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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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A SPECIAL ISSUE TO MARK THE EIGHTIETH BIRTHDAY OF R. E. HOLTTUM, 1975
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THE
GARDENS’ BULLETIN SINGAPORE
Vol. XXX 10th October, 1977
Dr. R. E. HOLTTUM: An Appreciation by 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
1
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 by 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 by
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
7
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 by 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 ~
Zz
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
by 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
13
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 by 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).
17
18
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.
4
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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 by
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).
21
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
t
‘
Iridescent Plants 27
9
2)
c
=
1S)
s
2
wd
=
5
ST]
a
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 by
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.
31
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.
36
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.
38
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
>i.
ae
a3
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.
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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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6
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Plate 2.
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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. by
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
45
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”’.
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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.
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Gardens’ Bulletin, Singapore — XXX (1977)
37
Humata pectinata
58
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
59
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 . §
A - a e bs at
=a i ie eek A eee ie
oh ES? Rerniny Ae - < vies 1 ee gh wet
at eae ae
; ee <2
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.
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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
by 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.
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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
a 4
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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44
Calamus caesius and Calamus trachycoleus Compared by 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
75
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
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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).
77
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