 (US).

SAMAR, Gandara, Price & Hernaez 158.

PANAY. Capiz, Mt Bulilao, Martelino & Edafio BS 35645 (US), 35652 (US),

35693 (US), 35730 (US).

Ecology: Epiphytic on trunks and large branches in wet forest, 300-1200 m.

Notes: I place here with doubt Merrill 6101 known to me by a single sterile

specimen pustular beneath. According to the field label, a Bornean specimen at

Kew, matching the type, was also epiphytic with short-creeping rhizome (R. E.

Holttum, in litt.). Related is the New Guinean D. pseudoparasitica v.A.v.R. with

relatively slender and long-creeping rhizome and dimorphic fronds.

Distribution: Sumatra, Malaya, Borneo (Java?).

12. Dryopteris purpurascens (B].) Christ, Philip. J. Sci. 2C (1907) 213, nomen

tantum; Copel., Leafi. Philip. Bot. 3 (1910) 807; Backer & Posthumus, Varenflora

voor Java (1939) 47, p.p. Aspidium purpurascens Bl., Enum. Pl. Jav. (1828)

169. —— Nephrodium divisum auct. non Hook.: Copel., Polyp. Philip. (1905)

26. —— D. subarborea auct. non (Bak.) C. Chr.: Christ, Philip. J. Sci. 2C (1907)

214, p.p.; Copel., Leafl. Philip. Bot. 3 (1910) 807; Fern Fl. Philip. 2 (1960)

284, p.p.

MINDANAO. Mt Apo, Copeland 1138 (MICH, NY, US), 16/4a (MICH), s.n.

5 Sept. 1932 (MICH), Elmer 11418 (MICH), 11550 (F, GH, MICH, NY, US),

11842 (F, GH, MICH, NY, US), Williams 2479 (NY, US); Mt Matutum, Cope-

land s.n. 30 April 1917 (MICH), Ramos & Edafnio BS 85302 (GH).

Ecology: Terrestrial, petrophytic, or epiphytic on mossy trunks, 1500-2000 m.

Notes: Elmer 11842 has a different appearance because of maroon color when

dry and finer teeth of segments; it may represent another species. Mindanao

specimens which I am treating as D. purpurascens differ from the next species, D.

permagna, by being somewhat laxer and less dissected with broader usually deltoid

teeth and inconsistently catadromous or anadromous architecture, as well as the.

paraphyses. The holotype of Aspidium purpurascens Bl. (photo MICH, US, Morton

neg. 1304) consists of two separate pinnae evidently from the same frond. labeled

as collected in the Moluccas by Zippelius. Blume’s variety Beta (photo MICH, US,

Morton neg. 1305) seems to be identical, possibly parts of the very same frond,

consisting of a frond apex and the basal portion of a basal pinna, but is labeled

Java, collector Blume. Both are quadripinnate, and anadromous throughout. While

Philippine Dryopteris 249

no Philippine materials are an exact match to the type of D. purpurascens, I refrain

from describing the Mindanao plant as new without further information. Among

other subtle differences, Javan plants have relatively numerous persistent narrow

paleae along axes below.

Distribution: Java, Celebes, Moluccas (Borneo?).

13. Dryopteris permagna Price, sp. nov. —— D. subarborea auct. non (Bak.)

C. Chr.: Christ, Philip. J. Sci. 2C (1907) 214, p.p.; Copel., Fern Fl. Philip. 2

(1960) 284, p.p.

Rhizoma crassum, erectum, paleis usque ad 42 X 7 mm cellulis marginalibus

aliquot eminentibus ferens. Stipites usque ad 103 X 1.2 cm. Lamina subquinque-

pinnata, ovata, usque ad 138 cm longa, herbacea. Pinnulae primariae pleraque in

positione catadromica portatae. Pinnae basales amplissimae, usque ad 64 cm

longae, petiolus usque ad 10 cm longis, pinnula basali basiscopici usque ad

31.5 cm longa. Pinnulae ultimae apice rotundatae denticulatae, lateribus lobatae vel

dentatae, lobis acutis erectopatentibus usque fere patentibus. Costae, costulae

venaeque infra pilis persistentibus multiseptatis appressis praeditae. Sori medialis,

indusia 0.5-1 mm diametro, glabra. Paraphyses inconspicuae, hyalinae, non

capitatae,

HoLotypus: Luzon, Mt Santo Tomas, Price 16J]] (PNH).

Rhizome stout, erect. Paleae to 42 X 7 mm, long-acuminate above a relatively

broad lower portion, brown, concolorous, margins with small blunt projections

consisting of the outcurved portion of one cell or two adjacent cells. Stipe light

brown, with smaller narrower paleae, to 103 cm long and 12 mm thick, with 15

vascular bundles near base, the two adaxial on either side of the stipe-groove much

the largest. Lamina ovate, to 138 cm long, olive-green, herbaceous, quadripinnate or

subquinquepinnate at base, pinnae and major pinnules long acuminate, most or all

pinnae with basal primary pinnules in catadromic position, secondary pinnules in

anadromic position except sometimes for the larger ones of basal pinnae, tertiary

pinnules and quaternary segments nearly all in anadromic position. Major axes

beneath with deciduous narrow fibrils, persistent appressed reddish-brown multi-

septate hairs c. 0.3 mm long borne along costae, costules and veins beneath,

sometimes a few on upper surface. Basal pinnae largest, to 64 cm long on stalks

to 10 cm long, basal basiscopic pinnule to 31.5 cm, stalk to 1.8 cm, not very

much larger than the succeeding, basal acroscopic pinnule to 20.5 cm, very slightly

shorter than the several succeeding. Largest secondary pinnule 10 cm long, tertiary

pinnules to 2.5 X 1.7 cm, largest quaternary pinnules broadly adnate or connate,

to 10 X 5 mm. Third pair of pinnae from base 45 cm long, fifth pair 33 cm long

with stalk 2 cm, tenth pair 12 cm long, ultimate pinnules with broad blunt

denticulate apices and spreading-ascending pointed lobes or teeth. Sori about medial,

indusia 0.5-1 mm across, brown, glabrous, paraphyses shorter than sporangia,

apices hyaline, not or hardly expanded.

LUZON. Mt Polis. Oliver PNH 30576; Mt Data. Ramos & Edario BS 40285 (US):

Mt Pauai, Sulit PNH 5120 (MICH): Bontoc, Ramos & Edanio BS 37834 (US),

37847 (US), 37985 (US), 38064 (US); Mt Ugo. Ramos BS 5777 (MICH, NY,

US); Haight’s Place, Mearns BS 4194 (MICH, US); Baguio, Loher 832 (US); Mt

Santo Tomas, Price 1611, 1862.

Ecology: Terrestrial in mountain forest, 1400-2200 m.

Notes: Ramos & Edafio BS 37847, very congested and nearly glabrous, is placed

here with some doubt. This species, apparently endemic to Luzon, is very similar

to Polynesian specimens of D. arborescens (Bak.) O. Ktze., which is characterized

by broad pale brown paleae on costae, the paleae attached parallel to the costa.

250 Gardens’ Bulletin, Singapore — XXX (1977)

Another similar Polynesian plant, D. maxima (Bak.) C. Chr., is anadromous,

very finely dissected, with supramedial sori. Some New Guinean specimens

are similar to those of Polynesia; none are quite the same as anything in the

Philippines.

Acknowledgements

I am grateful to the directors and curators of the following institutions for

the use of facilities, loan of specimens, and other forms of assistance: A, C 4

F, GH, KYO, MICH, NY, PNH, SING, TAI, US. I am especially indebted to

Prof. T. J. Rimando of the University of the Philippines at Los Bafios for much

help when collecting in the Benguet highlands of Luzon.

Fertile-sterile Leaf Dimorphy in Ferns

Warren Herb Wagner, Jr. and Florence S. Wagner!

Department of Botany, University of Michigan

Ann Arbor, Michigan, U.S.A.

In addition to his many other contributions, Dr. R. E. Holttum has been

unquestionably one of the world’s leading authorities on the taxonomy of ferns with

fertile-sterile leaf dimorphy, especially the lomariopsid group. Inspired in part by

reading his papers, we became interested several years ago in the over-all subject

of fertile-sterile dimorphy and began the study to be reported here. As will be

seen, there are numerous disagreements in the interpretations of dimorphy, and

this represents an effort to review these as well as other aspects of the general

phenomenon.

Dimorphy of fertile and sterile fronds is widespread and bears upon both the

systematics and ecology of ferns. It is an expression of a trend found throughout

vascular plants, a trend which achieves its ultimate in seed plants. Recognizing

that the term “‘dimorphy” may sometimes apply to other than fertile-sterile (i.e.,

various types of heterophylly of vegetative leaves in ferns), for convenience the

term here will be confined to differentiation into sporophylls (spore-producing

fronds, fertile Blatter) and trophophylls (purely photosynthetic fronds, Laub-

blatter). Profound alterations may appear in the evolutionary transition from

primitive monomorphy, including transformations in morphology, anatomy, physio-

logy, and ecology. These changes are thought to possess adaptive value. However,

this idea is not universally accepted, and there is little or no experimental evidence

to support it. At present, in fact, most of our conclusions are intuitive and hypo-

thetical, and we strongly need rigorous tests. We hope that the ideas we present

here will help to stimulate such experimentation.

As many as one in five of all fern species possess what may be called fertile-

sterile dimorphy, but this figure has to be arbitrary because of so many transitions

that occur between monomorphy and dimorphy. Where the dimorphic condition

is strongly expressed, taxonomists have traditionally utilized it as a useful character

in species discrimination. In some instances, taxonomic separations have been

created even at the generic level on the basis of sporophyll-trophophyll dimorphy.

The analysis of foliar dimorphy invokes a multitude of biological questions

such as what parts undergo changes, what are the effects on sporangial arrangement,

what developmental processes underlie formation and timing of fertile expressions,

what — if any — ecological advantages are conferred, and to what extent

dimorphism is valid as a systematic character, The following report is an outcome

of our observations in the field, laboratory, and herbarium, together with a survey

of the literature. The most thorough and most recent survey seems to be that of

F. G. Dickason in 1946. Because of the widely scattered nature of references to

this subject, we fully expect that some important items have been overlooked. We

also anticipate that various ideas expressed here will be subjected to future

1 Research carried out in connection with National Science Foundation Grant DEB-75-

03550, ‘Evolution and Taxonomic Applications of Foliar Characters.” Various individuals

and institutions made available specimens for this study, especially Michael G. Price. Janice

Glimn Lacy provided many of the microscope preparations and the diagrams, and various

individuals aided in the development of this manuscript. To all we express our warm thanks.

251

252 Gardens’ Bulletin, Singapore — XXX (1977)

reappraisal, If, however, this article serves to stimulate critical research, its purpose

would be fulfilled.

All degrees of morphological and anatomical divergence exist between fertile

and sterile fronds. In fact, one of the reasons the term “‘monomorphic”’ is rarely

used may be that practically all ferns are strictly speaking at least slightly

dimorphic. Steps in dimorphic divergence are well exemplified in numerous genera.

In eastern North America, for example, we encounter a good illustration in the

woodfern genus, Dryopteris. D. goldiana (Hook.) Gray has hardly any detectable

difference between soriferous and non-soriferous mature fronds. D. cristata (L.)

Gray, with erect, tall fertile fronds, and spreading, short sterile fronds, shows

strong differentiation. D. spinulosa (O. F. Muell.) Watt is intermediate, Other good

examples of such sequences are met with over a broad taxonomic spectrum, such

as, in Polypodioideae, Loxogramme, Microgramma, and Pyrrosia; in Blechnoideae,

Blechnum; and in Dryopteridoideae, Davallia, Humata, Nephrolepis, Oleandra,

and Polystichum?.

It should be understood that the term ‘“‘dimorphism”’ does not necessarily

imply a precise morphological parallelism. The changes involved are probably

often the result of analogous, rather than homologous, transformations, and different

examples show different degrees of involvement of petioles, blades, pinnae,

sporangial arrangements, histology, and physiological reactions.

The sporophyll-trophophyll differentiation may embrace whole fronds or only

parts of fronds, and in the latter, it may be the tip that is fertile, the middle of the

frond, or the base. These modifications are diagrammed in Figure 1. Interfoliar

dimorphy, constituting complete differentiation of whole fronds is the most

common type. However, because of the commonly transitional nature of the

manifestations, many fertile fronds are merely “‘subdimorphic” (term as used by

Dickason, 1946) in the sense that the changes displayed are incomplete in kinds

and amounts. A “holodimorphic”’ situation includes total loss of vegetative function

in the sporophyll. However, the latter is hard to achieve because during the early

stages of maturation the rachises and costae of all fertile fronds (at least those

known to us) are to some extent green and photosynthetic. Whole-frond differentia-

tion into sporophylls may follow two routes. It may be the culmination of an

evolutionary trend in which at first only the frond tip is fertile. Dickason (1946)

suggests that there is a progressively earlier shift from the vegetative to the

reproduction condition of the leaf starting at the apex (i.e., Fig. 1, B to D) mediated

by an hypothetical “‘sporogen” growth substance. On the contrary, one can just

as well imagine modification of the whole leaf progressing from the inception of

the leaf, bottom to top.

Where fertile-sterile differentiation appears in separate parts of the same leaf,

“the term hemidimorphism is used instead of dimorphism’’ (Dickason, 1946). The

hemidimorphic leaves with terminal fertile pinnae are the most widespread. In

Osmundaceae, this condition is illustrated by the familiar O. regalis L.; in Poly-

podioideae, by the genera Paltonium and especially Belvisia; in Grammitidoideae,

by Grammitis Sect. Xiphopteris; and in Adiantoideae, by Llavea. Two extremes in

the Dryopteridoideae are observed in the eastern North American Polystichum

acrostichoides (Michx.) Schott and Dryopteris ludoviciana (Kze.) Small. As in

other forms of hemidimorphism, the fertile pinnae even when they are

photosynthetic at the time of spore production as they are in D. ludoviciana ——.

are shorter-lived than the sterile, and turn brown and die well before the sterile

pinnae.

Examples of hemidimorphism in which only the medial pinnae are the fertile

ones are known best in Osmunda, and most familiarly in O. claytoniana L.. which

is designated in North America by the colloquial name of “‘interrupted fern”

2 The family-subfamily classification used in this paper is that of W. Wagner (1973).

Leaf Dimorphy in Ferns 253

because its fertile pinnae discharge their spores and fall off early in the season

(usually by early or middle July), leaving fronds in which the lower and upper

regions of photosynthetic pinnae are set apart by a length of naked rachis.

Ferns with the hemidimorphic condition effecting only basal pinnae are

taxonomically a motley lot. In the striking cyatheoid genus Thyrsopteris, known

in the living flora only from Juan Fernandez Islands, several pairs of lower pinnae

are fertile, these extremely skeletonized (i.e., lacking lamina) but bearing sori with

conspicuous cup-like indusia. More abundant ferns with modified basal pinnae

occur in Ophioglossaceae and Schizaeaceae subg. Anemioideae. In the genus

Anemia we find a spectrum running from only subtly subdimorphic basal pinnae

all the way to fully differentiated basal pinnae. Accompanying this progressive

laminar reduction and skeletonization of the basal pinnae pair, there is increased

tendency for them to become erect and tower above the spreading sterile pinnae.

In the most evolved forms in Anemia, the sporangial masses are held well above

the photosynthetic portions of the leaves, and effectively the same result is achieved

as in those dimorphic leaves in which the sterile fronds are flat and form rosettes

(cf. Fig. 1, C, hemidimorphic, with Fig. 2, D, holodimorphic). The alterations in

Anemia have been described in some detail by Mickel (1962, 1967). One of the

interesting features in specialized members of this genus is the close association

of the fertile basal pinnae in point of apparent origin with the second pair of

pinnae, i.e., the lowest of the sterile pinnae, a condition to which Mickel refers as

“fertile pinnae approximate to sterile.” On the basis of comparative morphology,

we can conclude that the fertile petiolule bases have fused with the main petiole

up to the first sterile petiolules. A parallel condition is seen in the single fertile

spike of certain Ophioglossaceae, e.g. Botrychium virginianum (L.) Sw.

Mickel (1967) describes the probable evolution of the fully fertile fronds of

certain species of Anemia as following a pathway opposite that suggested by

Dickason (see above) for other holodimorphic ferns. Most Anemia taxa produce

a single pair of fertile basal pinnae, but in A. rutifolia Mart. there is more than

one pair of basal fertile pinnae, and in A. millefolia Gardn. all of the pinnae are

fertile. We do not believe that the two models are necessarily in conflict, nor the

idea, suggested earlier, that in some cases the sporophyll may have been holo-

dimorphic from the beginning. There are thus three different possible routes: (a)

change directly from monomorphic leaves to holodimorphic (Fig. 1, A-D); (5)

change from monomorphic to hemidimorphic, with the fertile pinnae apical, to

holodimorphic (Fig. 1, A-B.D); and (c) from monomorphic to hemidimorphic

with the fertile pinnae basal to holodimorphic (Fig. 1, A-C-D). Monographers

should be prepared to encounter any one of these routes in a given evolutionary

line.

Morphologically the single fertile spike of Ophioglossaceae (Botrychium,

Helminthostachys, Ophioglossum) resembles the fertile basal pinna condition in

Anemia, differing from it in the apparent fusion to form a single primordium and

mature structure. This interpretation ——— the ‘“‘Fusion Theory” or “‘Peltation

Theory’’ —— to explain the fertile spike, is the most parsimonious and requires

the least number of assumptions *. Unfortunately, the morphological intermediates

that would directly confirm the phylogeny of the fertile spike have been lost from

the fossil record. All we have are data from living forms to support this interpreta-

tion ——— the pattern and origin of vascular strands, and the occasional grapefern

specimens (especially in Botrychium subg. Sceptridium, considered primitive on

other grounds) of the Anemia condition with separate fertile basal pinna pairs, as

3In the welter of interpretations to which the fertile segment of Ophioglossaceae has been

subjected, one of the most unusual is that of Wieffering (1964), who, regarding Ophioglossum,

treats the sterile segment of the frond as a “trophophyll’ and the fertile spike as comprising

a “fertile stalk” terminated by a “strobilus”!

254 Gardens’ Bulletin, Singapore — XXX (1977)

well as intermediates, There are a few examples of adder’s-tongue ferns, Ophio-

glossum, with the sterile blade reduced or suppressed entirely. However, the belief

that the rare South African species, O. bergianum Schlecht. has complete sporophyll-

trophophyll separation (as believed, for example, by Copeland, who keyed it out

on this basis in 1947) has proved to be erroneous. In our own observations of this

curious plant we find an illusion of two separate fronds that is created by a very

low attachment of the fertile spike to the sterile segment.

Although the details are not agreed upon, the consensus seems to be that the

sporocarps of the Marsileaceae are modified basal pinnae in which the sporangia

are encapsulated. Unless it can be established that the sporocarps are de novo in

origin, comparative morphology dictates that the homology to basal pinnae is the

most likely explanation of these very distinctive structures, especially if it is agreed

that Marsileaceae are specialized leptosporangiate ferns. Gupta (1956, 1962) has

brought together much of the information bearing upon the nature of the sporocarp.

Also, he has found (pers. comm.) very interesting forms of Marsilea minuta L.

which have numerous sterile pinnae borne along the leaf axis, suggesting a reversal

to a more primitive condition. It is easy to speculate that the hemidimorphic

condition of Marsileaceae may have more than a coincidental resemblance to that

of the Anemioideae.

Attention should be called to the less spectacular but substantial changes that

often occur in the specialization of distinct trophophylls. The obviously striking

aspect of many sporophylls quickly attracts attention from morphologists and

taxonomists, but often the trophophylls themselves do not remain merely as sterile

remnants but little changed from the original double-duty leaves of the mono-

morphic ancestors. The uncommon curly-grass fern, Schizaea pusilla Pursh, of

northeastern North America has trophophylls which are wiry, twisted organs

concentrated at the base of the plant. In various polypodioid genera, such as

Microgramma in the New World and Pyrrosia in the Old, there seems to be an

inverse correlation between changes in the sporophyll and those in the trophophyll:

As the sporophyll becomes longer and narrower, the trophophyll becomes shorter

and wider. Holttum (1954) illustrates Pyrrosia nummularifolia (Sw.) Ching, an

extreme example in its genus with the sterile fronds “‘very shortly stalked, almost

circular to broadly ovate.” In this example, the fertile fronds are more similar to

the norm of the non-dimorphic or subdimorphic fronds of the genus than are the

sterile. Figure 2, D, illustrates this type of change in trophophylls.

In a few of the paddle-ferns, Elaphoglossum, the trend described above is

reversed: The trophophylls become linear, extremely long and narrow, and pendent,

while the sporophylls retain their lanceolate outlines. Splendid illustrations of such

specialized hanging trophophylls are the New World tropical Elaphoglossum

siliquoides (Jenm.) C. Chr. and especially E. herminieri Bory ex Fée.

In teleological terms, the “‘need’’ for certain leaf shapes required by carrying

out both duties of photosynthesis and sporulation seems to be eliminated if

sporophylls evolve, permitting the trophophyll to become more adapted to its role.

In Dryopteris cristata, the sterile leaves are smaller and more spreading, so that they

form a “‘light-catching” rosette at the base of the plant in contrast to the bulk of

the species of Dryopteris in which the sterile fronds are less differentiated from the

soriferous fronds. In general, in terrestrial forms, the flatter and rounder the

trophophyll, the more efficient it will be photosynthetically. This seems to be true

also in those epiphytic ferns, such as the creeping polypodioids, in which the

trophophylls “hug” the branches and twigs of the host trees and shrubs.

A somewhat neglected component of comparative morphology of the sporophyll

is the modification of sporangial distribution. The over-all trend is toward the

concentration of sporangia, but this concentration is achieved in a variety of ways.

The sori may keep their organization and simply become more tightly grouped.

Leaf Dimorphy in Ferns 255

a( Hh

Figure 1. Diagrammatic representation of fertile-sterile dimorphy. 4. Prmitive condition:

monomorphic. D, F. Complete dimorphism of sporophyll and trophophyil, showing different

relative sizes of sporophylls. B,C, E. Hemidimorphic conditions: B Fertile parts terminal

E. Fertile parts medial. C. Fertile parts basal.

They may, on the other hand, lose their soral organization and fuse into a con-

tinuous “‘sheet” of sporangia spread over the sporophyll lamina, or the entire

sporangial area may become skeletonized, losing lamimar webbing completely and

producing clusters of single-veined axes. The sporangia may be exposed, even m

the youngest stages of development (as in certain acrostichoid ferns). or they

may be protected by indusia, paraphyses, or by inrolling of laminar tissue to form

more or less persistent sheltering capsules. Also, these various modifications may

be combined in different ways.

Compacted but still discrete sori are observed in diverse taxonomic groups.

The exposed epidermal areas between the sori are reduced so that finally no bare

can be seen and the sori are in contact on all sides. For polypodioid sori

this is well illustrated in the genus Pyrrosia. While it is true that after spore

discharge members of this genus may appear to have truly acrostichoid soriation,

256 Gardens’ Bulletin, Singapore — XXX (1977)

Figure 2. Diagrammatic representation of relative differentiation of sporophylls and

trophophylls, A. Primitive condition: monomorphic. B,C, D, Dimorphic. C. Extreme change

in sporophyll, the characters (height, orientation, exaggerated). D. Extreme change in

trophophyll, specializations of size, shape, and orientation strong.

this is actually an illusion. Sori of the ancestors of Pyrrosia were probably of the —

type seen today in Pleopeltis well separated, exindusiate, and round. An

outstanding example of nearly confluent indusiate sori is present in the eastern

North American Polystichum acrostichoides (L.) Schott. Here the situation

resembles that in Pyrrosia in that after spore discharge the fertile pinnae become

falsely acrostichoid, Prior to spore discharge, however, not only are the discrete

sori recognizable, but each is provided with a conspicuous peltate indusium.

Leaf Dimorphy in Ferns 257

Truly acrostichoid soriation results from the origin of sporangia upon a

continuous placental surface which extends across the abaxial surface of the blade

or pinna. Even in young stages it is not possible to make out individual sori.

Although there are some disagreements over details of interpretation and termi-

nology (cf. W. Wagner, 1965, pp. 299, 300). recognition of this type of soriation

is generally accepted as a valuable tool in generic taxonomy. Ferns with acrosti-

choid soriation may or may not have laminar contraction associated with it. In

Elaphoglossum (Aspleniaceae subg. Dryopteridoideae) and Acrostichum (Adianta-

ceae), the fertile blades or pinnae may be only slightly contracted or not contracted

at all. However, they tend to be more ephemeral than the sterile blades or pinnae,

becoming curled and dry before the sterile segments. Acrostichoid soriation in

unrelated taxa probably has had different modes of origin. In Acrastichum the

ancestors were very likely gymnogrammeoid ferns lacking indusia. The sori were

linear and followed the veins, and they were separated by exposed laminar surface.

By extension of the sori into the naked areas and finally becoming confluent, the

acrostichoid condition came about, the sporangia making up a continuous sheet,

clothing the entire laminar undersurface. On the contrary, the history of Elapho-

glossum may have followed a very different route. If we accept the comparative

evidence from many characters, Elaphoglossum is a derivative of ferns with round,

indusiate sori (like those of Dryopteris and Polystichum). The steps in the evolution

of Elaphoglossum from its probable progenitors may then be suggested to have

followed this route: (1) assumption of the epiphytic habitat, (2) simplification

of the frond, divided to undivided, (3) approximation of sori, from remote to

continuous (as seen today in Polystichum acrostichoides). (4) loss of indusia, and

(5) unification of placental tissue all across the laminar surface to become acrosti-

choid. Some of these steps may have taken place in a different order. The

important point is that the acrostichoid condition may be the result of convergent

evolution and not necessarily be a signal of systematic affinity. Surely the examples

known in the polypodioid ferns (e.g., Belvisia, Christiopteris, Dendroglossa, and

Platycerium) arose from very different ancestors from those given above.

Bower pointed out long ago (1923) that some acrostichoid ferns have “no

special vascular provision for the nutrition of the sporangia”, whereas others

develop elaborate modifications involving two parallel vascular systems ——— the

normal venation plus a specialized receptacular venation on the abaxial side. The

two are in connection with each other, of course. The state of having two parallel

vascular systems is termed “diplodesmic”, and is especially well shown by

acrostichoid examples in the Polypodioideae (Bower, 1928).

What we here call the botryoidal condition, in which the sporangia form

clusters upon more or less naked and branched axes, is best shown in three groups

—Ophioglossaceae, Schizaeaceae subg. Anemioideae, and Osmundaceae. Here

the flat laminar condition is partially or entirely lost, and the sporangial masses

become three-dimensional and glomerulate. Assuming that lamina-borne sporangia

are ancestral (see discussion below) we may conclude that the primitive condition

in Osmundaceae is that displayed by Todea and in Anemioideae by Mohria or

certain members of Anemia itself. The only presumed primitive or intermediate

conditions found in Ophioglossaceae are occasional forms or terata in which the

sporangia are borne upon partly or fully developed lamina. This is especially well

shown in certain species (e.g., Botrychium matricariifolium A. Br. and B. simplex

E. Hitche.) in which a percentage of every population shows sporangia arising

from the sterile segment margins.

Sporophylls which have the encapsulated or angiosporangial condition are

especially well developed in Plagiogyriaceae and Aspleniaceae subf. Blechnoideae.

In these ferns the sporangia are covered, at least partially, by the inrolling of the

pinna margins. They are protected until they are fully mature and ready to

discharge the spores. To study early stages of sporangial development it is necessary

258 Gardens’ Bulletin, Singapore — XXX (1977)

to cut away or pull back the folded-over lamina. The most highly evolved

encapsulated structures are those of the onocleoid ferns, especially Matteuccia and

Onoclea, in which the sporophyll capsules do not open until months after the

sporangia have matured.

Fertile-sterile differentiation at the histological level has been largely uninvesti-

gated and we are currently trying to determine the scope of the anatomical changes

associated with sporophylls. By using the frequent intermediates that occur as

abnormalities in many dimorphic taxa, it is possible to detect gradients and

establish homologies. Techniques of clearing and staining portions of the leaves

are especially suited to this type of study. A brief summary of our observations

is as follows: Narrowing of the segments of sporophylls is accomplished by the

progressive reduction of the lamina until, in extreme cases like the botryoidal

types described above, the axes become terete and there is no trace of laminar wings.

Associated with this there is a large increase in the ratio of over-all surface to

volume, allowing for more rapid drying. With reduction in lamina we usually

notice an increase in the width of the vein intervals and a simplification of the vein

pattern. The latter is accomplished by eliminating some of the vein branchings

and most or all of the areoles, as in Lorinseria areolata (L.) Pres] (Blechnoideae)

and Onoclea sensibilis L. and Onocleopsis hintonii F, Ballard (Dryopteridoideae,

Labouriau, 1958; Lloyd, 1971). In Lomariopsis the sporophylls have free-ending

veins, lacking the marginal connections of the trophophylls (Holttum, 1939).

Comparative investigations of the vein intervals of nearly two dozen taxa were

made by Lommasson and Young (1971), but they did not compare conditions in

sterile and fertile fronds.

There may be simplification in other laminar tissues as well. The epidermis

may be softer in sporophylls than in trophophylls, and may tend to produce fewer

stomates. The epidermal cells may be larger and straighter-walled. The mesophyll

may have less elaborate spongy parenchyma that is composed of larger, thinner-

walled cells. In most strongly dimorphic taxa, the cells lose their chlorophyll at

maturity, appearing en masse a whitish, yellowish, or brownish color. Where the

acrostichoid condition occurs, the sporangial initials are no longer localized in

punctate or linear receptacles, but become spread over the abaxial surface, forming

a continuous placental area both along and between the veins. As indicated earlier,

the sporophyll venation of certain acrostichoid ferns may actually be more complex

than that in the trophophylls.

The physiological study of sporophyll-trophophyll differentiation is still in its

infancy. One thinks of such names as Labouriau, Sharma, Steeves, Sussex, Wardlaw,

and Wetmore in connection with processes of initiation and development of fertile

leaves. White (1971) has reviewed much of their work. There seems to be little

question that formation of sporophylls is controlled hormonally, and that their

timing is related to periods of light and perhaps heat, although there is a vast

amount of research needed to clarify the factors involved. Various extraneous

influences may interfere with the normal course of sporophyll development, causing

intermediate fronds to form. There are scattered reports on effects of injuries

brought on by such agents as fire, flood, mowing, destructive substances (e.g., tar),

and the trauma of transplanting, in inducing intermediate fronds, but there are

very few controlled experiments to warrant general conclusions, It does seem that

destruction of many or all of the actively photosynthetic fronds of a given plant

will have a tendency to produce imperfect expressions of sporophylls in the new

crop of fronds. In Polystichum acrostichoides, the partially fertile, subdimorphic

pinnae of “forma incisum (Gray) Gilbert’? apparently result from abnormal new

growth of late leaves after injury of the normal fronds of the season (Wagner,

Farrar, and McAlpin, 1970).

Leaf Dimorphy in Ferns 259

In very strongly marked dimorphy, the fertile frond becomes less responsive

to light and more responsive to gravity. The typical spreading orientation of the

trophophyll in which the blade approaches a position perpendicular to the direction

of light is absent in the sporophyll which is now so negatively geotropic that it

becomes totally erect. A simple and quick experiment to demonstrate tropism of

fertile parts may be made with Botrychium virginianum (L.) Sw. If whole leaves,

including both the fertile and sterile segments, are cut off at ground level during

the active growing season before the sporangia have fully matured or dicharged,

and laid in pans of water in various positions, we have found that at room tempera-

ture in only 10 to 20 hours all of the fertile spikes will twist and become upright,

no matter what the position of their associated sterile segments, illustrating dramati-

cally the “‘urge”’ to raise the sporangia to the highest position possible.

The tissues of the sporophylls, being simpler than those of trophophylls and

having a more flimsy structure, are more readily dried out. Their vertical position

in relation to sun and wind, in contrast to the trophophylls which are closer to the

substrate and more protected, also promotes dessication, Presumably the sporophylls

have “‘cheaper construction” partly because they are more ephemeral and, lacking

photosynthesis, have no value to the plant after their function of spore production

is finished. Support by the rest of the plant is withdrawn when the sporophyll’s job

is done, and the fertile frond accordingly turns brown, dries out, and falls away.

This is shown especially well in single hemidimorphic leaves. The differentiated

fertile tips of Osmunda regalis and Polystichum acrostichoides dry up and die long

before the sterile parts of the same leaf. In Botrychium, Ophioglossum, and Anemia,

the photosynthetic pinnae continue to function as if nothing had happened, even

though major parts of the leaf, the fertile segments, have collapsed and turned to

twisted debris.

The seasonal timing of sporophylls and trophophylls may differ profoundly

in taxa occurring in the same geographical areas and habitats. Nozu (1968) studied

periodicity in Japan of members of nine genera with respect to sterile-fertile frond

maturation, and reported that in a given species they may differ by as much as six

months, Usually the sterile fronds mature first during the course of the year, but

in Osmunda japonica Thunb. the fertile fronds mature a month or so earlier than

the sterile.

A most curious state of affairs is found in the periodicity of Botrychium

lunarioides (Michx.) Sw., a rarity of the southeastern United States known as

‘Winter Grapefern.” The frond arises above ground in November and the sterile

segment expands to its full size by sometime in January. The fertile segment, on

the contrary, remains curled over at the ground level and stays soft and meriste-

matic for nearly three months, only expanding and become erect in March. Spore

discharge occurs in late March and then the entire leaf, including both fertile and

sterile segments, dies in April. The plants are dormant, and completely buried

under ground, from April to November.

Of what adaptive value is fertile-sterile leaf dimorphism? It is found in plants

of such widely different habits and habitats that it is practically ubiquitous. The

condition is found in giant ferns (e.g., Acrostichum) and midget ferns (e.g.,

Peltapteris), in thick-textured ferns (e.g., Polystichum) and thin-textured ones (e.g..

Trichomanes). Dimorphy occurs in ferns with simple leaves (Pyrrosia) and with

compound leaves (Maxonia). It exists in taxa that are terrestrial, taxa that are

epiphytic and taxa that are aquatic. It is found in taxa with upright stems, with

creeping stems, and with climbing stems. Xerophytes and hydrophytes, arctic and

tropical ferns —— all have dimorphic representatives, the common species as well

as the rare.

260 Gardens’ Bulletin, Singapore — XXX (1977)

For what ecological reasons has dimorphic evolution occurred in such a

gamut of ferns? One can, of course, ask the same question regarding other

pteridophytes ——- Equisetopsida and Lycopodiopsida which have made

analogous changes of shoots rather than leaves. In all likelihood, similar forces have

operated to produce strobili (analogous to apically hemidimorphous fronds) and

whole fertile shoots (analogous to fertile-sterile segregation of whole fronds, for

example Equisetum arvense L. and Lycopodium carolinianum L.).

Some major extrinsic factors to take into account are seasonality, radiant

energy (e.g., heat, light), role of air and water, and biotic factors (e.g., shading,

grazers, fungi). The plant functions of most importance here are photosynthesis,

spore production, and spore dispersal and establishment.

All of the following adaptations may contribute to the differentiation of sterile and

fertile fronds:

1. To elevate spores of the fertile fronds into the wind stream for maximum

dispersal.

2. To permit more extreme drying effects to enable sporangia to open and

close efficiently. .

3. To spread out photosynthetic blades in horizontal positions for maximum

reception of light.

4. To place photosynthetic blades in most humid surroundings to counteract

water loss.

5. To bring photosynthetic blades closer to the substrate where there are

higher carbon dioxide concentrations during day time.

6. To enable differentiation of seasonal timing for most favourable period for

spore reproduction and most favourable period for vegetative assimilation.

7. To separate fertile from sterile parts if one or the other is more liable to

grazing or fungal attack.

Most authors agree that sporophylls tend to bring the spores into a position

where they can better be carried away by the wind. Holttum (1938) describes the

common situation which ‘takes the form of a somewhat contracted lamina in the

fertile fronds, which stand erect, on longer stipes than the broader sterile fronds

which bend away in a rosette around them, ... This form of dimorphism un-

doubtedly helps the distribution of spores, raising the sporangia well above the

surrounding leaves and exposing them to drier air and any slight wind that may

exist.””

Copeland (1906) emphasized the drying aspects in his analysis. In his words,

‘“‘Dimorphism, whether merely begun, or highly developed, whether a character of

whole fronds or of their parts, has in all cases the object of permitting the proper

dryness of the mature sporangia without an improper desiccation of the vegetative

structures, This is done by merely raising the reproductive structures farther above

the substratum; or (rarely) by special structural devices, such as notched margins;

by a restriction of the assimilating surface of the reproductive frond or region, so

that it may be sacrificed in emergency; or by a more complete elimination of the

vegetative structures in constitutionally ephemeral fertile fronds.”

_ As already stressed, origin of sporophylls is accompanied often by increasing

divergence or specialization of the trophophylls or trophophyllary blade parts. This

Specialization invokes progressive broadening of the blade together with re-

orientation into spreading or horizontal position. The blade becomes shorter and

if the fronds are clustered a rosette is formed (Fig. 2, D). In long-creeping

thizomes, the much reduced trophophylls are scattered and often cling to the

substrate. The modification of the trophophylls provides maximum photosynthetic

surface for the area of the blades and orients them so that they have a moister,

Leaf Dimorphy in Ferns 261

more protected environment. Under some circumstances, greater assimilation of

carbon dioxide may be possible due to diffusion from the substrate of the respira-

tory products of bacteria, fungi, animals, and roots.

We need to know more about optimal times of appearance of sporophylls and

trophophylls. Time of spore production, for example, may be highly adaptive, but

we will not know this until we understand more about life cycles in the natural

environment. In the Great Lakes area of North America, Hill and W. Wagner

(1974) determined that thin-walled green spores are discharged in the spring and

very thick-walled, non-green spores are discharged in the fall for the most part.

This may have to do with germination and establishment, as the former group of

spores are adapted for quick germination on mineral medium and the latter for

slow germination on organic medium. The best season for release of thin-walled

spores may be the spring when the greatest moisture and rainfall are available.

Comparable conditions prevail in the tropics with respect to wet and dry seasons.

Drought conditions in tropical dry seasons may call forth extreme adaptations

of the trophophylls, exaggerating those described above. as found in Anemia

Sect. Trochopteris (A. elegans (Gardn.) Presl, A. eximia Taubert) with extremely

small, flat laminae (cf. Mickel, 1962). The polypod, Microgramma heterophylla

(L.) Wherry: illustrates an elaborate form of heterophylly in a vining fern. This

tropical American fern grows over dry limestone rocks and produces short, clasping,

ovate leaves with blunt tips, in contrast to a more upright, elongate form of sterile

leaf, and the long, erect, narrowly lanceolate fertile leaves. The completely fiat.

rosette-like sterile segment of the winter grapefern, Botrychium lunarioides, is

probably adapted to growth only during winter months and is a device to capture

maximum heat and light as well as to protect the tissues from sudden drying or

cold shocks.

Whether or not sporophyll-trophophyll divergence has adaptive value with

respect to parasites is presently purely speculative, but such a possibility should

not be ignored. Collectors notice frequently, especially in the tropics, that soral

structures seem to be especially liable to molds and to tiny insects that feed

upon the spores and sporangia. Conceivably there are adaptive advantages to

isolating the parts most likely to be eaten from those which photosynthesize. There

is also a chance that the eating of sori and fertile pinnae by small animals may

play some role in dispersal, especially of those spores that germinate beneath the

soil or under leaf litter.

Although the onocleoid ferns are commonly cited by morphologists to

illustrate foliar dimorphy, they are, in fact, unusual and not typical in several

respects. Lloyd (1971) in his recent monograph of the systematics of the onocleoid

ferns compared the fertile and sterile fronds of the three genera involved, but he

did not compare them with other fern genera. Too little is known as yet about the

rare Mexican genus Onocleopsis in its natural setting to evaluate it, but the two

other genera, Onoclea and Matteuccia, show a number of unique features: The

fertile leaves are (1) shorter than the sterile or equal to them, (2) their tissues are

more dense and sclerified, (3) the sori, which are indusiate, are tightly encapsulated

at maturity, and (4) the sporophylls persist much longer than the sterile fronds,

which are early-deciduous in the autumn. What governs the final opening up of

the bead-like pinnules is unknown, but in the eastern United States and Canada,

where there is a strong winter-summer seasonality, the fertile units remain rigidly

closed during the non-growing season and spore discharge is prevented. The hard,

upright fertile leaves persist through the winter, covered with, or protruding

through, the snow. The spores within stay green during their winter dormancy, and

when they are released early in the spring they are ready to germinate immediately,

thus taking advantage of the vernal moistness (Hill and W. Wagner, 1974).

Although the sporophylls of Onoclea may sometimes equal or even surpass the

262 Gardens’ Bulletin, Singapore — XXX (1977)

trophophylls in height, those of Matteuccia are relatively much shorter, one-third or

less the length of the trophophylls. In August and September, the sporophylls of

Matteuccia are hard to see, buried as they are among the tall trophophylls and

the other foliage of their swamp habitats. But in mid-winter all or most of the

herbaceous plants have wilted and collapsed, so that the sporophylls are entirely

exposed and easily swept by the wind. However, the actual discharge does not

occur until March and April. Obviously the morphological peculiarities of these

plants are correlated with the unusual seasonality of their spore production and

release.

DISCUSSION

The noted Cornell University morphologist of the first half of this century,

Arthur J. Eames, entertained some unusual theories about foliar dimorphism, In

general he read the sequence just the opposite of the traditional interpretation and

the one adopted here. In Eames’ words (1938), ““The segregation of vegetative and

reproductive functions in different parts of the leaf, or in separate leaves, represents

undoubtedly a primitive condition ...; from this ancient dimorphism there has

developed the condition often called monomorphism -~—— found in the

majority of ferns, a blade serving for both vegetative and reproductive functions

... And from this there has developed a new dimorphic condition fertile and

sterile leaves separate; this change has taken place independently in various genera

...”’ He thus postulated “‘ancient” dimorphism, which is the primitive state, leading

to monomorphism, and then, once again to “‘modern’”’ dimorphism. (Cf. his

figure 175).

Eames cited three families Ophioglossaceae, Osmundaceae, and

Schizaeaceae as representing the primitive type. All of these are of the type

referred to in this paper as having “‘botryoidal” sporangial arrangement. Those of

‘“‘Polypodiaceae”” (ie., all higher leptosporangiate ferns) represent Eames’

‘“‘modern” type. We are not clear, however, upon what evidence he based his

conclusions. The Ophioglossaceae are so isolated and so lacking in fossil evidence

that we can only reason about what its course of evolution has been. We assume

that it was derived from monomorphic fern-like plants because of analogies with

repeated evolutionary sequences that we know in other groups. With respect to

Osmundaceae, Miller (1971) concludes that “‘isomorphism represents the primitive

state, incomplete dimorphism (only certain pinnae of a fertile frond modified for

sporangia formation) is intermediate, and complete dimorphism is advanced,” and

he calls attention to the fact that fragments of laminate fronds bearing osmunda-

ceous sporangia occur mainly in Jurassic sediments but range throughout the

Mesozoic. Miller notes that “‘such fronds are generally included in the form genus

Todites.”” Even though dimorphic pinnae in Osmundaceae were also found as early

as the Jurassic and Triassic, this does not mean that this condition was primitive,

any more than in families known primarily only today in which both monomorphic

and dimorphic pinnae occur.

Also, as regards Schizaeaceae, the investigations of Mickel (1962, 1967) tend

to contradict the conclusions of Eames. Mickel states that “‘although the change

to dimorphism seems spectacular, in Anemia it is apparently a relatively simple

and taxonomically minor one, which has occurred at least twice in the genus.” He

explains the unique leaf structure of typical anemias with their upright fertile basal

pinnae to be derived from the typical fern leaf with unmodified basal pinnae (as

in the closely related genus Mohria), and he describes a series of species, beginning

with Anemia colimensis Mickel, with nearly monomorphic pinnae, and continuing

through more and more differentiated forms, that finally terminates in such end-

points as A. rutifolia and A. millefolia.

In view of the patterns seen in Osmundaceae and Anemioideae, and the total

pattern of ferns in general, it is unlikely that there are two kinds of dimorphism,

Leaf Dimorphy in Ferns 263

‘primitive’ and ‘‘modern,” and that the latter arose from the former via a

monomorphic stage. Such an hypothesis appears overly complex in the face of the

over-all evidence. The hypothesis of one-way directionality is more economical and

more probable. And when one considers the numerous analogous trends as are

observed in other vascular plant orders such as Lycopodiales, Selaginellales,

Equisetales and Cycadales, for example, the idea of reversal from dimorphic to

monomorphic seems even less likely.

In his rather lengthy discussion of foliar dimorphy, Dickason (1946) wrote

that “‘Evidently dimorphism has not arisen in response to any given set of environ-

mental factors, and cannot be considered as adaptive.” He based his conclusion,

apparently, on the fact that dimorphy appears in plants of such widely different

habits and habitats. Dickason was also concerned with the fact that “dimorphic

and monomorphic species grow side by side.”” These conclusions are in keeping

with a general negativism on Dickason’s part with respect to adaptive values of

most phylogenetic trends in ferns. He did concede, however, that dimorphy is an

evolutionary advancement over monomorphy.

Whether Dickason’s conclusion that there is no adaptive value in dimorphy

is justified is questionable. Merely that foliar dimorphy occurs in a wide spectrum

of habits and habitats does not necessarily militate against its being adaptive with

respect to one or more of the factors outlined above. Furthermore, examples of

non-adaptive and adaptive character-states in taxa that grow side-by-side are

numerous in all plant communities. The point is that all plant species probably

have mixtures of both adaptive and non-adaptive or neutral attributes.

Taxonomically there is no question that dimorphism is an extremely helpful

character for making identifications. In Genera filicum, Copeland (1947) uses

fertile-sterile dimorphism as a key character for over fifty genera. Nevertheless, it

is not necessarily a valid character for establishing affinities or delimiting genera.

For one thing, dimorphy is not a single character but an ensemble of different

characters (Table 1), and any given expression invokes its own combination. Each

manifestation must be analyzed accordingly. Furthermore, dimorphy has arisen in

practically all families of ferns and in some of them probably several times. Table

2 summarizes the incidence in the various families and subfamilies of ferns. What

is designated as ““weak”’ involves only a few representatives of the taxon or only

slight subdimorphism or both. “Strong” involves a majority of members and

usually very distinctive sporophylls and trophophylls or well marked hemi-

dimorphism. At the family level, dimorphy seems to be most pervasive in

Ophioglossaceae, Plagiogyriaceae, Cheiropleuriaceae, Marsileaceae, and Salvinia-

ceae.

In spite of the fact that dimorphy has been used to distinguish a number of

fern genera, it is questionable whether it is of fundamental importance. Sometimes

the dimorphic taxon is connected by intermediates to monomorphic taxa. Tagawa

and Iwatsuki (Iwatsuki, 1961) erected the genus Dimorphopteris on the basis of its

perfectly dimorphic fronds, the fertile pinnae so contracted as to be linearimonili-

form. However, the type species, D. moniliformis from the island of Halmahera in

the Moluccas has been shown by Holttum (1972) to be an extreme in a series of

subdimorphic members of a section of the genus Pronephrium. Copeland, among

others, paid considerable attention to sporophyll-trophophyll distinction in his

classification, but this was apparently partially a result of his emphasis on con-

venience. In Table 3 are listed 12 genera (right-hand column) recognized by Cope-

land which may indeed be too little differentiated to justify separation from their

parent genera. 4

4]It may have some relevance that two recent students of the genus Equisetum, namely R. L.

Hauke and C. N. Page, have rejected shoot dimorphism as a determining character for

establishing sections and subgenera (Hauke, 1974).

264 Gardens’ Bulletin, Singapore — XXX (1977)

The major taxonomic value of foliar dimorphy in ferns as a whole is at the

species level. There are numerous examples of pairs of closely related species in

which one member is essentially monomorphic and the other dimorphic, Such

pairs deserve close examination; because of their genetic similarity their study

may give us valuable insights into the evolutionary and adaptive significance of

this phenomenon.

Table 1. Parallel and convergent changes associated with sporophyll-trophophyll differentia-

tion. The primitive state for each character is undifferentiated or nearly so.

LEAF SIZE

ORIENTATION

PETIOLE LENGTH

BLADE OUTLINE

BLADE CUTTING

LAMINAR WEBBING

LAMINAR MARGINS

VENATION COMPLEXITY

VEIN INTERVAL

MESOPHYLL

EPIDERMIS

SORIATION

ENCAPSULATION

SEASONALITY

DURATION

Strongly unequal

Sporophyll erect, trophophyll spreading

Unequal

Sporophyll narrow, trophophyll wide

Sporophyll usually more divided; trophophyll

usually less divided (but exceptions)

Sporophyll with reduction or loss of lamina;

trophophyll unchanged or with increase

Sporophyll commonly smoother; trophophyll

more toothed

Sporophyll most commonly simplified by vein

loss; trophophyll more complex

Sporophyll veins more remote; trophophyll

veins more approximate

Sporophyll non-green, soft; trophophyll green,

more compact and rigid

Sporophyll with fewer stomates and _ larger,

smoother-walled cells; trophophyll with

normal epidermis

Sori concentrated, often confluent into acrosti-

choid condition or grouped into botryoidal

condition

Sporangia, at least while young, covered by

rolled lamina

Sporophyll separated from trophophyll in time

of appearance, usually earlier but may be

later

Sporophyll short-lived; trophophyll long-lived.

Rarely otherwise

Leaf Dimorphy in Ferns 265

Table 2. Parallel and convergent evolution of foliar dimorphy in ferns. Relative incidence

and main types in different families or subfamilies. Letters refer to dimorphic types in

Figure 1.

STRONG TO

MODERATE COMPLETE

ABSENT TO WEAK

Polypodioideae (B, D) Ophioglossaceae (C)}

Marattiaceae (D)

Gleicheniaceae Lygodioideae (B) Osmundaceae (B, D, E)

Matoniaceae Dryopteridoideae (B,D, F) Plagiogyriaceae (D)

Dipteridaceae Blechnoideae (D) Anemioideae (C)?

Loxogrammeoideae Ceratopteridoideae (D)

Marsileaceae (C)4

Salviniaceae (D)®

Grammitoideae (B, D)

Schizaeoideae (D)?*

Adiantoideae (D)

Vittarioideae

Cyatheoideae

Hymenophylloideae

Lindsaeoideae

Asplenioideae

Azollaceae

1 Fertile spike composed of fused basal fertile pinnae.

2 Fertile basal pinnae not fused, except in some cases to the petiole.

3 Fertile blade drastically reduced and petiole exaggerated, Sterile petioles maintained at

maturity in Schizaea pusilla Pursh.

4 Type “C” if sporocarps are modified fertile pinnae.

5 Sporophyll is the immersed root-like leaf.

Table 3. Genera of close affinity which were discriminated by Copeland (1947) on the

basis of fertile-sterile leaf dimorphism.

MONOMORPHIC OR

SUBDIMORPHIC

PARENTAL GENUS

DIMORPHIC SEGREGATE

POLYPODIACEAE

Pleopeltis Lemmaphyllum

Pleopeltis Marginariopsis

Colysis Dendroglossa

Aglaomorpha Merinthosorus

Crypsinus Pycnoloma

ADIANTACEAE S. l.

Pteris Neurocallis

CYATHEACEAE S. 1.

Trichomanes Feea

ASPLENIACEAE S. 1,

Phanerophlebia Cyclodium

Ctenitis Psomiocarpa

Tectaria Tectaridium

Tectaria Hemigramma

Tectaria Quercifilix

266 Gardens’ Bulletin, Singapore — XXX (1977}

LITERATURE CITED

Bower, F. O. 1923. The Ferns. Vol. I. Analytical Examination of the Criteria of

Comparison. Cambridge U. Press.

Bower, F. O. 1928. The Ferns. Vol. III. The Leptosporangiate Ferns. Cambridge

WaPress,

Copeland, Edwin B. 1907. The comparative ecology of San Ramon Polypodiaceae.

Philippine Jour. Sci. 2 (no. 1): 1-76, pl. 1-4.

Copeland, Edwin B. 1947. Genera filicum. Chronica Botanica, Waltham, Mass.

Dickason, Frederick Garrett. 1946. A phylogenetic study of the ferns of Burma.

Ohio Jour. Sci. 156 (no, 2): 73-108.

Eames, Arthur J. 1936. Morphology of Vascular Plants. Lower Groups. McGraw-

Hill Co., New York.

Gupta, K. M. and T. N. Bhardwaja. 1956. Indian Marsileas: their morphology

and systematics. Jour. Bombay Nat. Hist. Soc. 53: 423-444, pl. 1-3.

Gupta, K. M. 1962. Marsilea. Botanical Monograph No. 2 Council of Sci. and Ind.

Res., New Delhi.

Hauke, Richard L. 1974. The taxonomy of Equisetum ——— an overview. New

Botanist 1: 89-95.

Hill, Royce C. and W. H. Wagner, Jr. 1974. Seasonality and spore type of the

pteridophytes of Michigan. Mich Bot. 13: 40-44.

Holttum, R. E. 1938. The ecology of tropical pteridophytes. Jn Verdoorn, F.,

Manual of Pteridology, pp. 420-450. Martinus Nijhoff, The Hague.

Holttum, R. E. 1939. The genus Lomariopsis in Madagascar and the Mascarene

Islands. Notul. Syst. 8: 48-62.

Holttum, R. E. 1972. Studies in the family Thelypteridaceae. The genus Proneph-

rium Pres]. Blumea 20: 105-126.

Iwatsuki, Kunio. 1961. Taxonomic studies of Pteridophyta VI. Acta Phytotax.

Geobot. 19: 1-11.

Labouriau, Luiz Gouvéa. 1958, Studies on the initiation of sporangia in ferns.

Arquivos do Mus. Nacional, Rio de Janeiro 46: 119-202.

Lommasson, Robert C. and C. H. Young, Jr. 1971. Vascularization of fern leaves.

Amer. Fern Jour. 61: 87-93.

Lloyd, Robert M. 1971. Systematics of the Onocleoid ferns. Univ. Calif. Publ. Bot.

61: 1-93. |

Mickel, John T. 1962. A monographic study of the fern genus Anemia, Subgenus

Coptophyllum. Iowa State Jour. Sci. 36: 349-482.

Mickel, John T. 1967. The phylogenetic position of Anemia colimensis. Amer. Jour.

Bot. 54: 432-437.

Nozu, Yoshitomo, 1968. Studies on the leaves of the Ferns. I. The dissimilarities

of the fertile and sterile fronds in some ferns. Jour. Fac. Sci., Univ. Tokyo

Sec. III. 10: 13-27, pl. 2.

Wagner, W. H. Jr. 1965. Paraphyses: Comments on recent papers. Taxon 14:

299-302. |

Leaf Dimorphy in Ferns 267

Wagner, W. H., Jr. 1973. Some future challenges of fern systematics and phylogeny.

In Jermy, A. C., Crabbe, J. A., and Thomas, B. A. (ed.), ““The Phylogeny and

Classification of the Ferns.’ Bot. Jour. Linnaean Soc. 67 (Suppl. No. 1):

245-256.

Wagner, W. H., Jr., D. R. Farrar, and B. W. McAlpin. 1970. Pteridology of the

Highlands Biological Station Area, Southern Appalachians. Jour. Elisha

Mitchell Sci. Soc. 86: 1-27.

White, Richard A. 1971. Experimental and developmental studies of the fern

sporophyte. Bot. Rev. 37: 509-540.

Wieffering, J. H. 1964. A preliminary revision of the Indo-Pacific species of

Ophioglossum (Ophioglossaceae). Blumea 12: 321-337.

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A Note on Dendrobium serpens (Hk.f.) Hk.f.

GUNNAR SEIDENFADEN

Botanical Museum, Copenhagen

In his “Orchids of Malaya’, Holttum (1964 : 319) lists Dendrobium serpens,

adding to his description that “‘more information is wanted about this species”.

We listed it in “Orchids of Thailand’? (Seidenfaden & Smitinand 1960: 244)

based on Holttum’s information that Dendrobium virescens Ridl. should be con-

sidered a synonym, and remarked that the plant was found once in Tomoh in

Peninsular Thailand just north of the Malayan border but we had never ourselves

seen the plant.

In the summer of 1970 Smitinand and I went on an orchid hunting expedition

to the southernmost provinces of Thailand. Among the plants collected near the

border of Malaya at Waeng Forest Station west of Sungai Kolok was a trivial

Dendrobium, which was brought to the hothouses in Copenhagen in the hope that

it would produce the flowers necessary for the identification. No flowers appeared

for years, but when in the spring of 1975 we experimentally raised the average

humidity in the houses, several of the plants showed increased activity, and, in

May 1975 it produced flowers, making it possible to identify it as Dendrobium

serpens by means of Holttum’s keys. This led to the small study related below,

giving some bits of the information wanted by Holttum.

During the period of printing of the orchid part of ““The Flora of British

India” Hooker f. continued to receive new material, both in the form of new

collections in the field and as drawings, mainly from the Calcutta herbarium. This

necessitated the addition of a supplement to his treatment, found in the sixth

volume of “Flora of British India’? containing a lot of important, although too

often overlooked, information.

Among the drawings received from Calcutta was one of a Dendrobium of the

Pedilonum section, said to represent a plant growing at ‘“‘Rangoon’’, brought by

Mr. Gilbert, a resident of Moulmein, where he succeeded Parish, and on a more

modest scale tried to take up the work of this great botanist, sending to Europe

herbarium specimens of orchids, often accompanied by sketches of which several

are found in Reichenbach’s herbarium in Vienna. In the present case Gilbert must

have sent a living plant, because when Hooker f. received the drawing from

Calcutta, to it was attached a single dried flower, said to have developed in the

Garden at Calcutta in November 1884. As to my knowledge the species has never

turned up in the Pegu area, I feel we must not exclude the possibility that the plant

was not at all found “at Rangoon” but rather, brought up to Rangoon from

Moulmein for shipment to Calcutta. Hooker f. called the plant D. panduriferum.

Another drawing received from Calcutta, made after a plant collected by

Kunstler in Perak and flowering in Calcutta in May 1884, prompted Hooker f.

to describe it as a variety of Gilbert’s plant, calling it Dendrobium panduriferum

var. serpens. Of this he evidently got no plant material and had in his description

to rely solely on the drawing. This too he later reproduced (1895:10, PI.16), at

which time he reached the opinion that the two entities were different enough to

justify considering the Perak plant a separate species, Dendrobium serpens.

Very little information on these two taxa has appeared since. Grant (1895: 108)

copied Hooker f. in his Burma list, while Ridley in his paper on the orchids of

269

270 Gardens’ Bulletin, Singapore — XXX (1977)

GT 7517

RIDLEY 14303

MACHADO

Fig. 1. Dendrobium serpens (Hk. f.) Hk. f. a. inflorescence from a fresh Thai specimen,

b. flower from the Temengo specimen, c. flower from the type specimen of D. virescens, d.

epichile and e. flower without lip and mentum from the Thai specimen.

Dendrobium ser pens 27]

the Malay Peninsula (1896) overlooked Kunstler’s Perak plant which he sub-

sequently listed in his “‘Materials” (1907:47), and, in his “‘Flora of the Malay

Peninsula” (1924:47) we learn that he had identified one of his own collections

as being Kunstler’s plant, giving the locality as Ulu Temengo (= Temengor) on

an island in the Kertai River.

Already in 1896 Ridley had, however, introduced a new taxon, Dendrobium

virescens, based on a collection made by Machado further to the Northwest, at

Tomo (spelled by Ridley as Tomoh) which I believe is on the Thai side of the border

and, in the “‘Materials’’ we learn that he had identified a plant collected at Taiping

in Perak by Curtis, — the latter is repeated in the 1924-paper.

In his large monograph on Dendrobium, Kranzlin (1910) lists Dendrobium

virescens and D. panduriferum without adding any new information, he places

them in two different subsections of the Pedilonum-section, although they might

be considered conspecific, cfr. below. Dendrobium serpens he choses to consider

a synonym of Reichenbach’s Dendrobium ionopus, a plant said by Reichenbach to

have come from Burma. Hooker f. had originally suggested that his var. serpens

could be the same as D. ionopus, but later (1895:11) he indicated that he was

convinced of the contrary; Kranzlin seems to disregard this viewpoint, while both

Ridley and Holttum ignore Kranzlin’s opinion.

In response to “more information” sought by Holttum, I got with the kind

assistance of Dr. Chang the available material from Singapore, at the same time

receiving from Vienna the material kept in the Reichenbach herbarium under the

name of Dendrobium ionopus.

There seems no doubt that the recently found Thai plant is identical with both

the Tomo plant which is the type specimen of Dendrobium virescens Ridl. and

with the plants from Ulu Temengo and Taiping. The most characteristic features

are details of the lip, see Fig. 1. There is a half-moon shaped transverse callus at

the base of the epichile at the side of which the edges fold into ears continuing

as longitudinal somewhat converging folds on the blade of the epichile. There is

no fattening on the epichile, the whole of it is of very thin texture, which is not very

clear on the drawings Hooker f. received from Calcutta; based only on these,

wane ails the blade as “‘cordate above and below, slightly constricted at

the middle”’.

When comparing the original drawing which was the only basis for Hooker’s

analysis, with the plate published by Hooker of Dendrobium panduriferum it is

important to recall that here the details of the flower are based on Hooker’s own

dissection of a flower he received with the habit drawing. It is tempting to suggest

that after moistening or cooking the flower, any folds and ears in the thin material

would vanish, and a lip like the one found in the material we have from Thailand

and Malaya of D. serpens would look pretty much like Hooker’s drawing of the

lip of Dendrobium panduriferum, described by Hooker as “‘deeply constricted with

auricle-like sidelobes”, the transversal callus is the same. I am therefore quite

convinced that D. panduriferum and D. serpens are the same species. The only

depiction of a flower in front view in the habit drawing of D. panduriferum (drawn

in India after fresh material) shows, in my opinion, an epichile indistinguishable

from the one shown in the Indian picture of D. serpens, and the other differences

mentioned by Hooker seem not very convincing. I feel, however, that it would be

too hasty to reduce D. serpens to a synonym, perhaps it is safer to wait until fresh

material has been obtained of plants from ‘‘Rangoon’’; repeating Holttum’s words

that “‘more information is wanted’’, my preliminary summary on D. serpens

therefore reads as follows:

Dendrobium serpens (Hk. f.) Hk. f., Ann. Bot. Gard. Calc. 5: 10, T. 16, 1895.—

ee 1907:51; idem 1924:47; Holttum 1964: 319; Seidenfaden & Smitinand

1960: 244.

272 Gardens’ Bulletin, Singapore — XXX (1977)

Dendrobium panduriferum var, serpens Hk. f., Fl. Brit. Ind. 6: 186, 1890.

Dendrobium virescens Ridl., Journ. Linn. Soc. 32:259, 1896—F. N. Williams

1904: 368; Ridley 1907:51; Kranzlin 1910:100; Ridley 1924:47.

Dendrobium ionopus auct. non Rchb, f.: Kranzlin 1910: 122 (p.p.)

Distribution: Thailand: Legeh at Tomoh (Macado s.n. SING!, type of Dendrobium

virescens); Near Waeng Forest Station, Narathiwat 250m (GT 7517 Cl).

Malaya: Perak (drawing in Calcutta after a plant sent by Kunstler, type); Ulu

Temengo on an island in Kertai River (Ridley 13303 SING!); Taiping (Curtis

s.n. SING)!); presumably also Karangan, Kedah (Haniff s.n. SING!, insufficient

material).

Now, on Dendrobium ionopus Rchb, f. of which Kranzlin considered Dendro-

bium serpens a synonym:

The material of this taxon in Herb. Reichenbach consists of two sheets. The

one carrying the handwritten original description (No. 41205) contains a single

dissected flower which, under the number 3769, had been sent by Low in November

1882 and said to originate in Burma, further, some rough coloured sketches by

Reichenbach of that flower, and a copy of one of Day’s drawings. I later saw the

original of that drawing in Day’s rich manuscript collection in Kew, made in

November 1882, and a note gives the same information that it was imported by

Mr. Low from Burma. The other sheet (No. 41204) contains an inflorescence also

sent by Low, with whom it flowered in 1881; it is noted that the collector was

Boxall and that it originated in the Philippines.

The flowers on the two sheets are identical. The most important characteristics

are the two low triangular lateral calli near the edge at the base of the epichile

(these are both in Reichenbach’s and Day’s drawings coloured dark red-purple),

and the obtuse tooth in the “‘spur’’, see Fig. 2. The flower is quite different from

that of D. serpens, and Kranzlin was wrong.

But seeing my sketches of D. ionopus, Dr. Garay called my attention to the

figure of a taxon described in 1908 by Kranzlin as a new species, Dendrobium

epidendropsis, based on one of Loher’s plants. This is clearly the same as Low’s

plants. D. epidendropsis has later been recorded several times from Luzon.

Here again, it is at hand to be suspicious about the old information. Con-

sidering the general tendency during the later part of the last Century to be less

exact (and often with purpose misleading) on the origin of commercially introduced

plants, I feel convinced that the flower Low sent to Reichenbach and Day in 1882

as “from Burma” is from the same plant or the same consignment as the one sent

a year before as being a Boxall-collection from the Philippines, and I feel we

should exclude the record from Burma.

Finally, I would suggest that the photograph reproduced under the name

Dendrobium O’Brienianum by Davis & Steiner (1952: 120) looks more like

Dendrobium ionopus; Kranzlin’s drawing of D. O’Brienianum (1910: Fig. 5 N-O)

has a much more slender mentum, also the photograph shows a pair of dark spots

somewhere at the base of the lip, not mentioned in Kranzlin’s colour description,

but, naturally, the photo is not sharp enough for a sure identification.

pee My knowledge on Dendrobium ionopus could therefore be summarized as

ollows:

Dendrobium ionopus Rchb. f., Gard. Chron. 1882, 2: 808.—

Hooker f. 1890: 732, Grant 1895: 100; Kranzlin 1910: 122. (excl. syn. D. serpens).

Dendrobium epidendropsis Krzl., Orchis 2: 79, Fig. 12, 1908, syn. nov. — Ames

1909: 598; Kranzlin 1910: 106; Ames 1915: 123; idem 1925: 348.

Dendrobium ser pens 273

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RR tt Ch

5mm

H.R. 41205

Fig. 2. Dendrobium ionopus Rchb. f. Type specimen. a. the only mutilated flower, with

dorsal sepal, a petal and part of a lateral sepal missing, lip removed, b. epichile of lip, c.

tooth in mentum.

Dendrobium O’Brienianum auct. non Kranzlin: Davis & Steiner 1952: 120?

Distribution:? Burma: s.n. (Low 3769 Herb, Reichenbach 41205!

improbable).

, type, origin

Philippines: sine loc. (Boxall s.n. Herb. Reichenbach 41204!; Loher s.n., type of

D. epidendropsis; Lyon 118 and 161, fide Ames); Lamao River, Bataan, Luzon

(Borden 2109 and Elmer 6839, fide Ames); Rizal, Luzon (Ramos 3060, fide

Ames).

Literature Cited

Ames, O., 1909: Notes on Philippine orchids with descriptions of new species 1.

Phil. Journ. Sci. 4, 5: 593-600.

. 1915: Orchidaceae 5. The Genera and Species of Philippine Orchids.

Boston, 271 pp.

. 1925: Enumeration of Philippine Apostasiaceae and Orchidaceae. Bur.

Sci. Publ. No. 18: 252-463. Manila.

Davis, R. S. & M. L. Steiner, 1952: Philippine Orchids, New York, 270 pp.

274 Gardens’ Bulletin, Singapore — XXX (1977)

Grant, B., 1895: The Orchids of Burma. Rangoon, 416 pp.

Holttum, R. E. 1964: Orchids of Malaya. Flora of Malaya 1. 3rd. ed. Singapore.

Hooker, J. D., 1890: The Flora of British India 5: 667-864, 6: 1-198. Ashford,

Kent.

. 1895: A Century of Indian Orchids. — Ann, Roy Bot. Gard. Calc. 5:

1-68, Pls.

Krinzlin, F., 1910: Orchidaceae-Monandrae-Dendrobiinae, in Engler: Das

Pflanzenreich IV. 50. II. B. 21.

Ridley, H. N., 1896: The Orchideae and Apostasiaceae of the Malay Peninsula.

Journ, Linn, Soc. Bot. 32: 213-416.

. 1907: Materials for a Flora of the Malayan Peninsula 1: 7-233.

Singapore.

. 1924: The Flora of the Malay Peninsula 4: 4-233. Ashford Kent.

Seidenfaden, G. & T, Smitinand: The Orchids of Thailand 2, 2: 185-326 Bangkok.

Williams, F, N., 1904: Liste des Plantes connues du Siam. Bull. Herb. Boiss.

2. ser. 4: 362-372.

Notes on the Systematy of Malayan Phanerogams XXV.*

Araliaceae

BENJAMIN C. STONE

Department of Botany, University of Malaya,

Kuala Lumpur

Abstract

Two genera are reduced to synonymy, Wardenia King to Brassaiopsis Decne. & Planch..

and Acanthophora Merr. to Aralia L. Critical notes on some other genera, some new

combinations, and two new species, Brassaiopsis minor and Schefflera singularis, are published.

A brief growth analysis of Arthrophyllum diversifolium sensu King is presented.

In the course of preparing the treatment of Araliaceae for Vol. 3 of the “Tree

Flora of Malaya” a number of problems involving not only the recognition and

delimitation of species, but also of genera, have arisen. Two undescribed species

have been encountered and are published here. Equally important, the status of

two hitherto monotypic genera, Wardenia King and Acanthophora Merr., has been

reassessed, and it has been concluded that they should be reduced to synonymy

under Brassaiopsis Decne. & Planch. and Aralia L. respectively. The largest genus

of the family in Malaya, Schefflera J. R. & G. Forst., is currently being revised

by D. G. Frodin of the University of Papua New Guinea. In his dissertation

(Cambridge University 1971) he has taken a broad view of Schefflera, incorporating

within it three genera recognized by Ridley in his Flora of the Malay Peninsula

(vol. 1, 1922), Brassaia Endl., Scheffleropsis Ridl., and Tupidanthus Hk. f,. &

Thoms. In a recent publication (1970) Philipson has reviewed the genus Gastonia

and remarked on a species which occurs in Malaya, which was not in Ridley’s

Flora. The genus Acanthopanax Miq. has been found to occur in Malaya, the sole

Malayan species being A. malayana Henderson (Gdns Bull. Sing. 7: 105-6, pl.

23. 1933). Ridley himself described a second species of Hederopsis King some years

after his Flora was published (H. major Ridl. in Kew Bull. 124. 1929). The genus

Polyscias Forst. is not in Ridley’s Flora, but is represented in the Peninsula by

several species found in cultivation.

Thus at present there are known to occur in Malaya ten genera of Araliaceae:

Acanthopanax, Aralia, Arthrophyllum, Brassaiopsis, Dendropanax, Gastonia,

Hederopsis, Polyscias, Schefflera, and Trevesia. If Tupidanthus is maintained as a

distinct genus, as advocated by Philipson (pers. comm.), the total is eleven.

The genus Aralidium Mig. was assigned to Cornaceae in Ridley’s Flora, but

placed in Araliaceae by Corner (Wayside Trees of Malaya 1: 154. 1940 & 1952).

After considerable preliminary study it is deemed preferable to exclude it from

Araliaceae on account of its lack of resin canals, among other characters. Whether

it should rejoin Cornaceae or find some other position is a problem still under

study.

275

276 Gardens’ Bulletin, Singapore — X XX (1977)

Aralia L.

Ridley reported three species of Aralia from the Malay Peninsula, A. thom-

sonii Seem., A. armata (Wall. ex G. Don) Seem., and A. ferox Miq. Merrill

(Philipp. J. Sci. 13, Cr, 316-318, 1918) showed that Philippine and Celebes

specimens determined as A. ferox Mig. were not that species at all, and because

of the climbing habit created a new genus Acanthophora for these plants with A.

scandens Merr. as the sole species. Later, van Steenis (Bull. J. Bot. Buitenz. ser. 3,

17: 390, 1948) added further determinations and included the “‘A. ferox” of

Ridley’s flora, adding the genus Acanthophora to the known Malayan Araliaceae.

The species Acanthophora scandens Mert. is certainly distinct. It has recently

again been collected along the Ginting Highlands Road, Pahang (Stone 12016).

A study of this material and a review of the pertinent literature suggest however

that the generic status for this species is highly dubious, its sole claim to it being

the lianoid habit. Since most species of Aralia are shrubs or trees (or herbs) the

liane habit does stand out. On the other hand there are many genera known which

include species with this habit along-side others which are trees or shrubs, witness

Schefflera, and outside the Araliaceae, Derris (Leguminosae), to name but two

examples. In this view I am supported by D. G. Frodin (Misc, Rec. Fl. Mal.

Found. III: 8. 1973).

In any case there is a nomenclatural reason why the name Acanthophora

Merr. 1918 cannot be used, even if the generic concept it denotes were considered

worth retaining. This is the generic name Acanthophora Lamouroux 1813, applied

to a taxon of Rhodophyceae (Red Algae). This name was obviously overlooked

by Merrill, when he created what was, in fact, a later homonym.

Aralia scandens (Merr.) T. D. Ha, Novost. Sist, Vysh. Rast. 11: 229, 1974.

Acanthophora scandens Merr. Philip. J. Sci. C, 13: 316-318. 1918.

Aralia ferox sensu Ridley (non Miquel) in Fl, Mal. Pen. 1: 873. 1922.

(Not Acanthophora Lamouroux, 1813 (Algae) ).

Distrib. — Malaya, Philippines, Celebes. In Malaya: Perak, Pahang, and

Selangor. Fig. 1.

Arthrophyllum Blume

Among the Araliaceae this is one of the more clearly distinct genera, rather

easily recognized by its pinnate leaves and 1-celled ovary and fruit. The species

are another matter. They are poorly described, mainly because of poor inadequate

collections, and badly discriminated. Most are rather difficult to distinguish. Those

in the Peninsula in particular are in a state of confusion because of the too-

Optimistic classification by Ridley. In the Flora (I: 885) Ridley lists 8 species.

These cannot be reached through the key and cannot be recognized from the

descriptions which are too brief and contain errors of fact. The available specimens,

including types and syntypes, even when examined, are not always decisive. Of

the 8 species listed by Ridley, four seem to me reasonably clear; the others are

to be considered as synonyms or as varieties of these four.

The main point of this note is to describe in some detail the basic growth

pattern of the largest of the Malayan Arthrophyllums. This plant was called A.

diversifolium by Clarke, King and by Corner, while Ridley called it A. ovalifolium.

Since it never possesses bipinnate leaves, it does not correspond very well with the

former. Apparently it is not the same as the latter. The nomenclature will be treated

In a separate paper by W. R..Philipson. At any rate, it is clear that throughout

the lowlands of Malaya, there is a single, rather uniform tall Arthrophyllum species, .

277

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278 Gardens’ Bulletin, Singapore — XXX (1977)

which often enters waste ground as a pioneer regrowth plant. Its trunk eventually

branches sparsely, and the trees may reach a height of over 15 m. This is the plant

referred to here in the following description.

Arthrophyllum sp. — A. diversifolium sensu King, Mat. Fl. Mal. Pen. in J. R.

As. Soc. Bengal, 47, II (1): 59. 1898, non Blume; sensu Corner, Wayside Trees

of Malaya 1: 155, 1952. — A. ovalifolium sensu Ridley, Fl. Mal. Pen. 1 : 885,

1922, non Jungh. & De Vriese. Fig. 2.

A. congestum Ridl. J. Fed. Mal. St. Mus. 10: 137. 1920; Flora 1: 887. 1922.

Lectotype: SELANGOR: Klang Gates, Ridley 13421. (SING).

Although Ridley’s binomial A. congestum applies unquestionably to this plant

there seems little doubt that it is to fall into synonymy when a wider study of the

group is made, and this is to be reported on by Professor Philipson in this volume.

Ridley attributed the name A. ovalifolium to Miquel, but the authors, as

stated by Miquel, were Junghuhn and De Vriese. Miquel’s description does not

seem to be a different interpretation. In describing A. congestum Ridley notes the

flowers as with 4 petals, but examination of the type collection shows that this is

wrong; the flowers, as would normally be the case, are 5-merous. There are no

other characters to distinguish this from the common, arboreous lowland species

which Ridley calls A. ovalifolium. In accepting this name for a Malayan species,

the chance of error is not small, and it remains to be checked against authentic

specimens, At any rate, the species intended here is the most abundant of the

Malayan lowlands, at least on the west coast, and is usually, when mature, a

branched tree up to 14 m tall with a trunk reaching 30 cm diameter. The leaves

on the main erect (orthotropic) stems are closely spiralled; they are large, once-

pinnate, reaching a length of 2 meters, with 12-14 pairs of large leaflets (Fig. 2-2).

Like Ridley, I have never seen a bipinnate leaf in any Arthrophyllum; on this

basis I have refrained from identifying our plants with A. diversifolium Bl. or its

synonyms, though this is the name used by King in the ‘Materials’ and Corner in

“Wayside trees of Malaya (1940)” and found on a number of herbarium specimens

in SING and KEP. According to the available descriptions A. diversifolium has at

least some bipinnate leaves. It is possible that hasty observation of our plants would

suggest that they too bear bipinnate leaves, but this arises through confusion of the

axillary determinate branches (which bear the flowering shoots) with leaves. These

axillary branches (Fig. 2-3) are more or less plagiotropic (though not usually

horizontal), and bear at their widely spaced swollen nodes opposite bipinnate but

reduced leaves, or nearer the tip, whorled and even further reduced leaves; in the

axils of the latter arise the short flowering shoots (Fig. 2-3 and 2-9), which

themselves have 1-3 nodes with pairs of simple leaves (Fig. 2-6). The resemblance

of the lateral branch to the rachis of one of the big leaves is striking, but its

position and smaller number of nodes serve to identify it. Since the leaves borne

on the lateral branches are much smaller, and with successively reduced numbers

of nodes, they are easy to collect, and hence are found on most herbarium sheets;

but they are not really representative of the plants, as besides being smaller, they

are not or scarcely asymmetric, which is the usual condition of the leaflets on the

lower, big leaves. Also, their number of lateral nerves may be smaller. These

Caeamaattes make identification of such fragmentary collections difficult and

esitant.

Plants of this species generally remain unbranched until they are about

2.5 m tall; at this stage they may branch, but the branches are recapitulations of

the trunk, i.e. orthotropic with spiral phyllotaxy and more or less equivalent. These

may grow to an equal length, then flower. Flowering is by inception of the axillary

lateral branches with distichous or whorled phyllotaxy, as described above; but

new orthotropic shoots arise from other axils providing relays. The lateral shoots

are strongly determinate in growth and reach about the same length as the big

Araliaceae 279

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Fig. 2. Arthrophyllum sp. (A. diversifolium, sensu King). Aspects of its morphology. | :—

First-order shoot. 2:—Pinnate leaf of first-order shoot. 3:—Second-order shoot.

4:—Medial leaflet from leaf of first-order shoot. 5:—Medial leaflet from lowest

leaf of second-order shoot. 6:—Simple leaf from third-order (flowering) shoot.

7:—Cross-section of leaf rachis. 8:—Axillary bud. 9:—Apex of first-order shoot

bearing second-order shoot branches (PB) and their axillant leaves (LF) or their

scars (LS). 10:—Floral details. Stone 12156.

280 Gardens’ Bulletin, Singapore — XXX (1977)

pinnate leaves of vegetative stems. Their resemblance to rachises has been mentioned

above; in fact, the immediate appearance suggests that the flowering umbels arise

in the axils of leaflets, but close study dispels this impression. The big lower

leaves always have at least 10 nodes, but the upper leaves on the lateral branches

may have as few as 1 or 2 nodes (these leaves usually immediately subtend the

flowering shoots). The reversion to spiral arrangement at the apex of the lateral

branch is abrupt and striking. On the flowering shoots, there are umbels in the

axils of the leaves on the median nodes as well as clustered at the apex. The

inflorescence, therefore, consists, strictly speaking, of a pseudumbellate cluster of

simple umbels, although superficially the appearance suggests a 2- or even 3-com-

pound umbellate structure.

The leaflets of a single leaf often have unequal laminal areas. The whole row

of leaflets on one side (e.g. the “‘left’’ or “‘right’’ side) may be slightly larger than

those on the other side of the rachis. However, if there is this ‘overdevelopment’

of the right side of a leaf, then the next higher leaf of the spiral often has its left

side similarly overdeveloped, and the successive higher leaves reveal an alternation

continuing this pattern.

The nodes of the leaf rachis are closer nearer the leaf apex, and the internode

distance is correlated approximately with leaflet width, being about the same or

slightly more than the width of the nearest leaflets. The lower internodes elongate

first and most rapidly.

The stipules are short and connate into a narrow ligular rim, ciliate in the

very youngest stage but soon glabrate. Internodes on the orthotropic shoots are

so short that the sheathing bases of the petioles are in contact.

The fruits are about 5-6 mm long, globose to broadly ellipsoid, capped by

the remnant calyx rim and the short persistent conic style.

Seedlings. Seedlings of this species are epigeal, with a pair of opposite bluntly

ovate cotyledons about 1 cm long. The first several true leaves are simple, ovate-

acuminate; these are eventually followed by pinnate leaves.

Brassaiopsis Decne. & Planch.

(Wardenia King, syn. nov.)

The genus Wardenia with its single species W. simplex King has been found so

far only in a few localities in Perak and Selangor and hitherto has been considered

monotypic. F. R. I. Kepong Herbarium has several specimens of Wardenia simplex

King, the type species of the genus cited below. These collections supplement the

original from Ulu Kerling, Perak, made by Kunstler over 80 years ago.

Two collections made in 1955 and 1969 in Trengganu, appear to represent a

second species of this genus. It is described as a ‘fleshy treelet’ and differs from

W. simplex in several obvious features, its elliptic acuminate cuneate (rather than

ovate subcordate) leaves, much shorter and more slender petioles, and diminished

inflorescence with little if any branching, and perhaps its longer pedicels and

sparser puberulence. The material is rather incomplete; no fruits are available. The

flowers are evidently past anthesis and the ovaries starting to enlarge. They clearly

show a bilocular ovary, each cell with a single ovule. This conflicts with King’s

Original generic description of Wardenia which calls for a unilocular ovary that

develops a dissepiment later, separating the seeds, but agrees with the diagnosis

of Brassaiopsis. According to Philipson (Bot. J. Linn, Soc, 63, Suppl. 1: 90, figs.

1-6. 1970) the original observations may have been wrong as the type collection

definitely has 2-locular young fruits. The floral character of “‘ovary 1-celled.

2-ovulate”” should be treated with caution, however, as in any case it ‘disappears’

in the fruit phase which is more normal for Araliads, i.e. there is one seed per

loculus. The ovular character may not differ therefore from the situation in

Araliaceae 281

Brassaiopsis Decne. & Planch. (including Fuaraliopsis Hutchinson. which is

Araliopsis Kurz non Engler). It would appear that Wardenia differs from

Brassaiopsis only in having simple leaves. However, simple-leaved species of

Brassaiopsis have already been described. Philipson (1951)* for example describes

B. castaneifolia from Burma, which has simple leaves. Thus the generic distinction

does not appear to be tenable. The venation pattern in the simple-leaved species

is quite easily reconciled with that of the compound or palmately lobed-leaved

species.

In both B. (Wardenia) simplex and B. minor, and in B. polyacantha and B.

elegans, the young parts are covered with a rufous furfuraceous indument.

Microscopic examination reveals that this is made up of numerous slender

brachiate to stellate hairs with a variable number of branches and considerable

variation in size. These structures are more or less ephemeral and the stems,

leaves, etc., become glabrescent with age. These hairs are virtually identical in all

four species, though their abundance varies. They are densest on the innovations

and buds of B. polyacantha.

The sheathing base of the petiole and the adnate stipules are similar in all

four species. The petiole base is obliquely invaginated and the basal margins

abruptly produced into a pair of lobes at a certain distance away from the base,

as in shown in the accompanying figures. This pattern also shows up in the lowest

inflorescence bracts, which appear to be much modified leaves with the blade and

petiole reduced effectively to the sheath and ligule, although a prolongation

representing the blade (and showing lateral nerves) but tightly cylindrical-subulate,

may be present.

The venation patterns in the leaves of all four species is comparable; the

lowest lateral nerves are always inserted on the midrib by means of an outcurved

junction. They are elevated beneath. There are comparatively few, i.e. widely

spaced secondaries often with subparallel intermediate nerves between them. The

lowest pair of lateral nerves tends to run up at or very near the blade margin and

justifies the term ‘subtriplinerved’. In the palmate-leaved species these nerves give

off leaflets which are thus asymmetrical. The overall veining pattern is campto-

dromous.

Consideration of these further characters, indument, petiole-ligule-stipules, and

veining, adds weight to the decision to reduce Wardenia to Brassaiopsis.

In R. Viguier’s anatomical study of Araliaceae (1906) Brassaiopsis was

included (in the ‘Shefflerineés’ p. 96) and a characterization of its petiole anatomy

was presented (the species not specified). For Wardenia, Viguier apparently had

no material and he was unable to compare it anatomically with other genera. (He

expressed doubt that it pertained to Araliaceae and suggested that it might belong

to the Cornaceae). However, such doubt is unnecessary and it is preferable to

reduce Wardenia to the status of a synonym of Brassaiopsis. This is also the view

of D. G. Frodin (Misc. Rec. Fl. Mal. Found. III: 8, 1973).

Key To MALAYAN SPECIES OF BRASSAIOPSIS

1. Leaves palmately lobed or divided

2. Leaves palmately 5—9-lobed, large to very large. to 60 cm wide; small

EE ye tn SRE ec i eine fie aie ie ea ats B. polyacantha

2. Leaves digitate with 3 or 5 or up to 3 leaflets, medium, to 25 cm. wide.

3. Leaflets 3-5: inflorescence branches with terminal umbel and a

ES EO OS Ae Or Ee ee oe B. elegans

3. Leaflets 5—9: inflorescence branches racemose-umbellate ............

Se eeebirerrsis . givissd. . aatcawadadd. Wasik -% Reds Ous 3s. B. glomerulata

* Phillipson, W. R. 1951. Contrib. to our knowl. of Old World Araliaceae. Bull. Brit.

Mus. (N.H.) Bot. 1 (1): 1-20.

282 Gardens’ Bulletin, Singapore — X XX (1977)

1. Leaves simple.

4. Leaves ovate-subcordate, to 35 cm long or more; petioles

stout elongate, 15 cm long, 5 mm thick. ............ B. simplex

4. Leaves elliptic, cuneate at base, smaller, to 25 cm long; petiole

to 7 cm Jong. 2 mm COCK far, tt acotk ans cae B. minor

ENUMERATION OF MALAYAN SPECIES OF BRASSAIOPSIS

|. Brassaiopsis polyacantha (Wall.) Banerjee, Indian Forester 93: 341. 1967.

B, palmata Kurz, J. As. Soc. Bengal 39, 2: 77. 1870. King, Mat. Fl. Mal.

Pen. in J. As. Soc. Bengal 67: 61. 1898; Ridley, Fl. Mal. Pen. 1: 887.

1922.

Malayan collections: KEDAH: Gunong Inas F. R., Sg. Kepang, alt. 3000 ft.,

tree 40 ft. tall, 3 ft. girth, fruit faintly bluish with white spots, in terminal raceme

2ft. long, 7.2.1968, Whitmore F RI-4639 (KEP); FRI-4644 (KEP). — Ulu Pantai

Mulik, 22.3.1938, Sow KEP-34612 (KEP). PAHANG: Cameron Highlands, Gunong

Brinchang, alt. 6000 ft., 4.5.1964, M.E.D. Poore KLU-8126 (KLU) — Telom Valley,

21.8.1931, Jaamat KEP-25199 (KEP). — Bukit Tinggi village, Bentong Rd., tree

30 ft., 8.2.1966, Kochummen KEP-97777 (KEP). — Bentong Rd. between Genting

Sempah and Bukit Tinggi, alt. 1500 ft., 21.12.1968, Stone 8319 (KLU). — Bentong

Rd. just beyond Genting Sempah, 1800 ft. alt., 2.2.1968, Whitmore FRI-4623

(KEP). PERAK: Maxwell Hill Road, 54 miles, 29.10.1969, Kochummen F RI-2860

(KEP),

Poore 8126 differs in having more coriaceous leaves, appressed stellate hairs,

thornier petioles. It may be a different species.

2. Brassaiopsis glomerulata (BI.) Regel, Gartenfl. 12: 275. 1863.

B. speciosa Decne, & Planch. Rev. Hort. 4, 3: 106. 1854. Hedera glome-

rulata Blume, Bijdr. 1: 872. 1826. — Aralia glomerulata (Bl.) DC. Prodr.

4: 265. 1830.

Distribution: From India and S. W. China to Malaysia, Sumatra, and Java.

MALAYA: PAHANG, J. H. Burkill & R. E. Holttum SF-7882 (SING); s.n.

in 1922 (SING). NeGri SEMBILAN, Md, Nur SF-11729 (SING).

This species may be recognized by its stature, for it becomes a tree up to 10 m

tall; by its distinct (petiolulate) leaflets; and by the long inflorescence branches

bearing umbellules of about 25-35 flowers.

3. B. simplex (King) B. C. Stone, comb. noy. Wardenia simplex King, Mat.

Fl. Mal. Pen. in J. As Soc. Bengal 67: 60. 1898. Ridley. Fl. Mal. Pen.

1: 887. 1922. — Fig. 3.

MALAYA: Perak, Slim Hills F. R., ridge top, 2500 ft. alt., unbranched

small tree 15 ft. tall, trunk with distant thorns, inflorescence terminal, 4 Sept. 1966,

Whitmore FRI-794 (KEP). SELANGOR, Compt. 12, Chadangan F. R., Ulu

Selangor, hill forest, 1200 ft. alt., tree 10 ft. tall, girth 4 in., 27.9.1966, Chelliah

KEP 98147 (KEP).

4. B. elegans Rid]. Fl. Mal. Pen. 1: 888. 1922. — Fig. 4.

MALAYA: PAHANG; Cameron Highlands, J. Wyatt-Smith s.n. (KEP).

SELANGOR: Genting Sempah Ridge trail, 2500 ft. alt., 11.11.1970, Stone 9578 —

(KLU).

5. Brassaiopsis minor B. C. Stone, sp. nov. — Fig. 5.

Arbuscula, ramulis 5 mm diametro. Folia simplices elliptica marginibus in

partem mediam distalem serratis; petiolo 6-8 cm longo c, 1.5—-2 mm diametro basi

breviter vaginantio; lamina tenue coriaceo basi subtriplinervio nerviis c. 6-8

Araliaceae 283

FRI-1274]

Fig. 3. Brassaiopsis simplex (King) Stone. (Wardenia simplex King). Leaf, inflorescence,

floral details, stipule detail, pubescence. FRI-12741.

284 Gardens’ Bulletin, Singapore — XXX (1977)

0 90

4 9

= D

- *,

A\ a ‘ s

o hi Yee

- A

= 4

YX .

4 @

Fig. 4. Brassaiopsis elegans Ridl. Leaf, inflorescence, floral details. Stone 9578.

Araliaceae 285

Fig. 5. Brassaiopsis minor Stone. Leaf, stipules, bracts (left: dorsal, right: ventral),

inflorescence, and floral details (B':— the lowest inflorescence bract with a stipular-

ligulate base). Type collection.

286

Gardens’ Bulletin, Singapore — XXX (1977)

Fig. 6. Comparison of Schefflera singularis Stone with two Dendropanax species

A:—Schefflera singularis, leaf and floral details, type collection.

B:—Dendropanax maingayi King, leaf and floral details, KEP 66583. and fruit,

Poore 1072.

C:—Dendropanax lancifolius Ridl. cfr., leaf and floral details, Symington 25458.

Araliaceae 287

paribus, usque ad 25 x 12 cm, apicem versus abrupte acuminato, basi cuneato.

Inflorescentia terminalis, brevis, 4-5 cm longis, bracteatis, pauci-vel non-ramosis,

umbellatis; bracteolis deltoideis c. 1-1.5 mm longis rufo-puberulento-ciliolatis;

pedicellis c. 15 mm longis; calyce 5-dentato; ovario rotundato-lobulato, depresso,

biloculare, stylis binis unitis columna 1 mm longo formantibus. Loculi ovarii 2,

uniovulati.

Holotype: MALAY PENINSULA: TRENGGANU, Gunong Pahang, Ulu

Brang, camp 3, humid deep valley on granite; alt. 3500 ft., fleshy treelet, flowers

white; 21.9.1969, Whitmore FRI-12741 (KEP).

Additional spec. examined: TRENGGANU: 34th mile. Kuala Trengganu, Besut

Rd., lowland forest, (W. side), woody 2’ high, 8.9.1955, J. Sinclair & Kiah SF-

40775 (SING).

This is another simple-leaved species of Brassaiopsis. Originally I had con-

sidered it as a new Wardenia. The flowers show clearly that the ovary is bilocular

and there is a single ovule in each loculus. The slender shorter petioles, elliptic-

cuneate leaves and shorter inflorescence quickly distinguish this from B. simplex.

Hederopsis King

Type species: Hederopsis maingayi King, Hk. FI. Brit. Ind. 2: 739. 1898.

Originally described from rather inadequate material, this genus (endemic in

Malaya and apparently Sumatra, monotypic) has in the last few years been

collected rather widely and is in fact locally common in a few localities. It has been

suggested by Frodin that the genus could be merged with Macropanax Miq. which

appears to differ only in its usually 2-3-locular ovary, this structure in Hederopsis

being normally 5-6-locular. The difference is a slight one, but King’s genus can be

retained for the time being until a broader-based comparison can be made. The

leaves are palmate-digitate, often with 5 leaflets, or with 3, or even 4, and sometimes

(on flowering branches) reduced to 1 leaflet. The slender petioles may reach a

length of 56 cm, and the terminal leaflet (always slightly longer than the laterals)

a length of 31 cm and a width of 12 cm. The margins are dentate with small rather

distant teeth. The petiolules may reach a length of 15 cm (again shorter on the

lateral leaflets). — Fig. 7.

The spirally disposed leaves have a rather short sheathing base about 1 cm

long or less; when very young the margins are ciliate with pale brown multiseptate

hairs, which are often laterally adnate in flat ‘bundles’. The juvenile stems are

similarly rather densely hairy, but the hairs are ephemeral and all mature and

submature shoots are glabrous. The base of the inner surface of the sheath shows

a purplish spot adjacent to the axillary bud. The bud is compressed, purplish,

pubescent, and bears 2 imbricate minute prophylls (which are more or less deltoid

when flattened); these are equivalent to the leaf-sheath and show the ciliate margin

and, often, a more or less produced apical elongation sometimes bearing three tiny

lobes (suppressed leaflets). The outer face is usually rather pubescent. Similar

prophylls occur on the terminal shoot in the uppermost region during renewed

growth following the resting period, which is evident from the succession of several

long internodes followed by a short series of very crowded internodes on the main

branches and leafy shoots.

Individual plants of this species become attractive small trees with a rounded

crown, the trunk sometimes exceeding 90 cm girth, often fluted at base, giving

off a few main branches; the whole tree may reach 15-20 m height.

In Ulu Langat (Selangor) above the Reservoir (Ponsoon) there are several

of these trees, copiously fruiting in January-February 1975, The fruits are sub-

globose and still green when ripe.

288 Gardens’ Bulletin, Singapore — XXX (1977) |

Fig. 7. Hederopsis maingayi King. Leaf and floral details, and fruit.

Araliaceae 289

Spec. examined: MALAY PENINSULA: KeEpaH: Ulu Muda Forest Reserve,

Compartments 115 & 116, 1500 ft. alt., a tree 40 ft, high, girth 24”, bole fluted, with

small buttresses, bark rugose, lenticellate, inner bark thick brown, with watery

exudate, wood white, fruits green. 21.1.1969, Y. C. Chan FRI-6782 (KEP).

PERAK: Bintang Hijau F. R., secondary forest, tree 50 ft. tall, 26” girth, flowers

pale yellow, faintly aromatic, cut bark smelling of resin, bole slightly fluted,

22.11.1966, Rahim Ismail KEP-98531 (KEP). KELANTAN: Ulu Sungei Lebir

Kecil, 500 ft. alt., 17.9.1967, P. F. Cockburn FRI-7123 and 7123 (KEP). SELANGOR:

Ulu Langat, 2 miles from Kg. Injin Satu, tree 50 ft. high. 17.4.1963, Abdul Samat

196 (KIU). — Ulu Langat Forest Reserve, tree 30 ft. high. 9.3.1966 Whitmore

FRI-106 (KEP). — Ulu Langat, Bukit Lakwak, Kg. Panson, n.v. (Temuan)

““Kayu pacha” 4.10.1958, Gadoh, Phytochem. Survey 1015 (KEP). — Ulu Langat

above NEB works, stream valley, tree 10 m x 35 cm, fruits green, 7.2.1975,

M. M. J. van Balgooy & B. C. Stone 2249 (KLU etc.). PAHANG: Ulu Tembeling,

lowlands, 28.5.1931, Henderson SF. 24550 (SING). — Ulu Gali, Raub, 27.9.1929,

Kalong KEP-20274 (KEP). — Cameron Highlands, K. Mensum to Boh Estate,

24.9.1971, H. S. Loh FRI-17399 (KEP). — Same locale, 3.3.1972, alt. 2000 ft.,

tree 40 ft., girth 30”, Y. C. Chan FRI-19932 (KEP).

Scheffiera J. R. & G. Forst.

Schefflera singularis B. C. Stone, sp. nov. — Fig. 6A.

Liana glabra innovationibus inflorescentiaque excepto, pilibus perminutis

stellato-quinque-brachiatis; ramis usque ad 1 cm diametro teretibus cortice griseo;

ramulis foliaceis usque ad 6 mm diametro in sicco longitudinaliter canaliculato-

ruguloso; foliis simplicibus elliptico-obovatis marginibus valde sed anguste

revolutis apice obtusis vel minute apiculatis vel rotundatis vel retusis, basi cuneatis,

rigide percoriaceis costa in canaliculo costali utrinque leviter elevato, nerviis

invisibilis; petiolo 5-20 mm longo. 1.5— mm crasso in sicco ruguloso basi ligulato

ligulo 5 mm longo marginato; lamina 2-12 cm longo 1.5-6 cm lato apiculo

0.5-2.5 mm longo vel nullo; nerviis lateralibus c. 5-7 paribus sed omnino obscuris,

nerviis basalibus marginam formantibus. Inflorescentia terminalis bisumbellatis

(composito-umbellatis) usque ad 3 cm longis axibus griseo-albide furfuraceis

demum glabrescentibus bracteis deltoideis c. 4 mm longis bracteolis c. 1-2 mm

longis scarioso-ciliato-marginatis. Umbellae c. 5-10-floriferae, pedicellis c.

4 x 0.8 mm, calyce obconico 3 x 3 mm margine obscuriter 5-dentato minutissime

ciliolulato, petalis 5 deltoideis 2 x 1.8 mm intus obscuriter carinatis apicibus

unguiculatis, staminibus 5 filamentis albidis c. 1 mm longis antheris submedifixibus

ovatis 1 mm longis erectis, disco leviter elevato, stylopodio late conico truncato

vix 1 mm alto 2 mm lato, stigmatibus 5 obscuris radiatis ad apicem stylopodii

sessilibus. Fructus immaturis glabratus obconico-turbinato-hemiglobosus 5 x 4 mm

calyce undulato alte-marginatus stylopodio centrale 1 mm alto.

Holotype: MALAYA: PaHANG, Gunung Ulu Kali, summit area, 5000-5800

ft. alt., elfin forest with Dacrydium, big shrub with thick brittle leaves pale tawny-

greenish beneath, 23.7.1967, B. C. Stone 7175 (KLU).

Other specimens examined: MALAY PENINSULA, PAHANG, Gunung Ulu

Kali, summit area, shrub in mossy forest, 19 Sept. 1967, J. Dransfield KLU-8068

(KLU); same location, 27.6.1971, Mahmud b. Sidek and W. R. Stanton KLU-

16312 (KLU). Ginting Highlands Rd. 4.5 miles (c. 3000’), 12.6.73, Mohd. Shah &

Md. Ali MS. 2987 (climber? — fis. dark green in bud), (SING). Isotypes in

I and KLU.

290 Gardens’ Bulletin, Singapore — XXX (1977)

This very distinct endemic species so far has been collected only on the

summit area of the mountain Gunung Ulu Kali, in the Ginting Highlands of

Pahang, on the Selangor border. At first it was considered to be a new species of

Dendropanax but after much discussion and hesitation it is here described as a

Schefflera. In nearly all features this species agrees with the formal definition

of a Dendropanax, but differs in having a perceptible though limited ligular process

at the base of the petiole, similar to that in at least most Scheffleras. In addition

the exceedingly coriaceous leaf, with virtually invisible venation, appeared

inconsistent with the condition in Dendropanax, which at least in the Malaysian

species, is characterized by rather evident and subtrineryed leaves, as may be seen

in D. maingayi and lancifolius, illustrated in Fig. 6 B and 6 C. Schefflera singularis,

as shown in Fig. 6 A, not only has invisible nerves and evident stipular ligule, but

a more oboval laminar shape than the two species of Dendropanax. Nonetheless,

in floral characters there is almost no difference between the new Schefflera and

the two Malayan Dendropanax species, as the figures cited also display.

Within Schefflera the vast majority of species possess compound leaves with

trifoliolate or palmate leaflet arrangements, Rarely, as in Schefflera lanceolata Ridl.,

simple leaves occur, though even in this species ojder branches may show

trifoliolate leaves. Another such example is Schefflera simplicifolia Merr. of the

Philippines. Among Malesian species I know of no other similar cases, though

they may occur. Thus Schefflera singularis stands apart from nearly all other

Malayan Scheffleras by its simple leaves.

Another difference lies in the form of the inflorescence, which in Schefflera is

usually a compound umbel, i.e. an umbel of umbels or umbellules, while in

Dendropanax the umbel is often simple; but there are also species of Dendropanax

with compound umbels.

The genus Schefflera includes species which are climbers, others which are

shrubs or trees, the latter often epiphytic. Dendropanax species are apparently

mostly shrubs or trees. Schefflera singularis is a high-climbing woody liana.

A comparison of generic diagnoses in such standard works as various floras,

treatise* on the Chinese Araliaceae (Sargentia 2. 1942), and Vol. 2 of Hutchinson’s

“Genera of Flowering Plants” (1967), does not appear to reveal any completely

dependable differential features to distinguish Schefflera from Dendropanax except

perhaps the presence of a developed stipular ligule. The degree of connation or

separation of the style-tips seems to be highly variable within Schefflera and not

a diagnostic character.

In the light of these facts it is evident that further work is necessary to clarify

the distinction between Dendropanax and the small number of Schefflera species

with predominantly simple leaves. One may in fact question whether Dendropanax

lancifolius Ridl. is not itself misplaced in Dendropanax. (In passing it may be noted

that the generic name Dendropanax, although treated as neuter by the original

authors, has usually been considered masculine in gender, and thus the epithet is

spelled here Jancifolius rather than lancifolium as it appears in Ridley’s Flora).

Acknowledgements

_it is a pleasure to record the debt owed to Prof. W. R. Philipson of the

University of Canterbury for stimulating discussions concerning Araliaceae and

for his several suggestions which have improved and augmented this paper. I am

also grateful to D. E. Frodin, University of Papua New Guinea, and Prof.

? Bye. i. Li.

Araliaceae 291

I. V. Grushvitzky, Komarov Botanical Institute, Leningrad, for their helpful

advice. Thanks are also due to the Director and Curators of the Botanic Gardens,

Singapore, and the Forest Research Institute, Kepong, Malaysia, for the opportuni-

ties to study the specimens of Araliaceae in their care. Shamsuddin bin Abdul

Jalil, a former Botany Honours student in the University of Malaya, assisted in

this work by carrying out a useful preliminary study of the Malayan species of

Arthrophyllum as an Honours thesis under the writer’s supervision, and his

contribution is gratefully acknowledged.

Reference

Philipson, W. R. 1970. The Malesian species of Gastonia. Blumea 18: 490-495.

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A Note on the Cytology of Botrychium lanuginosum

and the Occurrence of the Genus in Malesia

A. C. JeERMy* and T. G. WALKER**

Summary

Over the past twenty-five years intensive plant collecting has shown two species of

Botrychium (B. lanuginosum Wall. ex Hook. & Grev. and B. australe R. Br.) to be present in

eastern New Guinea, the former hitherto confined to Central—S.E. Asia and reaching to the

Central Himalaya and the latter Australasian with a possible representative in S. America. A

further species, B. daucifolium Wall. ex Hook. & Grev., is distributed from Sri Lanka

(Ceylon) to the Philippines and Sulawesi (Celebes) with questionable outlying records in Fiji

and Samoa, reaching S. China in the North, and replaced by a taxon of doubtful specific

identity in Japan (B. japonicum (Prantl) Underw.). B. lanuginosum was found in large quanti-

ties in the kunai grasslands at 1600 m in the Finisterre Mountains of E. New Guinea and

material fixed in the field has proved to be octoploid m = 180) and also hexaploid,

suggesting the tetraploid cytotype recorded by other workers in India and Sri Lanka may

also be present in New Guinea.

INTRODUCTION

The appearance of Botrychium in S.E. Asia and Malesia is seasonal. Its

ecological requirements are little known but its appearance may be linked with

the increased rainfall seen just before the full monsoon develops. For this reason

limited collections have been made as the aerial parts of the leaf quickly become

yellow and die down after the spores are shed. Three species have so far been

recorded in Malesia, B. lanuginosum Wall. ex Hook. & Grev., B. daucifolium Wall.

ex Hook. & Grev. and B. australe R. Br. The first and last species are components

of well drained soil in the open anthropogenic grasslands in Malesia (from

1600 — 2700 m in New Guinea) and on similar grassy banks or more rarely in

woods on the Asiatic mainland. B. daucifolium is a species of more shady situation

often in open woodland in the Himalaya and on the open floor of the rain forest

in Borneo and Sulawesi. Betty Molesworth Allen (1959) discovered a colony at

1500 m on Gunong Perdah in an opening in the tall forest, growing in a scrub of

ogg i and young Cyathea. She records the soil as being a black sticky

umus.

KEY TO MALESIAN SPECIES OF Botrychium

1 Fertile segment arising from the rachis of the sterile blade, ++ equal in length

to.adjament. sterile pinnac ........ 000? aaah onan atee....... 1 lanuginosum

1 Fertile segment arising from the petiole below the sterile blade; the stalk of

the fertile segment 1 — 2 times as long as the sterile blade

2 Fertile stalk + equalling sterile blade; upper pinnae linear-lanceolate or

triangular, pinnatifid or dentate only ....................0.ccccceee eee 2 daucifolium

2 Fertile stalk at least twice as long as the sterile blade; upper pinnules ovate,

Se kv eer ne EA Pep, ee \ nee 3 australe

* Botany Department, British Museum (Natural History), London, England.

** Department of Plant Biology, University of Newcastle upon Tyne, England.

293

294 Gardens’ Bulletin, Singapore — XXX (1977)

Cytology, Botrychium 295

1 B. lanuginosum Wall. ex Hook. & Grev. Icon. Fil. tab. 79 (1829).

The name first appeared in Wallich’s Catalogue of 1828 without a description

and was described by Hooker, the figure by Greville (/.c.) drawn from material

supplied by Wallich. Ballard (1940) discusses the typification and describes the

specimen in Greville’s herbarium at the Royal Botanic Garden Edinburgh from

which the two lower pinnae were removed, thus giving rise to the abnormal! plant

seen in his plate 79.

The species ranges from the Central Himalaya across China to Taiwan, south

to Sri Lanka on the Indian subcontinent and in Malesia to Luzon, Sumatra, Java

and New Guinea. The species is distinct in having a broadly deltate tripartite

sterile blade, the lobes usually 3-4 pinnatisect, and with the fertile scape

arising from the rachis above the insertion of the two lower pinnae (see Figs

1 & 2). The bud, rachis and especially the costal axils are usually sparse to

densely covered with long unbranched hairs although some fronds in the New

Guinea population mentioned below were almost glabrous. Hayata (1914) has

described the glabrous form as B. leptostachyum, later to be combined by Nakai

(1925) as a variety of B. Januginosum, but in view of the wide variation in the

above population both are of doubtful status. The degree of dissection of the sterile

blade varies, younger plants being usually less compound, but always more

dissected than either B. daucifolium or B. australe. There is no apparent correlation

between ploidy level and degree of cutting or size of sterile frond (see Figs 1 & 2).

2 B. daucifolium Wallich ex Hook. & Grev. Icon. Fil, 2, tab. 161 (1830).

Wallich (1828) listed the first gatherings of this (No. 49) as B. subcarnosum

and later by the name Hooker took up for it. The specimen drawn is in the

Edinburgh herbarium and other plants of the same gathering at the British

Museum (Natural History) London,

The species ranges from a similar area in the centra) Himalaya east to Taiwan

and south to Sri Lanka. In Malesia it is found throughout the archipelago to

Sumbawa Is. (G. Batulanteh), Celebes (G. Bonthain) and Mindanao (Mnt Apo)

and Negros (Cuernos de Negros). Allen (1959) found two populations in the

Cameron Highlands area of Pahang. There are specimens in the Kew Herbarium

from Upola, Samoa and Nadavivatu, Fiji which compare well with material of

B. daucifolium from Asia, but a closer study of more recent material needs to be

made.

B. daucifolium is distinct in having thin, but not membranous, lamina tissue

and a less dissected frond in which the upper pinnae are barely cut (see Fig 3)

but lobed with the segments toothed. The fertile spike arises from the petiole about

10 cm or more below the insertion of the blade and the fertile segments are strict,

not spreading as in the other two species. The species varies in its hairness, some

species in Borneo being quite densely pubescent on the rachis and costae, Allen

(1959) mentions that the very young fronds on her Malayan plants were covered

with copious pale hairs. Himalayan plants tend, in herbarium specimens at least,

to be more glabrous, as do the adult plants of Allen, who gives a fine general

photograph of the species (/. c. facing p. 251).

Material from Japan, E. China and some from Taiwan, being more mem-

branous, more deeply toothed and with the fertile stalk somewhat longer, has been

separated off as B. daucifolium var. japonicum Prantl, raised by Underwood

Facing page

Figs. 1-4. Silhouettes of Botrychium. 1 & 2. Silhouettes of Botrychium lanuginosum from

Moro, Finisterre Range, New Guinea showing range of dissection. 1: Jermy 3887

(x 4); 2: Walker 8333, octoploid plant from same population (x 4). fs = fertile spike.

3: B. daucifolium Gunong Dempo, Sumatra, C. J. Brooks 15881 (x 4). 4: B. australe

Western Highlands District, New Guinea J. S. Womersley, NGF 15213 (x 4).

296 Gardens’ Bulletin, Singapore — XXX (1977)

(1898) to a species. It is certainly related in its morphology to the typical mainland

asian B. daucifolium. A small form was described by van Alderwerelt van

Rosenburg (1911) from Preanger Regencies, Java under the name of var. parvum.

Later in 1913 the same author described yet another variety, subbasalis, which was

not only smaller than usual for B. daucifolium but had the fertile spike arising

towards the base of the stipe as in B. ternatum and B. australe. Clausen (1938)

considered this, from the published description only, to be a possible new species.

We have not studied van Alderwerelt van Rosenburgh’s type and must reserve

judgement but it is our experience that juvenile and depauperate Botrychiums can

appear very abnormal.

3. B. australe R. Br. Prod. Fl, Nov. Holl.: 164 (1810).

Collected and described by Robert Brown from material from Tasmania and

New South Wales. The specimen in the herbarium of the British Museum (Natural

History) from Parametta, Port Jackson can be taken as typical but the specimen

at Kew is more complete (it contains roots) and is best to be designated lectotype.

The species ranges from South Austra'ia to New Zealand and north into south

Queensland where it was recorded.as B. ternatum (Thunb.) Sw. by Bailey (1883).

It differs from the latter in its dissection of the thicker lamina which has hyaline

marginal cells.

The sterile leaf is 10-20 cm long, ternate, with the central leaflet larger and

more compound. The upper pinnae of this leaflet are ovate and cut to the costa,

the resulting segments glabrous, linear and dentate at the truncate apex; the tissue

is fleshy and the lamina margin is made up of two to three rows of hyaline cells. The

compoundly branched and spreading fertile segment arises in the lower half of the

stipe to a height of 30 cm. (see Fig. 4).

The map in Fig. 5 shows the range of the species in Australia. Its presence

in Papua New Guinea in the Southern and Western Highlands Districts is a logical

extension of this range. It grows in high altitude anthropogenic grasslands

(Miscanthus-T hemeda associations) and around 2700 m in Danthonia-Poa grass-

land and Cyathea scrub and in two localities together with B. lanuginosum. The

taxonomy of this australasian complex is little understood and is being reviewed

by New Zealand workers at present and their results are awaited with interest.*

THE CyToLocy oF B. lanuginosum IN NEw GUINEA

Whilst collecting fern material for subsequent taxonomic and cytological studies

in Madang District, east New Guinea in 1964 the authors showed children in

Moro, a village at 1680 m at the head of the Gusap River in the Finisterre

Mountains, an illustration of Botrychium, Within the hour the children returned

with armfuls of B. lanuginosum collected from amongst the kunai (Imperata

grass) on old garden sites. This is a good example of the dangers of this method

of collecting easily recognisable but possibly rare species. Luckily this population

could withstand such collecting and some 50 specimens were taken to show the

range of variation. Seven specimens thus collected had meiotic material and were

fixed in the field.

One specimen (Walker 8333) showed exactly 180 bivalents at meiosis (Plate 1)

and three others (Walker 8332, 8334, 8336) 270 chromosomes and hybrid meiosis.

The base number for Botrychium is well established as 45 and thus these New

Guinea specimens were octoploid and hexaploid respectively. This contrasts with

other counts for the species from the Indian subcontinent where Verma &

Loyal (1960) recorded n=90 in a Himalayan specimen and Ninan (1956)

a Se Re RENN Mintle a

* Dr J. Braggins, University of Aukland, N.Z., is of the opinion (pers, comm., 1976) that the

New Guinea specimens constitute a new subspecies.

Cytology, Botrychium 297

recorded n=90 and 2n=180, also for N India; Manton and Sledge (1954) showed

a similar situation in Sri Lanka (n=90). Hence all previous counts are at the

tetraploid level. The fact that we have hexaploid material showing hybrid

behaviour suggests that the tetraploid may also be present in New Guinea.

)

ee ee ee Se ee ee ee

B. lanuginosum

B. australe

5. Distribution of the three species of Botrychium found in Malesia.

298 Gardens’ Bulletin, Singapore — XXX (1977)

REFERENCES

Allen, B. Molesworth (1959) Malayan Fern Notes, A preliminary note. Gard.

Bull. Singapore 17: 251-252.

Bailey, F. M. (1883) A synopsis of the Queensland Flora. Brisbane.

Ballard, F. (1940) Notes of ferns and fern allies IJ. Kew Bull. 1940: 29-31.

Clausen, R. T. (1938) Monograph of the Ophioglossaceae. Mem. Torr. Bot. Club

19 (2): 3-177.

Hayata, B. (1914) Icones Plantae Formosae 4: 134. Taipei.

Manton, I & Sledge, W. A. (1954) Observations on the cytology and taxonomy

of the pteridophyte flora of Ceylon. Phil. Trans. roy. Soc. Lond. 283B: 127-185.

Ninan, C. A. (1956) Cytology of Ophioglossaceae. Curr. Sci. 25: 161-162.

Nakai, T. (1925) Bot. Mag. Tokyo 39: 192.

Underwood, L. W. (1898) American ferns I. The ternate species of Botrychium.

Bull. Torr. Bot. Club 25: 521-541.

Van Alderwerelt van Rosenburg C. R. W. K. (1911) New or interesting Malay

ferns 3. Bull. Jard. Bot. Buitenz. ser. 2 (1): 3.

(1913) New or interesting Malay ferns 5. ibid. ser 2 (11): 6 et

tab. 3.

Verma, S. C. & Loyal, D. S. (1960) Chromosome counts in some ferns from Naini

Tal. Curr. Sci. 29: 69-70.

Wallich, N. (1828) A numerical list of dried specimens of plants in the East India

Company museum collected under the superintendence of Dr Wallich, London.

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Plate 1. Permanent aceto-carmine squash of metaphase I of a spore mother cell of Botrychium

Ranugeniosts (Walker 8333) together with explanatory diagram below. Magnification

1000.

A Revision of the Malesian Species of Arthrophyllum BI.

(Araliaceae)

W. R. PHILIPSON

Department of Botany,

University of Canterbury,

New Zealand.

The genus Arthrophyllum forms a well-defined group of species within the

Araliaceae. Its principal distinctive characters are the single-celled ovary and the

arrangement of the umbels on specialized flowering branches. The vegetative

shoots bear pinnate or bipinnate leaves in spiral phyllotaxis. Branches do not

develop until flowering occurs, when a terminal] tuft of lateral branches develops.

These bear opposite leaves of reduced size and terminate in a whorl of secondary

branches which may themselves bear umbellules or which may branch again

before doing so (fig. 1). Vegetative growth is continued by one or more innova-

tions, with spirally arranged leaves, arising below the crown of flowering branches.

The geographical range of the genus almost coincides with that of Flora

Malesiana, but extends slightly further west to the Nicobar Islands and northwards

into Laos and Thailand, and the range does not include the Lesser Sunda Islands.

Apart from the treatment in Ridley’s Flora of the Malay Peninsula, no revision

of the species has appeared. The many taxa given specific names by early investi-

gators (most of which are here regarded as synonyms) were never critically

compared, so that knowledge of the genus has remained confused and fragmentary

and botanists have tended to avoid it. Consequently it has been found necessary

to describe several new species, while at the same time reducing several others

to synonymy.

The present revision is regarded as only tentative. Herbarium material is

abundant, but most specimens are fragmentary and therefore fail to illustrate the

_unusual branch-systems which characterize some species. Specific distinctions

seem to be unusually difficult to define on the basis of herbarium specimens, so

that knowledge of the group will remain superficial until a comprehensive field

study can be undertaken throughout the range of the genus.

The genus comprises two species-complexes, and also the anomalous

A. proliferum. One of these complexes includes seven shrubby species with simpler

inflorescences (no more than three degrees of branching), namely, A. maingayi,

A. montanum, and A. alternifolium (from the Malay Peninsula, with the first

species also from Sumatra and Borneo); A. papyraceum (from Sumatra); A.

pulgarense and A. cenabrei (from the Philippines); and A. kjellbergii (from

Celebes). The second complex consists of nine species of tree stature with larger

inflorescences (having four degrees of branching). This complex occurs commonly

throughout the range of the genus, and its subdivision into species presents con-

siderable difficulty. To the east A. macranthum is abundant on the mainland of

New Guinea, while on islands to the north a second species, A. pacificum, is

recognized. In the Philippines and northern Moluccas this complex is represented

by A. ahernianum, a species which also extends into the extreme north-east of

Borneo. The complex is more difficult to classify further west and south, that is,

in Celebes, Borneo, Java, Sumatra, and the Malay Peninsula. In the present

299

Gardens’ Bulletin, Singapore — XXX (1977)

300

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Arthrophyllum ‘301

revision, in addition to the widespread A. diversifolium, five entities from this

area are given specific rank: three species endemic to Borneo (A. crassum, A.

collinum, and A. ashtoni; one most distinctive species, A. angustifolium, which

is rare in Borneo and the Malay Peninsula; and a species, A. engganoense, which

is known only from the small island, Enggano, off the west coast of Sumatra. The

remaining great mass of material is here treated as a single species, A. diversifolium.

The variation and nomenclature of this widespread taxon is discussed separately

under the section on that species.

ARTHROPHYLLUM

Arthrophyllum Bl. Bijdr. Fl. Ned. Ind. ( 1826) 878. Mormoraphis Jack ex Wall.

Cat. n. 4931 (1831).

Sparingly branched trees or shrubs, unarmed. The leaves on vegetative shoots

spirally arranged, imparipinnate, those on flowering branches often opposite,

smaller, or reduced to a single leaflet; stipular sheath small. Inflorescence con-

sisting of compound umbels, either solitary and terminal or more commonly borne

on a cluster of specalized leafy branches arising from the axils of the uppermost

leaves; pedicels not articulated. Flowers hermaphrodite; calyx an undulate rim;

petals 4-6, valvate in bud; stamens 4-6, anthers curved, basifixed; ovary turbinate,

l-celled; disk fleshy, rising in the centre to the + sessile capitate stigma. Fruit

ovoid or spheroidal, often oblique; exocarp leathery; endocarp cartilaginous; seed

solitary, pendulous; endosperm deeply transversely ruminate.

About 18 species (17 in Malesia) extending from the Nicobar Islands and

in the larger islands or regions is as follows: Sumatra 4; Malay Peninsula 5; Java

1; Borneo 7; the Philippines 3; Celebes 2; Moluccas 2; New Guinea and Bismarck

Archipelago 3. The genus Eremopanax, with several species in New Caledonia,

may be congeneric,

KEY TO THE SPECIES ...

1. a. Inflorescence becoming paniculate by the successive development of

branches below the umbellules (figs. 3, 4) .................. I. A. proliferum

ge Be Bl) Ren ee rn oe 2

2. a. Inflorescence with four orders of branching (fig. 2) as ahes DE No 3

b. Inflorescence with three (or fewer) degrees of branching (figs. 5, 6)

PCRS SETH SEES EEE HEHH SHS EHEHEHHE HEHEHE EHH HET HEHTHEHEHEHESHE HEHEHE HEHEHE HEHEHE HEHE HEHEHE HEHEHE HHH HEHEHE HEB E ES

ee ec dha cr asncescccserccnnacusvscceseccee ce 4

b. Leaflets broader rie Re habia oe nares Sct diiet ries , ‘delieenednante 5

4. a. Leflets c. 4-7 cm NN OR ok ai ek. Se CO TIC 2. A. ashtoni

b. Leaflets much longer ............0....66, NRIs. 3. A. angustifolium

5. a. Pedicels (at early anthesis) + 10 mm long .................ecceeeeeee ees 6

b. Pedicels (at early anthesis) c. 5 mm long, or shorter Maye Be, 7

6. a. Peduncles of umbellules with bracts or bract-scars (fig. 13) .........:00008

AO Elie nck eels ate ANE Vilas care Fp eyigs pF 8S Cunt beg Hew ps Shaye gd since y «fp 4. A. ahernianum

b. Peduncles of umbellules (fig 14) without bracthint ‘goassieivol.wil........

Sich bedns 6s xs aeteRNGEe SMO) die Ws. UL. JAI. BALI 28. ‘5. A. engganoense

302 Gardens’ Bulletin, Singapore — XXX (1977)

7. a. Leaves associated with the umbels rotund (fig. 17) ..............:esueseeeeees

yp... o). .nenio8. ol aunehor. zvicage. oauds..:Ated. aes 6. A. collinum

b. Leaves (or leaflets) associated with the umbels ovate or elliptic ......... 8

8. a. Leaves associated with the umbels ovate, + fleshy, with the lower surface

smooth (lateral veins obscure) (fig. 18) .................. sence 7. A. crassum

b. Leaves (or leaflets) associated with the umbels + elliptic, coraceous or

9. a. Young parts glabrous ....../....0i tide cesee eee ece scene 8. A. pacificum

b. Young parts with rufous tOmentuM .........05...ceseee separ -cosceenezecrsens 10

10. a. Umbels at anthesis with numerous filamentous pedicels (fig. 15) .........

iniawas. ao. 22ve0!. oAT:: bomany-.acdwade 26-4808 ben 9. A. diversifolium

b. Umbels at anthesis with fewer stout pedicels (fig. 16) ...................0.05.

FT SSI? | BOS , SOLOS tee her ton Soe eect 10, A. macranthum

11. a. Leaflets membranaceous or chartaceous ................cceeee eee eee ene en ees » he

b: Leaflets coriaceous 2) 0/ 00225. 2002.2... PRP. 4001, GE ae ... 14

12. a. Mid-leflets about 16 cm long ................c cece seen ees 11. A. papyraceum

b. Mid-leaflets + 8 cm long or shorter ............... cece cece ee eeeeeeeeneenes 13

13. a. Leaflets usually 5-7 (fig. 7), primary inflorescence branches usually

short (+ 3-6 cm) and without articulations ............... 12. A. maingayi

b. Leaflets more numerous (fig. 10), primary inflorescence branches longer

(10-20 cm) with one or more leafy nodes ............... 13, A. kjellbergii

14. a, Leaflets c 5: ...celi iio eee 14. A. cenabrei

b. Leafléte more nWNCTOUS 55 Feats oan senda de atesagtds o-oo vnunet peeaiee sail ae 15

15. a. Petals & stamens B>.. .<cpts <i;44-. sis erseshilionan 15. A. pulgarense

b.... Petals & stamens. 4. Of) Bic, 103s coadesbomsvesss she ¥t “es rdead nadienea eee 16

16. a. Leaflets 6 cm long, or longer (fig. 8) .................000. 16. A. montanum

Leaflets shaster CBR) 39) sib dantbchocnpatiebhisebonds 17. A. alternifolium

1. Arthrophyllum proliferum Philipson, sp. nov.

Arbor foliis imparipinnatis. Inflorescentiae ex suis propriis ramis natae,

paniculatae factae ramis sub umbellulis iterum atque saepius ramificantibus. |

_ Medium sized tree, glabrous, Leaves of the vegetative stems spirally arranged,

imparipinnate, multijugate, 90 x 24 cm, of the flowering branches smaller with

fewer pinnae or usually simple; leaflets obovate-oblong, c. 16 x 7 cm, chartaceous,

margin entire, slightly revolute, base broadly cuneate to truncate, sometimes

oblique, apex acute, Inflorescences terminating specialized plagiotropic shoots,

bearing axillary flowering branches and ending in umbellules of a few flowers

below which pairs of whorls of branches conitnue the growth of the inflorescence

to produce an elongated panicle of umbellules.

_ Endemic to Papua New Guinea. Type: Territory of New Guinea, Morobe

District, Mauwata Banda L.A., Bulolo, Havel and Kairo NGF 17172 (Canberra).

In mid-mountain rain forest, reaching the canopy, on steep slopes, 300-1200 m.

The flower and fruit are typical of this well-defined genus, but the branching

of the inflorescence is unlike that found in all other species.

Arthrophyllum 303

Figs. 3-6. 3. A. proliferum; a portion of inflorescence in the fruiting state (x 4);

4, same, showing detail of ultimate branches of the inflorescence in the flowering

stage (x 1). 5. A. maingayi, terminal inflorescence (x 4). 6. A. montanum,

a single flowering shoot (x 4).

2. Arthrophyllum ashtoni Philipson, sp. nov.

Abor gracilis tomento fusco. Folia imparipinnata, foliolis lanceolatis vel

late-lanceolatis, 3-7.5 x 1-2 cm. Inflorescentia ramis in proporium officium

mutatis et in coronam terminalem dispositis, quarter ramificantibus.

A slender small tree, to 5 m high, young parts with brown scurfy tomentum.

Leaves of the vegetative stems spirally arranged, imparipinnate, multijugate, to

about 30 cm long; of the flower-bearing branches smaller with fewer pinnae, or

simple; leaflets lanceolate to broadly-lanceolate; 3-7.5 x 1-2 cm, thinly coria-

ceous, margin entire, revolute, base broadly cuneate, apex tapered to sub-caudate,

veins channelled above, visible below. Inflorescence a terminal cluster of specia-

lized leafy branches; the main rays variable in length in the same inflorescence,

ending in a whorl of secondary rays subtended by simple leaves; the secondary

rays 4-12 cm long, bearing simple leaves, usually in an opposite pair and

terminating in compound umbels; umbellules with about 8-10 flowers pedicels

4-10 mm long, furfuraceous,

Endemic to Borneo. Type: Brunei: Pagon ridge, Ashton Brun 2341 (Kew).

304 Gardens’ Bulletin, Singapore — XXX (1977)

Moss forest on sandstone ridge, and Kerangas forest, 1000-1550 m.

The small narrow leaflets are very distinctive. No other species with small

leaflets has inflorescences which branch to the fourth degree.

3. Arthrophyllum angustifolium Ridley in Journ. Fed. Mal. St. Mus. 10 (1920)

136.

A shrub or small tree, 5 m high, young parts rufous tomentose, becoming

glabrous. Lower leaves spirally arranged, imparipinnate, multijugate, about

70-90 x 30-40 cm; upper leaves reduced mostly unifoliate, opposite, broader,

with petioles about 2-4 cm long; leaflets coriaceous lanceolate, c. 15-22 xX

1.5-2.5 cm, tapering to an acute or obtuse apex, base cuneate, margin entire,

slightly revolute. Inflorescence a terminal cluster of specialized leafy branches,

the main rays 30 cm (or more) long, bearing simple leaves in opposite pairs

ending in a whorls of c. 10-12 secondary rays subtended by simple leaves; the

secondary rays 8-12 cm long, terminating in an umbel of 5-12 tertiary rays c.

2-3 cm long, each bearing an umbellule of c. 8-12 flowers, pedicels c. 5 mm long:

Malay Peninsula and Borneo. Type: Perak, Gunong Keledang, Ridley 9683

(SING).

Occurs in forest and old regenerated forest on peat swamp at low altitudes

or on ridges.

The lanceolate leaflets are unlike those of any other species, The grey bark is

minutely fissured and bears many small orange lenticels, The wood is soft and

white. The Peninsular and Bornean specimens are similar, except that the flower

buds are larger in the Brunei plant.

4. Arthrophyllum ahernianum Merr. in Philipp. Journ. Sci. 1 (1906) Suppl. 109.

— ‘tA. pinnatum Clarke” F. -Vill. Novis. App. (1880) 103 — Macropanax sp.

Vidal Rev. Pl. Vasc. Philip. (1886) 145. — A. sablanense Elmer in Leafl, Philipp.

Bot. 1 (1908) 331. — A. borneense Merr. in Univ. Calif. Pubs. Bot. 15 (1929)

231, non Baker. — A. elmeri Merr. in Webbia 7 (1950) 319. — A. merrilliana

Furtado, in Gard. Bull, 19 (1962) 185.

Tree up to 15 m high, young parts with rufous tomentum., Leaves clustered

at the ends of the branches, spirally arranged on the vegetative shoots, impari-

pinnate, multijugate, articulated at the insertion of the pinnae; up to 200 x 60 cm;

of the flower-bearing branches smaller with fewer pinnae, or simple; leaflets

ovate-oblong, occasionally oblong-lanceolate, up to 35 x 12 cm, membranaceous

or chartaceous, margin entire, revolute, base cuneate to rounded, usually oblique,

apex short accuminate, Inflorescence a whorl of specialized leafy branches forming

a terminal crown; the main rays up to 150 cm long (or more), bearing pinnate

leaves usually in 1-2 opposite pairs, ending in a whorl of secondary rays up to

30 cm long terminating in compound umbellules; umbellules with about 10-20

flowers; pedicels c. 10 mm long (at anthesis). | |

Extending from northern Borneo, throughout the Philippines to the northern

Moluccas. Type: Lamao Forest Reserve, Luzon, Meyer 2780.

Primary and second-growth forest, from lowlands to 1000 m.

This .species replaces the more westerly A. diversifolium which it closely

resembles. It is characteristically larger in all its parts, particularly in the size of

the individual flowers and the length of their pedicels. There are fewer flowers in

an umbellule. The distinction between these two species is not always easy to

make, especially when the material is fragmentary. A few specimens from the

Philippines e.g. Clemens 585, Merrill 1232 appear very similar to A. diversifolium,

and it is possible that this species extends beyond Borneo, However, at present

Arthrophyllum ri 305

these specimens are regarded as part of the range of variation of A. ahernianum.

Similarly, at least one specimen from southern Borneo (Buwalda 7973) approaches

A. ahernianum in appearance.

5. Arthophyllum engganoense Philipson, sp. nov.

Arbor foliis imparipinnatis. Inflorescentia ramis in proprium officium mutatis

et in coronam terminalem dispositis, quater ramificantibus; umbellulae pedicellis

circa 5-10, c. 10-15 mm longis sub anthesi.

A tree to 21 m high, becoming glabrous. Lower leaves imparipinnate,

multijugate, 60 cm long or more, petiole 24 cm long, 6 mm wide, petiolules

10-18 mm long; leaflets broadly elliptic to elliptic-oblong, c. 12-15 xX 6~7 cm,

base rounded with a short asymmetrical cuneate centre, apex shortly apiculate,

margin entire often undulate chartaceous. Flowering branches c. 40 cm long;

leaves opposite, simple or unifoliolate, ending in a whorl of. simple leaves

surrounding a compound umbel to 30 cm diam.; secondary rays about 8, c.

10-15 cm long, tertiary rays c. 8, slender c. 20-40 mm long, without bracts;

pedicels c. 5-10 per umbellule, c. 10-15 mm at anthesis.

South Sumatra, Enggano Isd. Type: Liitjeharms 4260 (Bogor).

Occurs in forest at low altitudes (up to about 100 m).

The two known collections of this species are very similar and contrast with

the widespread A. diversifolium because of the few flowered umbelles with long,

spreading pedicels.

6. Arthrophyllum collinum Philipson, sp. nov.

Arbor parva in prima aetate tomento rufo. Folia imparipinnate, foliolis

oblongis, late ellipticis vel rotundis. Inflorescentia ramis in proporium officium

mutatis et in coronam termnalem dispositis, .quarter ramificantibus; bracteis

rotundis, coriaceis.

Sparingly branched shrub or small tree up to 12 m high, all young parts with

dense rufous tomentum. Leaves of the vegetative stems spirally arranged, impari-

pinnate, multijugate, up to 60 (or more) X 32 cm; leaflets oblong, broadly

elliptic or rotund, up to 16 X 7 cm, coriaceous, margin entire slightly revolute,

base truncate to rounded, unequal, apex rounded, obtuse, or shortly and bluntly

apiculate, mid-rib prominent, lateral veins usually clearly visible below, upper

surface frequenty rugose. Leaves of the flowering branches smaller with fewer

pinnae or more frequently unifoliolate, leaflets more rotund and with longer

petioles. Inflorescence a cluster of specialized leafy branches forming a terminal

crown; the main rays up to 60 cm long, bearing one or more rarely two opposite

pairs of usually unifoliolate rotund leaves.

Endemic to Borneo. Type: Sabah, Kinabalu, Clemens 33195 (BM).

In forest and scrub from 600-2700 m, sometimes growing as an epiphyte in

the crowns of trees.

The bark is grey and smooth, the wood pale and soft, and the cut stems

exude a yellowish or orange latex. This species is characterized by the rotund

coriaceous leaves on the flowering branches.

7. Arthrophyllum crassum Philipson, sp. nov.

- Arbor parva in prima aetate tomento rufo. Folia imparipinnata, foliolis

ellipticis oblongis vel lanceolatis. Inflorescentia ramis in proprium officium mutatis

et in coronam termnialem dispositis, quater ramificantibus; bractea ovata

plerumque acuta, rigidior alutaequa naturam habens, margine valde Tevoluta,

nervis lateralibus obscurissimis. |

306 Gardens’ Bulletin, Singapore — XXX (1977)

Sparingly branched shrub or small tree up to 8 m high, all young parts with

dense rufous tomentum. Leaves of the vegetative stems spirally arranged, impari-

pinnate, multijugate, up to 100 (or more) X 50 cm, of the flowering branches

smaller with fewer pinnae or more frequently unifoliolate; leaflets elliptic, oblong

or lanceolate, up to 24 X 7.5 cm, coriaceous, margin entire, strongly revolute,

base cuneate to rounded, often oblique, apex narrowed to an acute often caudate

apiculum, mid-rib prominent, lateral veins faint to obscure. Inflorescence a cluster

of specialized leafy branches forming a terminal crown; the main rays up to 70 cm

long, bearing an opposite pair (or rarely more pairs) of unifoliolate or (less

frequently) pinnate leaves.

Endemic to Borneo. Type: Sarawak, Bako National Park, Chai $18020

(Leiden).

Swampy peat forest and heath woodland, both primary and disturbed, from

sea level to about 150 m. Some fragmentary collections (Banying ak Nyudong

S19404; Brunig S8720; Hewitt 770) from higher altitudes (1000 m) further

inland may belong to this species,

The leaves associated with the inflorescence are distinctively fleshy, have a

strongly revolute margin and a smooth lower surface with indistinct lateral veins,

and are ovate with rather acute apex. The leaflets of the pinnate leaves on the

vegetative shoots are also rather leathery with indistinct lateral venation. The

shrubs are occasionally epiphytic.

8. Arthrophyllum pacificum Philipson, sp. nov.

Arbor gracillis glabra, foliis imparipinnatis; foliolis plerumque ellipticis, c.

8-10 x 4-5 cm, membranaceis Inflorescentia ramis in proprium officium mutatis

et in coronam terminalem dispositis, quater ramificantibus.

A slender tree to 14 m high, glabrous. Leaves of the vegetative stems spirally

arranged, pinnate, multijugate, about 60 x 24 cm; of the flower-bearing branches

smaller with fewer pinnae or simple; blade elliptic, oblong or avate, about

8-10 x 4-5 cm, rather membranaceous, mafgin entire, very slightly revolute, base

abruptly cuneate, often oblique, apex obtuse, acute, or slightly apiculate, Inflores-

cence a whorl of specialized leafy branches forming a terminal crown to the

vegetative shoots; the main rays 35 cm long bearing small pinnate leaves in

Opposite pairs.

Extending from the Moluccas to the Bismarck Archipelago. Type: Bismarck

Archipelago, New Britain, Pomio subdistrict, Jsles et al NGF 34424 (Lae).

Primary forest on slopes of mountains, attaining Nothofagus mossy forest,

from 800-2000 m. Reported as common in most localities.

All specimens from islands to the north of New Guinea are similar, in having

more delicate foliage than O. macranthum from the mainland of New Guinea.

9. Arthrophyllum diversifolium BI. Bijdr. Fl. Ned. Ind. (1826) 879. — A.

javanicum Bl. l.c. — A. ellipticum Bl. 1.c. — Mormoraphis sumatrana Jack, ex

Wall. Cat n. 4931 (1831). — Arthrophyllum blumeanum Zoll and Moritz, Syst.

Verz. Pl. Zoll. (1845-6) 41. — A. ovalifolium Jungh. and de Vriese in Ned.

Kruid. Arch. 1 (1846) 19. — A. ovatifolium Jungh. and de Vriese in Ann. Sci.

Nat. 3, 6 (1846) 117. — A. dilatatum Miq. Fl. Ind. Bat. 1 (1856) 768. — A.

diversifolium var. lanceolata Miq. Fl. Ind. Bat. Suppl. 1 (1860) 340. — A.

blumeanum var. oblongatum Miq. in Ann, Mus. Bot. Lugd. -Bat. 1 (1863) 27;

var. ellipticum Mig. |.c.; var. ovalifolium Miq. l.c. — A. borneense Baker in Kew

Bull. (1896) 23. — A. congestum Ridley in Journ. Fed. Mal. St. Mus. 10 (1920)

137. — A. havilandii Ridley in Kew Bull. (1933) 494. A. rufo-sepalum Ridley

lic. (1946) 40. — A. rubiginosum Ridley l.c. 41.

Arthrophyllum 307

A small tree up to 14 m high, young parts with rufous tomentum. Leaves

clustered at the end of the branches, spirally arranged on the vegetative shoots,

imparipinnate or bipinnate (rarely tripinnate), multijugate, 150 cm long and 45 cm

wide (wider in bipinnate leaves); usually in opposite pairs on the inflorescence-

bearing branches and smaller with fewer pinnae or unifoliolate; leaflets ovate-

oblong or elliptic, up to 24 x 11 cm + coriaceous or membranaceous, margin

entire, slightly revolute, base truncate, rounded, or cuneate, often oblique, apex

shortly acuminate. Inflorescence a cluster of specialized leafy branches forming a

terminal crown which abscises after fruiting; the. main rays up to 150 cm long,

bearing pinnate (or more rarely unifoliolate) leaves mostly in opposite pairs; the

secondary rays up to c. 30 cm long, bearing mainly simple leaves in opposite pairs;

tertiary rays (peduncles) about 5 cm long, articulate about the middle; pedicels

+ 20, c. 5-12 mm long.

From the Malay Peninsula and Sumatra to Java, Borneo and Celebes. Type:

Java, Salak, Blume (Bogor).

From sea-level to 1600 m, in a wide variety of habitats, from dry sandy soil

to swampy humus. Occours in primary lowland and montane rain-forest and also

in secondary forest, heath-forest and waste land.

The very widespread A. diversifolium is variable in many characters, and many

comprise a number of geographic subspecies, but no basis for this is apparent at

present. Most individuals have the lower leaves simply imparipinnate, whereas

others have bipinnate, or rarely tripinnate, leaves. The flowers and inflorescences

of these forms appear to be identical, though raipd changes in the umbellules

after flowering produce a deceptively distinctive appearance in specimens at

different stages of development. Field experience over the whole range of the

species will be required to understand this interesting leaf-polymorphism. In

treating all forms as one species I am partly influenced by the fact that most

authors who have been familiar with the plants in Java (where both forms occur)

have regarded the complex as a single species (the fact that some authors have

recognized the variant from Salak as a distinct species does not affect the problem

of leaf polymorphism).

Apart from the strikingly different leaf forms just discussed, certain local

variants may eventually be shown to justify specific rank. A form growing on

Salak (near Bogor) has often been regarded as distinct (see, for example,

Hochreutiner in Candollea 2 (1925) 481, and Backer and Bakhuizen v.d .Brink,

FI. Java 2 (1965) 169). Indeed this form is the basis of the name A. diversifolium.

I retain this name in preference to the other two names published simultaneously

by Blume because it has been most consistently adopted since it was first used in

this comprehensive sense by Clarke. On the evidence available I do not consider

the Salak plants any more distinctive than many other local variants. It might be

considered that A. engganoense is also no more than another such variant, but its

facies is so marked that specific rank appears justified. It is possible that Ridley

was correct in distinguishing A. congestum, but the material is not good and

appears inadequate to confirm specific status. Five collections from Brunei and a

neighbouring district of Sarawak are all very alike and sufficiently distinct from

both A. diversifolium and A. crassum to suggest that they represent a further

species: these are Ashton S7840, Chew Wee-lek 981; Fuchs and Diederix 21182;

and van Neil 3895 and 4312. For the present these five gatherings are tentatively

retained as a form of A. diversifolium. Similarly, the two collections described by

Ridley as A. rubiginosum and A. rufo-sepalum are based on collections which are

not altogether typical of A. diversifolium, but which come closest to that species.

In. the absence of more supporting material, it is advisable not to retain them as

species. The first of these names (A. rubiginosum) has been widely used in

308 Gardens’ Bulletin, Singapore — XXX (1977)

identifications of Bornean ‘specimens, but the specimens concerned are either

typical A. diversifolium or belong to the distinctive A. crassum. Specimens from

Mt. Kinabalu described by Ridley as A. havilandii have bipinnate leaves, and

appear to conform well with A. diversifolium. This form was again collected on

Mt.- Kinabalu by Clemens and is also known from Sarawak (e.g. Morshidi

S24074). |

10. Arthophyllum macranthum Philipson, Bull. Brit. Mus. Nat. Hist. Bot. 1

(1951) 18.

Tree up to 25 m high, sparsely branched with leaves crowded at the ends of

the branches, all young parts with dense rufous tomentum. Leaves of the vegeta-

tive stems spirally arranged, imparipinnate, multijugate, up to 100 x 30 cm; of

the flower-bearing branches smaller with fewer pinnae, or simple; petioles; leaflets

ovate to oblong, up to 16 X 8 cm, coriaceous, margin entire, revolute, base

rounded, truncate, or cordate, very rarely cuneate, often oblique, apex obtuse or

bluntly apiculate. Inflorescence a whorl of specialized leafy branches forming a

terminal crown; the main rays up to 60 cm long, bearing pinnate leaves often in

opposite pairs; the secondary rays about 10-20 cm long, bearing simple leaves,

usually in opposite pairs, and terminating in umbellules or compound umbellules;

umbellules with about 10-15 flowers; pedicels stout 3-5 mm long to 10 mm or